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University’ Microfilms International 300 N. Zeeb Road Ann Arbor, Ml 48106 8308960 King, Donna Kay COMMUNITY METABOLISM AND AUTOTROPHIC ■ HETEROTROPHIC RELATIONSHIPS OF WOODLAND STREAM RIFFLE SECTIONS Michigan Stale University University Microfilms International PHD. 1982 300 N. Zeeb Road, Ann Aitoor, MI 48106 PLEASE NOTE: In all c a se s this material h as been filmed in the best possible way from the available copy. Problems encountered with this docum ent have been identified here with a check mark V 1. Glossy photographs or p a g e s ______ 2. Colored illustrations, paper or print______ 3. Photographs with dark background______ 4. Illustrations are poor copy______ 5. P ag es with black marks, not original copy______ 6. Print shows through a s there is text on both sid e s of page______ 7. Indistinct, broken or small print on several p a g e s . 8. Print exceeds margin requirem ents______ 9. Tightly bound copy with print lost in spine______ 10. . Computer printout pages with indistinct print_____ 11. P ag e(s)____________ lacking when material received, and not available from school or author. 12. P ag e(s)____________iseem to be missing in numbering only a s text follows. 13. Two pages num bered I___________ . Text follows. 14. Curling and wrinkled pages 15. Other University Microfilms International COMMUNITY METABOLISM AND AUTOTROPHIC-HETEROTROPHIC RELATIONSHIPS OF WOODLAND STREAM RIFFLE SECTIONS By Donna K ay King A DISSERTATION Submitted to M i c hig a n State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Kellogg Biological Station and Department of Fisheries and Wildlife 1982 ABSTRACT COMMUNITY METABOLISM AND AUTOTROPHIC-HETEROTROPHIC RELATIONSHIPS OF WOODLAND STREAM RIFFLE SECTIONS. By Donna Kay King Using an in situ chamber oxygen method at five first through third order riffle sites In Augusta Creek, Michigan, estimates of community metabolism were made on a monthly or seasonal basis (1973 1975). Combined site, net community productivity (NCP) ranged from -0.25 to 4.03 g O 2 m - 2d"l, community respiration (CR) from 0.24 to 3.67 g O 2 m -2d” l, and gross community productivity (GCP) from 0.09 to 5.35 g O 2 m _2(j_ l. The integrative parameters net dally metabolism (NDM) and ratio of GCP to 24 hour CR (P/R) ranged from -0.72 to 2.68 g O 2 m ” 2d” l and -0.017 to 2.5, respectively and remained constant at each site. Most parameters were consistently low and stable at first order sites, while autotrophic sites varied significantly. Light- saturated photosynthesis occurred between 15,000 to 24,000 lux; no photoinhibition was evident. Detrital standing crops were stable at each site and ranged from 146 to 592 g m” 2. Average ratio of coarse to fine particulate organic matter ranged from 0.12 to 0.4 with an inverse relationship with P/R. Epilithon values ranged from 1480 to 5030 g m- 2* 12% of the total particulate organic matter. Detritus averaged Epilithon was detrital based at first order riffles; other sites were algal dominated. Average ratio of epilithon to detritus ranged from 4.5 to 14.8 wi t h greatest seasonal differences at the autotrophlc sites. Stream order was most highly correlated with all dependent variables (NCP, CR, GCP, NDM) followed by light. Temperature and epilithon development were significantly correlated with all dependent variables; non-attached detritus had little correlation. Particle-sized sediment (4 mm, 1 mm, 250 ym, 75 ym, 0.45 vm) contributions to NCP and CR were highest on 0.45 ym detritus and 4 mm (NCP) and 75 ym (CR) inorganic sediments on an AFDW basis. Areal estimates indicated highest NCP on larger inorganic and organic particles; highest CR occurred on 0.45 ym detritus and 4 m m inorganic particles. CR was highest on all detrital fractions on an AFDW basis; epilithon had higher contributions to NCP and CR on an areal basis. In situ trends were similar to areal particle-sized estimates. The support of woodland stream riffle sections was dependent on epilithon development while the autotrophic-heterotrophic balance was dependent upon the floral composition. This work is dedicated to Matthew H. Hohn. Who introduced me to the finer (diatomaceous) things ii life. ACKNOWLEDGEMENTS I would like to express sincere appreciation to K en Cumm i n s , for the atmosphere created In his laboratory, which allowed a technician to "sprout" into a student. His flexibility, enthusiasm, encouragement and friendship are valued. I am also grateful for the Montana "experience" and the opportunity to participate in the comparative methods conference at the Stroud Water Research Laboratory; both were highlights while working on this degree. This study would not have been completed without the continued interest and support of m y guidance committee: Drs. Kenneth W. Cummins, Chairman; Niles R. Kevern; Michael J. Klug; Robert G. Wetzel; and Ma t t h e w H. Hohn. To each I am grateful for their contributions to the thesis and my education. Dr. George H. Lauff, Director of the Kellogg Biological Station, deserves special thanks for facilitating the final stages of data analyses through provision of computer facilities and the opportunity to return summers to KBS to teach Freshwater Algae. Robert H. K i n g ’s support and encouragement throughout the years are greatly appreciated as well as his technical assistance in the field every sunrise and sunset included in the study period. Thank-you Bob for being you. Invaluable assistance in computer programming and data output were provided by Steven Weiss, John Gorentz and George Spengler. ill Virginia Holt, Roger Ovink, Ken Wiest and LaRaine Wank provided technical assistance at various stages of the study; Arthur Wiest constructed the chambers and Gall G u t h did the chamber and site cross sectional drawings. Charlotte Seeley and Alice Gillespie prepared the final manuscript and made the final stages of thesis preparation "enjoyable". I would also like to thank Karen Dacey, Patricia Gage, Gordon Godshalk, Carolyn H a m m a r s k j o l d , Virginia Holt, M a r y Hughes, Marilyn Jacobs, David Mahan, Rosanna Mattingly, Marjorie Robbins, Robbins, Steven Charlotte Seeley, M a r y Shaw, George Spengler, Jay Sonnad, M a r y Lynn Teague, Dolores Teller and Arthur Wiest for their individual contributions, encouragement and friendship during the research and/or writing phases of this study. The financial support for this study was provided by a Department of Energy Grant DE-AT06-79EV1004 for which I am thankful. iv TABLE OF CONTENTS Page List Of T a b l e s ........................................................ vii List Of Figures....................... xli Introduction.......................................................... 1 M e t h o d s ................................................................ 20 Results And D i s c u s s i o n .......................................... 33 Physical Parameters ............................................. T e mperature................................................ Light (Irradiance)........................................ 33 33 35 Community M e t a b o l i s m ............................................ Gross Community Productivity............................. Net Community Productivity............................... Community Respirat i o n..................................... Net Daily M e t a b o l i s m ...................................... Detrital Standing C r o p .................................... Detrital (AFDW) Community M e t a b o l i s m ................... E p i l i t h o n .......................................... 43 49 56 60 72 75 83 88 Factors Affecting Rates Of Community M e t a b o l i s m ....... 98 Stream O r d e r ............................................... 100 L i g h t ....................................................... 101 Temper a t u r e ........................................... 108 Detrital Ash-Free Dry W e i g h t ............................. 113 Epilithon Ash-Free D ry W e i g h t ........................... 113 Sites ........................... 117 Particle-Size Community M e t a b o l i s m ........................... Particle-size estimates of Community M e t a b o l i s m (AFDW basis)................................ Particle-size estimates of community metabolism (Areal b a s i s )............................... v 118 119 130 Page Synthesis............................................................. 149 Temperature.............................. L i g h t ............................................................ Community Metabolism.............. Regression Analyses............................................ .............. Particle Sized Community M etabolism S ites............................................................ 149 150 151 156 158 161 Literature C ited ............................. Appendix A Appendix B Appendix C Appendix D Appendix E Appendix F Appendix G Appendix H Estimates of community metabolism (g O 2 m ” 2) t Augusta Creek, M i c h i g a n ................................ Summary of community metabolism estimates, Augusta Creek, M i c h i g a n ................................ 163 174 257 Organic sediment composition (gAFDW) for selected sites on Augusta Creek, M i c h i g a n ........... 271 Summary of physical parameters for selected riffle sections of Augusta Creek, Michigan........... 291 Summary of regression analyses from selected sites of Augusta Creek, M i c h i g a n...................... 298 Light saturation curves from selected sites of Augusta Creek, M i c h i g a n............................ 305 Community respiration and NCP rates at various temperatures for selected sites of Augusta Creek, Michigan........................................ 316 Estimates of particle-sized sediment community metabolism from selected sites of Augusta Creek, M i c h i g a n .......................................... 328 vi LIST OF TABLES Page Table 1. Table 2. Table 3. Table 4. Table 5. Table 6 . Table 7. Table 8 . Table 9. Physical parameters of selected study sites, Augusta Creek, Kalamazoo County, Michigan (1973-1977).............................................. 9 Percent coverage and abundance of herbaceous plants from ten meter transects established perpendicular to each bank at five sites of Augusta Creek, Kalamazoo County, Michlagn, July 1975 ................................................. 14 Abundant and common macroinvertebrates from selected riffle sections of Augusta Creek, Kalamazoo County, Michigan (1974-1975). ......... 19 Comparison of community respiration rates obtained during daylight and dark periods from selected riffle sites of Augusta Creek, Kalamazoo County, M i c h i g a n ............................. 26 Procedure for estimating parameters of community metabolism experiments....................... 29 Temperature fluctuations at sites of Augusta Creek, Kalamazoo County, Michigan (1973-1975).............................................. 36 Light intensity fluctuations (irradiance) at selected sites of Augusta Creek, Kalamazoo County, Michigan (1973-1975)........................... 39 Literature estimates of in situ community metabolism in streams (g O 2 m - 2d“ l).......... 44 Average community metabolism parameters for experimental runs completed at five riffle sites of Augusta Creek, Kalamazoo County, M i c h i g a n 1973-1975...................................... 51 vii Page Table 10. Table 11. Table 12. Table 13. Table 14. Table 15. Table 16. Table 17. Table 18. Annual and seasonal estimates of P/R ratio at two sites of Augusta Creek, Kalamazoo County, Michigan (1973-1975).*....................... 68 CPOM and FP0M detrital standing crops and CP0M/FP0M ratios from selected sites in Augusta Creek, Kalamazoo County, Michigan (1974-1975).............................................. 81 Annual average estimates of detrital community metabolism (mg O 2 g “ ^AFDW) for selected sites of Augusta Creek, Kalamazoo County, Michigan (1974-1975)........................... 85 Percentage of detritus and epilithon and epilithon/detritus (E/D) ratios of sediments from selected sites of Augusta Creek, Kalamazoo County, M i c h i g a n ....... 95 Summary of regression data for various dependent variables and the independent variable stream order for all sites of Augusta Creek, Kalamazoo County, M i c h i g a n ........... 202 Summary of regression analyses involving various dependent variables and the independent variable light at five sites from Augusta Creek, Kalamazoo County, Michigan ..... 104 Summary of multiple regressions involving the effects of temperature and light on individual rates of NCP and CR from all sites of Augusta Creek, Kalamazoo County, M i c h i g a n ............................... 105 Summary of regressions involving the effects of temperature on NCP, GCP, CR, and NDM from various sites of Augusta Creek, Kalamazoo County, M i c h i g a n ......................................... 110 Regression summary for the relationship between GCP, NCP, CR, and NDM and the independent variable detrital standing crop (AFDW) for five sites of Augusta Creek, Kalamazoo County, M i c h i g a n ....... 114 viii Page Table 19. Regression summary for the relationship between NCP, GCP, CR, and NDM with the independent variable epilithon for five sites of Augusta Creek, Kalamazoo County, M i c h i g a n ................................. . Table 20. Comparison of community metabolism parameters from in situ and Gilson Respirometer experiments from five riffle communities of Augusta Creek, Kalamazoo County, M i c h i g a n ........................................ 120 Average particle-sized community metabolism estimates (weight basis) from Gilson Respirometery for five riffle sites of Augusta Creek, Kalamazoo County, M i c h i g a n ............ 124 Ranked particle-size contributions to various parameters of community metabolism (weight basis) from five riffle communities of Augusta Creek, Kalamazoo County, M i c h i g a n .............................................. 125 Average estimates of community metabolism of various particle-sizes expressed on an areal basis for five riffle sites of Augusta Creek, Kalamazoo County, Mic h i g a n........ 131 Ranked particle-size contributions on an areal basis to estimates of community metabolism from five riffle communities of Augusta Creek, Kalamazoo County, M i c h i g a n ............ 132 Table 21. Table 22. Table 23. Table 24. Table A-l Estimates of Community Metabolism (g C ^ / m ^ ) , Augusta Creek, Mi c h i g a n ................................. 174 Table A-2 Estimates of Community Metabolism/gAFDW Detritus, Augusta Creek, M i c h i g a n ..................... 218 Table B-l Summary of community metabolism estimates for SMITH SITE, Augusta Creek, M i c h i g a n ............... 257 Table B-2 Summary of community metabolism estimates for B AVENUE, Augusta Creek, M i c h i g a n ................. 258 Summary of community metabolism estimates for UPPER 43, Augusta Creek, M i c h i g a n ................. 259 Table B-3 Ix Page Table B-4 Table B-5 Table Summary of community metabolism estimates for NAGEL S I T E , Augusta Creek, M i c h i g a n 261 Summary of community metabolism estimates for KELLOGG FOREST, Augusta Creek, M i c h i g a n ............... 263 B-6 Summary of community metabolism estimates on a detrital AFDW basis for Smith Site, Augusta Creek, Michigan .............. Table B-7 264 Summary of community metabolism estimates on a detrital AFDW basis for B Avenue Site, Augusta Creek, M i c h i g a n .......................................... 265 B-8 Summary of community metabolism estimates on a detrital AFDW basis for Upper 43rd Site, Augusta Creek, M i c h i g a n ................................. 267 B-9 Summary of community metabolism estimates on a detrital AFDW basis for Nagel Site, Augusta C r e e k , M i c h i g a n ................... ... ................... 268 Table B-10 Summary of community metabolism estimates on a detrital A F D W basis for Kellogg Forest Site, Augusta Creek, M i c h i g a n ............. ...... ............. 270 Table Table Table C-l Table D-l Table D-2 Table D-3 Table D-4 Table D-5 Organic sediment composition (gAFDW) for selected sites on Augusta Crreek, M i c h i g a n ........... 271 Summary of physical parameters for selected riffle sections of SMITH SITE, Augusta Creek, M i c h i g a n ...................... 291 Summary of physical parameters for selected riffle sections of B AVENUE, Augusta Creek, M i c h i g a n .................. 292 Summary of physical parameters for selected riffle sections of UPPER 43, Augusta Creek, M i c h i g a n .............. 294 Summary of physical parameters for selected riffle sections of NAGEL SITE, Augusta Creek, M i c h i g a n .................................................. 295 Summary of physical parameters for selected riffle sections of KELLOGG FOREST, Augusta Creek, M i c h i g a n .......................................... 297 x Page Table E-l Table E-2 Table E-3 Table H-l Table H-2 Regression parameters for average NCP, CR, G C P and NDM versus independent variables from data collected in Augusta Creek, M i c higan riffle sections................................ 298 Regression parameters for individual rates of NCP and CR versus various independent variables from data collected in Augusta Creek, Michigan riffle sections.............. 299 Regression analyses for various sites of Augusta Creek, Michigan.............................. 300 Estimates of mean hourly rates of community m etabolism based on Gilson Respirometery from five riffle sites of Augusta Creek, M i c h i g a n .................................................. 328 Areal estimates of community metabolism based on Gilson Respirometery experiments from five riffle sites of Augusta Creek, M i c h i g a n ............... 343 xi LIST OF FIGURES Page Figure 1. Figure 2. Figure 3. Figure 4. Figure 5. Figure 6 . Figure 7. Figure 8 . Figure 9. Augusta Creek watershed, Barry and Kalamazoo counties, Michigan (modified from M a n n y and Wetzel 1973).................................................... 6 Community metabolism study sites selected in the Augusta Creek watershed (Kalamazoo and Barry counties, M i c h i g a n ) ............................................... 8 Woody vegetation profiles of Stoith and B Avenue sites, Kalamazoo County, Michigan ( 1975).................................................. 10 Woody vegetation profiles of Upper 43rd, Nagel, and Kellogg Forest sites, Augusta Creek, Kalamazoo County,Michigan (1975)............. 12 Percent composition of particle sizes from riffle section inorganic sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975)............................................. 18 Plexiglas chamber designed for estimating community metabolism in stream riffle s e c t i o n s ............................ 21 Experimental design of in situ community metabo l i s m field experiments....................... 24 Laboratory procedure for determination of ash-fee dry weight (AFDW) of chamber sediments andmacroinvertebrates...................... 28 Experimental design for Gilson Respirometer estimates of community metabolism of particle sized s u b strate.................. xii 32 Page Figure 10. Figure 11. Figure 12. Figure 13. Figure 14. Figure 15. Figure 16. Figure 17. Figure 18. Figure 19. Average dally temperature values for selected sites in the Augusta Creek watershed, Kalamazoo County, Michigan (1973-1975)............................................ 34 Average daily light values for the Smith, B Avenue, and Upper 43 Rd sites of Augusta Creek, Kalamazoo County, Michigan (1973-1975)............................................ 37 Average daily light values for Nagel and Kellogg Forest Sites of Augusta Creek, Kalamazoo County, Michigan (1973-1975........... 38 Daily light patterns at Kellogg Forest, Augusta Creek, Kalamazoo County, Michigan on 25 July 1975........................................ 42 Average GCP (+ SE) estimates for selected riffle communities in the Augusta Creek watershed, Kalamazoo County, Michigan (1973-1975)............................................ 50 Average rates (+ SE) of NCP at selected riffle sites in the Augusta Creek watershed, Kalamazoo County, Michigan (1973-1975)............................................ 58 Rates of average CR (+ SE) for selected riffle communities in the Augusta Creek watershed, Kalamazoo County, Michigan (1973-1975)............................................ 62 Average P/R ratios (+ SE) for selected sites in the Augusta Creek watershed, Kalamazoo County, Michigan (1973-1975).............. 66 Average values (+ SE) of NDM for selected riffle sites in the Augusta Creek watershed, Kalamazoo County, Michigan (1973-1975)............................................ 73 Average estimates of detrital standing crop (+ SE) from selected riffle communities in Augusta Creek, Kalamazoo County, Michigan (1973-1975)............................................ 77 xiii Page Figure 20. Figure 21. Figure 22. Figure 23. Figure 24. Figure 25. Figure 26. Figure 27. Figure 28. Seasonal distribution of particle-sized detritus from selected riffle sections of Augusta Creek, Kalamazoo County, Michigan (1973-1975)............................................ 78 Estimates of NCP and CR on a weight basis from selected Augusta Creek riffle sections, Kalamazoo County, Michigan (1973-1975)....... 86 Estimates of GCP and NDM on a weight basis from selected Augusta Creek riffle sections, Kalamazoo County, Michigan (1973-1975)............................................ 87 Estimates of eiplithon development, obtained on a seasonal basis from selected riffle sites of Augusta Creek, Kalamazoo County, Michigan (1974-1975)......................... 91 Seasonal distribution of particle-sized epilithon from selected riffle sections of Augusta Creek, Kalamazoo County, Michigan (1974-1975)............................................ 94 Epllithon/Detritus ratios over an annual period for selected riffle sites of Augusta Creek, Kalamazoo County, Michigan (1974-1975)............................................ 97 Estimates of NCP from particle sized detritus and epilithon communities from Smith, B Avenue and Upper 43rd riffle sites of Augusta Creek, Kalamazoo County, Mi c h i g a n (1974-1975).................................. 121 Estimates of NCP from particle sized detritus and epilithon communities from Nagel and Kellogg Forest riffle sites of Augusta Creek, Kalamazoo County, Michigan (1974-1975)............................................ 122 Estimates of CR from particle-sized detritus and epilithon communities from selected riffle sites of Augusta Creek, Kalamazoo County, Michigan (1974-1975).............. 128 xiv Page Figure 29. Figure 30. Figure 31. Figure 32. Figure 33. Figure 34. Figure 35. Figure 36. Figure 37. Figure 38. Estimates of particle sized community metabolism of the detrital component of Smith riffle sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975).............. 135 Estimates of particle sized community m etabolism of the epilithon of Smith riffle sediments, Augusta Creek, Kalamazoo County, M ichigan (1974-1975).................................. 136 Estimates of particle sized community metabolism of the detrital component of B Avenue sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975)......................... 138 Estimates of particle sized community metabolism of the epilithon of B Avenue riffle sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975)..................... 139 Estimates of particle sized community metabolism of the detrital component of Upper 43rd riffle sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975).............. 140 Estimates of particle sized community metabolism of the epilithon of Upper 43rd riffle sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975)......................... 142 Estimates of particle sized community metabolism of the detrital component of Nagel riffle sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975).............. 144 Estimates of particle sized community metabolism of the epilithon of Nagel riffle sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975).................................. 145 Estimates of particle sized community metabolism of the detrital component of Kellogg Forest riffle sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975)............................................ 147 Estimates of particle sized community metabolism of the epilithon of Kellogg Forest riffle sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975).............. 148 xv Page Figure F-l Net community productivity at various light intensities for combined sites of Augusta Creek, Michigan (1973-1975).......................... 305 Net community productivity at various light intensities for Smith Site, Augusta Creek, Mi c h i g a n (1974-1975).................................. 306 Net community production at various light intensities for B Avenue Site, Augusta Creek, Michigan (1973-1975)......................... 307 Net community production at various light intensities for Upper 43rd Site, Augusta Creek, Michigan (1974-1975).......................... 308 Net community production at various light intensities for Nagel Site, Augusta Creek, M i c h i g a n (1973-1975).................................. 309 Net community production at various light intensities for Kellogg Forest Site, Augusta Creek, Michigan (1974-1975)................. 310 Net community productivity within the 0-5C temperature range for combined sites of Augusta Creek, Michigan (1973-1975)................. 311 Net community productivity within the 5-10C temperature range for combined sites of Augusta Creek, Michigan (1973-1975)............... 312 Net community productivity within the 10-15C temperature range for combined sites of Augusta Creek, Michigan (1973-1975)............... 313 Figure F-10 Net community productivity within the 15-20C temperature range for combined sites of Augusta Creek, Michigan (1973-1975)............... 314 Figure F-l1 Net community productivity within the 20-25C temperature range for combined sites of Augusta Creek, Michigan (1973-1975)............... 315 Figure F-2 Figure F-3 Figure F-4 Figure F-5 Figure F-6 Figure F-7 Figure F-8 Figure F-9 Figure G-l Rates of community respiration at various temperatures for combined sites of Augusta Creek, Michigan (1973-1975).......................... xvi 326 Page Figure G-2 Figure G-3 Figure G-4 Figure G-5 Figure G -6 Figure G-7 Figure G -8 Figure G-9 Rates of community respiration at Smith site, Augusta Creek, Michigan (1974-1975)........... 317 Rates of community respiration at B Avenue site, Augusta Creek, Michigan (1973-1975)........... 318 Rates of community respiration at Upper 43rd site, Augusta Creek, Michigan (1974-1975)............................................. 319 Rates of community respiration at Nagel site, Augusta Creek, Michigan (1973-1975).......... 320 Rates of community respiration at Kellogg Forest site, Augusta Creek, Michigan (1974-1975)............................................. 321 Net community productivity at various temperatures for combined sites of Augusta Creek, Michigan (1973-1975).......................... 322 Net community productivity at various temperatures for Smith site of Augusta Creek, Michigan (1974-1975).......................... 323 Net community productivity at various temperatures for B Avenue site of Augusta Creek, Michigan (1973-1975).......................... 324 Figure G-10 Net community productivity at various temperatures for Upper 43rd site of Augusta Creek, Michigan (1974-1975)...... 325 Figure G-ll Net community productivity at various temperatures for Nagel site of Augusta Creek, Michigan (1973-1975).......................... 326 Figure G - l 2 Net community productivity at various temperatures for Kellogg Forest site of Augusta Creek, M ichigan (1974-1975)................. 327 xvii INTRODUCTION As the sun proceeds slowly northward the temperate streams, a golden glow stones, and logs, announcing another 20 March 1974 Augusta warming and lighting results on rocks, diatomaceous spring... Creek Interest in primary production of autotrophic organisms ranges from world wide support of the human population to unraveling the complex functions of ecosystems (MacFaden 1948, Wetzel 1975). Although there was early notice of primary producers and their importance in aquatic systems, it was not until 1944 that in-depth inquiries were begun. Wetzel These early studies on lakes are summarized by (1965, 1975a) and Lieth (1975). lotic systems are minimal (Odum 1956). Early productivity studies on Lotic systems are extremely heterogeneous and dynamic in all aspects. physical, chemical, Besides the typical spacial, and biological variables common in lentic systems, one studying lotic systems encounters variability in canopy cover, substrate distribution, and current; which in turn affect niche availability and distribution of organisms at all levels of food webs. These variables, along with watershed effects through runoff, flooding, and perturbations by man, are reviewed by Hynes (1970) and Whitton (1973), and their applicability to measurement of primary productivity discussed by Hynes (1970) and Wetzel (1975b). Methodology for estimating primary productivity in aquatic systems has been extensively reviewed (Ryther 1956b, Goldman, 1 1969, Vollenweider 1969, Hall 1973, Hall and Moll 1975, Bott et al. 1982). At the time of this investigation (1973 - 1975) no published in situ investigations had been made using chambers under natural field conditions in streams. Since this study, however, in situ measurements of community metabolism have been reported (Gothberg and Karlstrom 1975, Marker 1976b, Naiman 1976, Litke 1978, Pennak and Lavelle 1979, al. 1981), Sumner and Fisher 1979, Bush and Fisher 1981, Hornick et and a comparative study of three chamber techniques, including the chamber designed for this study (King chamber), and two open water techniques for estimating community metabolism has been completed (Bott et al. 1975). Although in situ chamber methods are beginning to take the forefront in the determination of community metabolism, Pennak and Lavelle (1979) and Rodgers et al. (1980) reinforce the lack of such data and the need to continue with this type of approach to examine ecological questions concerning community metabolism. It was the purpose of this investigation to estimate in situ levels of community metabolism for selected riffle sections of the Augusta Creek watershed over a two-year period and to examine the autotrophic-heterotrophic relationships within these communities through measurement of autochthonous production and community respiration with subsequent calculation of the integrative parameters the ratio of gross community productivity to 24 hour community respiration (Pg/R24 = P/R) and net daily metabolism (NDM). First through third order (Strahler 1957) sites were examined to elucidate changes in community metabolism throughout the headwater portion of the watershed and the effects of light and temperature on the rates 3 of in situ productivity. Relationships of detrital and epilithon standing crops to community metabolism were also examined. The major hypotheses were that the shaded first order sections obtain minimal support through autotrophic p a t h w a y s , which would indicate a heterotrophic dependence, while the open second and third order reaches would be supported by an autotrophic energy base over the annual period and that clearing of riparian vegetation from third order sections would increase the autotrophic potential of the riffle community. Estimates of community metabolism have been shown to reflect environmental changes due to pollution (Odum 1956) or short term stress such as application of lampricides (Maki 1974). The P/R ratio and NDM are also of value in characterizing autotrophicheterotrophic relationships in aquatic systems. Odum (1956), using P/R ratios from open water estimates, indicated the significance of this ratio in the classification of stream communities based on their major carbon source. He classified communities which received large amounts of allochthonous carbon in which respiration was greater than gross production (P/R < 1) as heterotrophic, while those in which there was much autochthonous production of carbon (P/R > 1) were termed autotrophic. This relationship has been established as a physiological classifier of streams by Caspers and Karke in 1966 and 1967 as reviewed by Pavletic et al. (1976). Hynes (1970) commented on the little attention that this valuable parameter had received in lotic systems. Fisher and Likens (1973) expanded on this relationship by incorporating estimates of export and import. Minshall (1978) reported P/R < 1 in aquatic plant-dominated lotic reaches, which indicated the possibility of heterotrophic conditions in communities producing large amounts of autochthonous carbon. This carbon, however, is available to the community mainly when converted to detritus, often downstream, which indicates the heterotrophic nature of the riffle due to low attached algal development from shading by aquatic macrophytes. Estimates of NDM (gross community production minus 24 hour community respiration) yield a valuable quantitative measure of support or dependence of a community on allochthonous carbon sources (Bott et al. 1978, Vannote et al. 1980). The heterotrophic nature of headwater streams and the importance of allochthonous inputs of organic carbon such as leaves, twigs, flower parts, bark, fine particulate organic matter (FPOM), and dissolved organic matter (DOM) is well documented as is the decomposition of these materials by bacteria, fungi, and macroinvertebrates within the deciduous biome (Minshall 1967; Kaushik and Hynes 1968, 1971; Trlska 1970; Hynes 1970; Coffman et al. 1971; Cummins et al. 1973; 1977, 1978, 1980). Suberkropp and Klug 1974; Cummins 1974, 1975a, Suberkropp et al. 1976, Bell et al. 1978, Vannote et al. These studies indicate that heterotrophic stream sections receive as little as one percent of their carbon from autochthonous production. Observations of forested areas of Augusta Creek, a woodland Michigan stream, indicated epilithic, epipsammic and epipelic algae were present, as well as high populations of algal-feeding aquatic insect larvae (scrapers; Merritt and Cummins 1978) such as Glossosoma and Neophylax (Trichopterascaddisflies). Wetzel (1975b) and Minshall (1978) emphasized that in order to use an autotrophic-heterotrophic 5 classification system one must first know the magnitude of community production and not assume autotrophic insignificance. In situ measurements of community metabolism not only estimate productivity, but total community production, which includes both autotroph and heterotroph (bacterial, fungal, and invertebrate) metabolism (Wetzel 1975b, Bott et al. 1978, Pennak and Lavelle 1979). Such estimates are directly dependent on light, temperature, carbon dioxide, and oxygen levels and indirectly on inorganic and organic nutrient availability, which all affect photosynthesis and respiration (Lieth 1975; Wetzel 1975a, 1975b). Intensive investigation of the effects of these parameters on iri situ community metabolism has not been completed. The extent to which autotrophic communities develop and contribute to the structure and function of woodland streams needs to be addressed. DESCRIPTION OF STUDY SITE The pussywillows have popped their bud scales and the Cornus with a blushing glow awakes from it's dull red winter slumber! They now await the stirring of leaf buds... Spring is in the air at Augusta Creek. 5 March 1974 Augusta Creek, a woodland stream located in Barry and Kalamazoo counties, Michigan, was the site of this Investigation (Figure 1). This third order (Strahler 1957) stream system joins the Kalamazoo River in Augusta, which in turn flows westerly into Lake Michigan at Saugatuck. The watershed has been described by Mahan and Cummins (1974) as having a total length of 73.3 k m (39.3 mi) and a drainage area of 72.3 k m “ 2 (27.9 mi"2). The watershed includes a combination 6 A U G U S T A CREEK WA T E R S H E D f • Kir N ^ U ip . . -.ailil'iifr * i - N - t o XI N.T.1S..RJWR.9WI i 1 * \ BARRY * & & KALAMAZOO C O U N T I E S . MICHIGAN SITES: 2 4 J S = Smith I K ILO M ETER U = Upper 43rd if n i . 1 B ■ B Avenue — N = Nagel IplinarT ilir i s. T W 7 y # i ARE A: 6 7 . 8 3 KM s K - Kellogg Forest vk - -w SLO PE: 2 .0 3 % KM AVERAGE ELEVA TIO N: 2 6 6 U. K | l l M M ARSHLANO aklaata 1 ^ c W OODLAND 1 W ATERSHED BOUNDARY KALAMAZOO RIVfR Figure 1. Augusta Creek watershed, Barry and Kalamazoo counties, M ichigan (modified from M a n n y and Wetzel 1973). of forested, meadow, and agricultural reaches with approximately 50% of the drainage in agricultural use (King 1979, M ahan 1980). The slope is slight, 2.03 m ^ k m ” * (Manny and Wetzel 1973), and discharge measured at the United States Geological Survey gauging station in Augusta (Figure 2) ranged from 8.72 m “ 3s- l (308 cfs) in 1975 to 0.27 m “ 3s"l (9.5 cfs) in 1970 and averaged 1.19 n T ^ s - ! (42 cfs) Cummins 1974). King (Mahan and (1978) gives weekly velocities and discharge values for the B Avenue, Nagel and Kellogg Forest Sites from 1973 to 1974 and discharge records from 1976 to 1977 at these sites plus Smith and Upper 43rd Sites are cited in M ahan (1980). The range and average discharge values for the sites selected for this investigation are summarized in Table 1. Augusta Creek is characterized as a hardwater stream with pH values of 7.5 to 8.7, alkalinity of 160 to 230 m g " l l “ l, and total hardness of approximately 280 m g - ^l- ^ CaC03 (Manny and Wetzel 1973, M a h a n and Cummins 1974, King 1978). Inorganic nitrogen levels ranged from 2 to 5 m g ” ^l~^ and orthophosphate from 10 to 40 yg- ^l- ^ during a diel study by Manny and Wetzel (1973). The present investigation concentrated on riffle sections of two sites in Augusta Creek (B Avenue and Nagel) during 1973-1974, while riffles of five sites (Smith, B Avenue, Upper 43rd, Nagel, Forest) were studied in 1974-1975 (Figures 1 and 2). and Kellogg The Smith Site typifies a natural first order site within the Augusta Creek watershed and joins the B Avenue tributary before confluence with the Upper 43rd reach (Figures 1, 2, 3). The B Avenue first order reach originates in a tamarac (Larix larcina) area, flows southwest through an agriculturally perturbated section where cattle have free access to 8 AUGUSTA CREEK WATERSHED UPPEB43RP S MITH NAGEL KELLOGG FOREST U.S. Gauging Station Figure 2. Community metabolism study sites selected in the Augusta Creek watershed (Kalamazoo and Barry counties, Michigan). 9 Table 1. Physical parameters of selected study sites, Augusta Creek, Kalamazoo County, Michigan (1973-1977). TEMPERATURE °C SITE ORDER RANGE DISCHARGE m 3 s -1 x MAXIMUM MINIMUM X SMITH* 1 0-25.6 9.3 0.257 0.004 0.013 B AVENUE** 1 0-24.6 8.7 0.400 0.030 0.070 UPPER 43rd* 2 0-29.6 10.7 2.100 0.290 0.643 NAGEL** 3 0-28.4 10.7 3.30 0.400 1.130 KELLOGG FOREST** 3 0-27.2 10.5 3.470 0.430 1.200 * Data from Mahan (1980) ** Data from King (1978) and M ahan (1980) [SMITH KIT:] SMITH TREES 20- Upstream » = J = P= U= w 30 20 20 Channel SHRUBS AND VINES ( T ) CmWS.IPP. (Dogwood) Pirthenoti*se*ji*Hrepe(*ffil (Virginia Creeper) Bhes lypM n (Slaghom Sumac) BoMLSPl. (Wild Rose) B pIm sjpp, (Blackberry and Raspberry) Saijispp, (Willow) Samhnaw caaadansh (Common Elder) TBJtcodwdren r a d k a a (Poison Ivy) V JtP U n . (WHd Grape) \BAvcmm] B AVENUE I TREES A = Acer rnhrem (Red Maple) B = getulajatea (Yellow Birch) C = Carpimn carel Malta (Blue Beech) F = FrarinPSTPt. (Black and White Ash) U = Ulmps s t (Elm) Upstream Distance (m) Figure 3. 30 ftM L flh tm (Red Maple) JpplpHi niff* (Black Walnut) P ren e stp , (Cherry) Ulnreirukra (Siiopeiy Elm) Channel SHRUBS MIO VINES (T) AIim r j p . (Alder) C om otspp, (Dogwood) Lindera benzoin (Spice Bush) e«rth< n«P )m tJH riilH »fin (Virginia Creeper) Rlreityphjna (Slaghom Sumac) R .jretnli (Poison Sumac) BnPJPP. (Wild Rose) R nbnajpc, (Blackberry and Raspberry) tsiicodendren radhrens (Poison Ivy) VHMtpp- (Wild Grape) W oody vegetation profiles of Smith and B Avenue sites, Kalamazoo County, Mi c h i g a n (1975). 11 the stream, then flows Into Hamilton Lake (Figures 1, 2, 3). The second order Upper 43rd meadow reach Is within the tributary started by Gilkey and Fair Lakes and is downstream from Lawrence Lake (Figure 1); it joins the B Avenue - Smith tributary to form a third order system. The third order Nagel Site, located near the center of the drainage is an open reach influenced by swimming, fishing and vegetation removal (Figures 1, 2, 4). Along the east bank riparian vegetation was present, but in places the west bank was a mowed lawn (Figure 4). The Kellogg Forest Site is owned and maintained in a "natural" forested condition by Michigan State University and is open to the public for fishing. It is located downstream from the cleared Nagel site and provided a comparison of forested and cleared third order reaches. Percent cover, as well as the major woody canopy components, determined from transects (30 m x 1 n) established perpendicular to the stream banks are illustrated in Figures 3 and 4. The narrow Smith and B Avenue reaches were heavily shaded and contrast with the open m eadow reaches of Upper 43rd and Nagel Sites. The Kellogg Forest Site, was a shaded third order reach along Augusta Creek. An extensive analysis of the woody vegetation for these five sites was made by Mahan (1980). Of the 41 taxa observed in the herbaceous understory of the transects only five had abundant (> 50% occurrence) or common (> 10% occurrence) status (July 1974). Sedges (Carex spp.) were common or abundant at all sites except B Avenue, Grasses (Graminae) were common at Nagel Site, skunk cabbage (Symplocarpus foetidus (L.) Nutt.) was common at B Avenue, goldenrods spp.) at Upper 43rd site, and forget-me-knots (Solidago (Myosotis sp.) were w ¥ iUPPtR 4 3 r tK fr ] I U PPER 4 3 rd ! TREES P = Prm«s (Cherry) T = ThejsoccMentaNi (White Cedar) Upstream £ 30 20 10 Distance (m) m 5.4 m Channel 10 20 30 SHRUSSMtDtmtES (T) (r e la te s SP, (Hawthorn) Rosa spp. (Wild Rose) Sallrspp, (Willow) Toilceeendron radicals (Poison ivy) HERBACEOUS VEGETATION OHIT (A) I NAGEL | IWAGflAfrl I w TREES Upstream Po ■■agTTfa *iS“ 20 Distance |n ) 7.5m Channel Ik e l l o g g f o r e s t ! 20 * £01 X Figure 4. JgL. -f30 HERBACEOUS VEGETATION ONLY (*.> TREES F = fraalnes atari (Black Ash) Po = P o pehnN m ettM es (Trembling Aspen) S = Saljasp, (Willow) SHRUBS AMD VINES (T) ¥ Upstream Dtstanca (hi) 20 iKEUO G GfOH ESTKfTl s •» w ■+■ *> A = A tr u H r a n (Red Maple) Po = PppetestremeleMct (Trembling Aspen) SHRUBS AND VINES IT) Cb m u i b L (Dogwood) PhysocarpeE e p tR tm w (Nlnebark) Salls.spB, (Willow) 7.0 m Channel C tn v u a R . (Dogwood) Parthenoc lStes j l lomHWll (Virginia Creeper) e hym a iHttJBBUMKH (HinebarK) Rlh e s se . (Gooseberry) BPH IPB. (WHd Rose) S ll iU S t, (Willaw) S tP t l l W H . (Elderberry) 5phea_se, Woody vegetation profiles of Upper 43rd, Nagel, and Kellogg Forest sites, Augusta Creek, Kalamazoo County, M ichigan (1975). 13 common at the Kellogg Forest (Table 2). At Upper 43rd and Nagel Sites emergent vegetation was present as w e l l , including Peltandra virgin!ca (L.) Kunth. and Saggittaria latifolia Willd. During the summer at the Nagel Site, m u c h of the riffle was covered by submergent j>. l a t i f o l i a , Sparganium, Potamogeton pectinatus L . , and P. nodosis Poir. The physical composition of the sediments for the five sites, after Cummins (1962), was determined by particle-sizing into the following categories: sand pebbles (> 16 m m ) , gravel (> 4 m m ) , very coarse (> 1 mm), m e d i u m sand (> 250 Ur n) , and very fine sands (> 75 pm). All sites were dominated by the larger particle-sizes with 65 to 81 percent composition of 16 and 4 m m substrates (Figure 5). The Smith and B Avenue Sites were very similar and on the lower end of the average (65 and 66% respectively), while the Upper 43rd (76%), Nagel (81%) and Kellogg Forest (77%) Sites were similar and had a higher proportion of particles greater than 16 m m in diameter. Travertine deposits were noticeable on the rocks at Upper 43rd, and Nagel, which reflected increases in photosynthetic activity and the alkaline nature of the water; travertine was absent at Smith and B Avenue, and intermediate in development at Kellogg Forest. The macroinvertebrates, which accounted for greater than 10% of the standing crop over an annual period, included 19 taxa: five mayflies (Ephemeroptera), five caddisflies (Trichoptera), one riffle beetle (Coleoptera), six dipterans with a predominance of midges ( Diptera), the amphipod Gammarus, 3). and a water mite (Hydracina) (Table The B Avenue Site had the highest number of common taxa (13), although the overall diversity was highest in the open reaches (Table 3). Table 2. Percent coverage and abundance of herbaceous plants from ten meter transects established perpendicular to each bank at five sites of Augusta Creek, Kalamazoo County, Michiagn, July 1975. SMITH B AVENUE U P P E R 43rd TAXON COMMON NAME L R* TA** L R TA L R Bryophyta moss 8 10 R 3 2 R - - Arthrophyta Equlsetum sp. horsetails 1 - R 8 1 R - 7 Pterophyta Pryopteris sp. Onoclea sensibilis L. shieldfern sensitive fern 6 11 R - 14 R - common cattail grasses <1 1 R 1 2 R <1 10 R 2 50 2 R 72 28 A TA R KELLOGG FOREST NAGEL L R TA L R TA 1 R 2 4 R 4 1 R - - - 4 - R - - - 17 R 4 R C <1 4 R 3 54 C 44 C 1 1 - R R - - - - - - 3 1 R Magnoliophyta Typhaceae Typha latifolia L. Gramineae Cyperaceae Carex spp. Sclrpus atrovlrens Willd. validus Vahl. Araceae Symplocarpus foetidus (L.) Nutt. sedges 52 33 C great bulrush - - - skunk cabbage 1 - R 19 12 26 C - - - Table 2. (cont'd.) SMITH U PPER 43rd TA T AXON COMMON NAME L Iridaceae Iris sp. wild Iris 1 Urticaceae Urtica sp. nettle Ranunculaceae Caltha palustrls L. Ranunculus sp. Thalictrum sp. m a r s h marigold buttercup m e a d o w rue Brassicaceae Berteroa incana (L.) D C hoary allysum - Rosaceae Fragarla virginiana Duchesne. Potentilla sp. wild strawberry <1 cinquefoil - 1 1 Fabaceae Trifolium pratense L. red clover Geraniaceae Geranium maculatum L. wild geranium Balsaminaceae Impatiens balsamina L. jewelweed 1 8 <1 2 R* B AVENUE TA** TA 1 1 2 5 R R R R 2 - R 1 3 1 1 10 - R R R 1 2 R 2 TA - 2 R - <1 R - R R 9 - R 1 R <1 TA R R R 1 3 - KELLOGG FOREST NAGEL R - <1 <1 2 R <1 <1 R Table 2. (c o n t ’d .) SMITH TAXON COMMON NAME L Violaceae Viola sp. violet - Lythraceae L y thrum sallcarla L. purple loostrlfe - wild carrot 3 - Asclepidaceae Asclepias sp. milkweed - Boraglnaceae Myosotls sp. forget-me-not - Verbenaceae Verbena sp. vervain - water horehound - Aplaceae Angelica atropurpurea L. Clcuta bulblfera L. Daucus carota L. R* 1 TA** L R - 2 - B AVENUE R - TA 1 R - _ _ _ R - _ - _ _ - - U P P E R 43rd L R - - TA - - - 2 - 1 2 - - 1 1 - - - - - - 1 L R TA L R 1 — R 1 - - - 3 <1 R R - 10 5 - R R - - - R 1 <1 R _ - - - 53 1 1 R _ <1 <1 R 1 - - KELLOGG FOREST NAGEL R TA — R R - Primulaceae Lyslmachla clllata L. Laiaaraceae Lycopus amerlcanus Muhl. - - - - - _ - - - _ _ - _ - _ _ - - - _ _ - _ C _ — R Table 2. (cont'd.) SMITH T AXON COMMON NAME Prunella vulgaris L. self-heal Rubiaceae Galium sp. bedstraw Lobeliaceae Lobelia sp. Asteraceae Cirsium sp. Erigeron sp. Eupatorium m a c u latum L. E. perfoliatum L. Rudbeckia hirta L. Solidago sp. Taraxacum officinale Weber L U P P E R 43rd KELLOGG FOREST NAGEL R* TA** L R TA L R TA L R TA — R 1 — R — - — 1 — 10 1 R 1 3 R <1 <1 R - — — 2 R — — — 1 1 R thistle fleabane daisy 1 2 1 - R R joe-pye weed boneset black-eyed susan goldenrod 4 common dandelion <1 1 1 R R - 1 R 3 10 8 28 — R - - — — 5 3 R 5 1 R - <1 unknowns Total Taxa * L = left bank R = right bank B AVENUE 23 ** T A A C R = = = =* 16 L R TA R <1 — R 1 R 1 1 R 5 — R - - — R C 4 1 - 3 - - 1 - 1 3 R R R R - - R - 1 4 R 1 R 10 — R - — - 1 R - 4 R 1 6 R 21 total abundance (x percent coverage for the site) abundant = greater than 50% common = greater than 10%; less than 50% less than 10% 28 16 Figure 5 Percent composition of particle-sizes from riffle section inorganic sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975). 19 Table 3. Abundant and common macroinvertebrates from selected riffle sections of Augusta Creek, Kalamazoo County, Michigan (1974-1975). Taxon Smith B Avenue Upper 43rd Kellogg Forest Nagel EPHEMEROPTERA Baetidae Ephemerella Stenonema Caenls Hexagenia - C* C C C C C C C - - C C - C C C C C C C C TRICHOPTERA Hydropsychidae Psychomyeiodae Neophylax Glossosoma Protoptila - C C C - - C C C A C C A C - A A - - - - COLEOPTERA Elmldae DIPTERA Chironomldae Simulidae Atherix Antocha Erlocera Dlcranota « Jidk A A - C C A - - - AMPHIPODA Gammarus — ARCHNIDA Hydracina TOTAL (C and A) TOTAL SPECIES ”T ~ (15) * C = common ( > 10% < 50%) ** A “ abundant (> 50%) 13 (19) C ~ T (30) I T (30) C ~f~ (29) 20 METHODS Indeed the spring diatom bloom is such a pleasant snake free period on Augusta Creek... 5 M a r c h 1974 Estimates of community metabolism were measured over a two year period (1973-1975) in five selected riffle sections of Augusta Creek, Michigan, using an in situ chamber technique (Figure 2). Circular chambers, illustrated in Figure 6 , were constructed of 3.2 m m (1/8") thickness Plexiglas cylinders of telescoping diameters. The bottoms were 21.5 cm in diameter by five cm deep, while the lids were 22 cm in diameter by approximately 10.2 cm in height. A hole was cut in the lids for the insertion of a stopper in which a YSI (Yellow Springs Instrument Co.) self-stirring oxygen probe (Model 5420-A) was mounted. All bottoms and dark lids were opaqued with Krylon flat black paint and covered with two layers of black electricians tape applied in opposite directions to prevent light penetration (APHA et al. 1971). Chambers were 0.036 m “ 2 in area, held approximately 1500 ml of sediment and three liters of water. Chamber bottoms were filled with natural substrate and buried in riffle sections of Augusta Creek (Figures 1 and 2) where they were allowed to recolonize at least one month prior to experimentation. Just prior to an experiment YSI oxygen meters (Model 51-A temperature compensating) and oxygen probes were calibrated with oxygen probes in 100% humidity chambers which were submersed in the stream and equilibrated to stream temperature. Water samples were taken with a VanDorn water sampler and fixed in the field for Winkler titration of dissolved oxygen (APHA et al. 1971) at the beginning and OPAQUE L ID OXYGEN METER YSI 51-A SELF STIRRING OXYGEN PROBE ^ + Y S I //Y 5 4 ~ A C L E A R LID CHA M B E R BOTTOM Figure 6 . Plexiglas chamber designed for estimating community metabolism in stream riffle sections. 22 end of each experiment to check probe calibration. An average deviation of + 0.5 m g O 2 1 “ * (n = 38) was found from Winkler results for initial oxygens, while + 0.04 mg O 2 I -1 was the average deviation of final titrations (n «* 38). The precision of the YSI probe system was + 0.2 m g O 2 l - * (Yellow Springs Instrument Co.). Checks on probe drift and oxygen consumption of empty chambers were run at room temperature in the laboratory and in temperature controlled experimental stream channels with no significant changes observed (+ 0.2 m g O 2 1 “ *). A complete field procedure diagram is illustrated in Figure 7. At the start of an experiment, Plexiglas viewers were used to locate chamber bottoms in the substrate. Markers were not used so the bottoms would remain inconspicuous in the sediments, the first two or three bottoms located in the predawn light were selected as replicates, provided they were not washed out. This method of selection was considered random. Plexiglas lids were placed over each chamber and due to the closeness of fit, plus layers of electrician's tape, a tight seal was obtained (chambers could be lifted out and set on the stream bank with no water loss). Half-hour changes in oxygen and temperature were monitored using polarographic oxygen sensors equipped with electric stirrers. The stirrer provided a current within the chamber to better simulate stream conditions, although the flow was not laminar, it was sufficient to thoroughly mix the water within the chamber, as evidenced by the almost instantaneous dispersal of test dyes or observations of attached detritus and algae that indicated swaying to be similar to that outside of the chamber. Laboratory and field 23 Figure 7. Experimental design of in situ community metabolism field experiments. 24 FIELD PROCEDURE PLEXIGLAS CHAMBER BOTTOMS-------(21.5 c m diameter x 5 cm height) .FILLED WITH NATURAL SUBSTRATE BURIED IN RIFFLE SECTION COLONIZED 1 M O N T H OR MORE SUNRISE or START: OXYGEN PROBES CALIBRATED IN SITU ( 100% humidity calibration chambers) D AY 1 CHAMBERS STREAM 2 or 3 CHAMBER BOTTOMS COVERED W I T H LID (light or dark-22 c m diameter x 10.2 cm height) chamber volume - 3L OXYGEN AND TEMPERATURE MONITORED (1/2 hr. intervals) Light lid-sunrise to sunset (net community productivity) Dark lid- ~ 8 hrs (community respiration) r If 02 bubbles form or reading would go off scale (light or O 2 falls below 2 m g l -1 ( d a r k ) ^ ' SUNSET O R FINISH W A T E R SAMPLES FOR W INK L E R 02 initial a^d final OXYGEN MONITORED 1/2 hr intervals* (if probe was available) TEMPERATURE 1/2 intervals** INCIDENT LIGHT 1/2 hr intervals*** Gently siphon chamber water 63 itffl brass screen 1 Gently refill chamber stream water M E A SURE CHAMBER VOLUME REPLACE BOTTOMS IN RIFFLE M A P LOCATION OF EACH REPLICATE 1 Add material from screen I Initial 0? reading Continue DAY 2 REPEAT WITH OTHER LID SAME SEDIMENT SAME REPLICATES \ SEDIMENT TO LABORATORY (Figure 8 ) Figure 7. *YSI Model 51-A oxygen meter with M o d e l 54-A polarographic oxygen sensor **YSI Thermister JLJLJL Weston Photometer Model 756 25 studies have indicated currents of less than 15 to 2 cm s"* affect CR and mineral uptake in lotic systems (Odum and Hoskin 1957; Whitford and Schumacher 1961, 1964; Pfeifer and McDiffett 1975). The oxygen-probe stirrer is designed to create a current of at least 10 cm s“ l over the membrane (YSI Instrument Co.). Estimates of community metabo l i s m obtained with these chambers compared favorably (within expected variation for the reach) wi t h those in which laminar flow was provided (Bott et al. 1978). The stirrer was run in the field using 12 volt car batteries and a Wards Power Verter (Model 2V200B). Each experiment consisted of a two day sampling period. One day a light lid was placed over each replicate chamber to monitor net community productivity (NCP) from sunrise to sunset and the second day to monitor community respiration (CR) on the same substrate dark lids were employed (days were always consecutive). Dark runs were for variable lengths of time (generally d aylength), but continued until an oxygen change of at least 2 m g 1“ 1 occurred, which placed measurement error at 10% (+ 0.2 m g I"!). Dark runs followed by diel light runs were completed at the B Avenue and Nagel sites to compare CR rates obtained during daylight and dark periods. There were no measurable differences in the two rates, therefore daylight monitoring of CR continued (Table 4). Stream temperatures and levels of incident light were also monitored at half-hour intervals, using a YSI Thermister or a mercury thermometer and a Weston Photometer (Model 756), respectively. At experiment termination the chamber volumes were measured and the sediments taken back to the laboratory for removal of detrital content through decanting, and sorting of macroinvertebrates (Figure 26 Table 4. Comparison of community respiration rates obtained during daylight and dark periods from selected riffle sites of Augusta Creek, Kalamazoo County, Michigan. Dark Chamber Light Chamber-Night Date Site x g 0 2 /m^h“ l - range x g 0 2 /m^h- ! - range 07 June 1973 B Avenue 0.029 (0.021-0.037) 0.028 (0.025-0.031) 29 M a y 1974 Nagel 0.082 (0.063-0.100) 0.079 (0.068-0.089) 27 7). During the second year of sampling, the organic content of attached periphyton assemblages, and POM and algae incorporated in the travertine, was determined by ashing and reported as epilithon (g AFDW m “ 2; Figure 8). All POM was oven dried at 52 C to a constant weight (48 hours or greater). Ash free dry weight (AFDW) was obtained by ashing at 550 C for 24 hours (Figure 8). When epilithon AFDW was determined, detritus was gently elutriated from the inorganic sediments; then materials were collected separately, particle sized and A F D W determined for both components. Macroinvertebrates were removed from either entire chamber sediments or subsamples (40 ml of 500 ml) of a sediment slurry (Figure 8 ). Using Chi-Square analysis for random distribution of organisms, it was determined that subsampling was adequate (P < 0.05) to estimate this component (Elliott 1971). Macroinvertebrates were dried (52 C) and AFDW determined as for sediments. Duplicate estimates of community metabolism were completed on a monthly basis at the B Avenue site from January 1973 through May 1974 and at the Nagel site from August 1973 to August 1974. Seasonal estimates were made at five sites (Smith, B Avenue, Upper 43rd, Nagel and Kellogg Forest) from October 1974 through July 1975). Seasons were delineated as follows: Fall - September through November, Winter - December through February, Spring - M a r c h through May, and Summer -June through August. Parameters calculated are listed and defined in Table 5 (Hewlett-Packard 2100A Mini Computer). Regression analyses, involving individual estimates of NCP and CR and averages of the parameters listed in Table 5 with the independent variables light, temperature 28 LABORATORY PROCEDURE CHAMBER SEDIMENTS PARTICLE SIZED 16mm, 4mm, 1mm, 250 Jim, 75 Jim. ROCKS SCRUBBED FOR REMOVAL OF POM SMALL SEDIMENT WASHED AND DECANTED FOR REMOVAL OF POM I DECANTED FOR REMOVAL OF ORGANICS ORGANIC SIEVED — 75 jura brass screen LPOM (total or particle sized) -INORGANIC PARTICLES VOLUME BY WATER DISPLACEMENT t FILTRATE to 3 L VOLUME OF INORGANIC PARTICLES MEASURED BY W ATER DISPLACEMENT 3 100 ml SUBSAMPLES 0.45 jum MILLIPORE FILTERS FPOM MACROINVERTEBRATES REMOVED FROM WHOLE SAMPLE OR 3 SUBSAMPLES P O M ------------DRIED 50 C FOR 48 HOURS OR MORE MACROINVERTEBRATES J WEIGHED MUFFLED AT |50C for 24 HOURS WEIGHED FOR AFDW DETERMINATION TOTAL OR PARTICLE SIZED DETRITUS OR EPILITHON * ** Wild dissecting microscope model M-5 ( 50x). Mettler H-16 (organic) - Mettler P-163 or E-1000 Thermolyne model F-41730 muffle furnace. Figure 8. (inorganic). Laboratory procedure for determination of ash-fee dry weight (AFDW) of chamber sediments and macroinvertebrates. 29 Table 5* Procedure for estimating parameters of community metabolism experiments. Parameter Definition Net Community Productivity (NCP) O2 produced during the daylight period (sunrise-sunset) mg O 2 m ” 2d” l (hr, month) Final O 2 -Initial O2 x CV*x CF** Community Respiration (CR) m g O 2 m “ 2h“ l (day,month) O 2 used by the community-day and night. Initial O 2 - Final O2 x CV x CF run length (h) Gross Community Productivity (GCP) NCP + (CR/h“ l x Daylength) m g O 2 m " 2 d “ l (hr, month) "profit or debit” Net D a i l y Metabolism (NDM) GCP m “ 2d” l - 24 hour CR m“ 2d” l m g O 2 m " 2 d ” l (hr, month) P/R Ratio Photosynthesis/Respiration ratio GCP m " 2 d - l / 24 hour CR tiT^d- * Ash Free D r y Weight (AFDW) Organic content (g) of sample Ashed 550C (24 hours) Epilithon AFDW Organic content of inorganic sediment Total A F D W AFDW + Epilithon AFDW * CV ■ chamber volume JLJL * CF = conversion factor to m^ 30 and AFDW, were completed (Biomedical Computer Programs P-Serles (BMDP); Health Sciences Computing Facility, UCLA, NIH Special Research Resources Grant RR-3; V ax-11/780 digital computer, Kellogg Biological Station). The contribution of the epilithon as well as the detrltal flora of various sediment sizes to the NCP and CR during February - March, June and August at these five Augusta Creek sites (Figure 2), was determined using two Gilson Differential Respirometers Gilson 1963, Lawton and Richards 1970). (Model 20; Triplicate samples, picked macroinvertebrate free, of detritus and inorganic sediment particle-sized in 4 mm, 1 mm, 250 pm, 75 pm, and 0.45 pm categories, were run at the ambient stream temperatures under 15,000 lux (GE Power Groove bulbs) for estimates of NCP and in darkness for estimates of CR (Figure 9). Samples were allowed to equilibrate under the lights for one hour prior to 15 minute estimates over a three hour period. The respirometers were then darkened for an equilibration period and a subsequent three hour dark estimates. (1 hour), All vessel contents were filtered on pre-weighed 0.45 p m Millipore filters, dried at 52 C, weighed, and ashed at 550 C for 24 hours for determination of AFDW (Figure 9). Estimates of NCP and CR were expressed as pi O 2 g AFDW“ 1 and g O 2 m - 2d“ l (G. L. Spengler modification of Petersen's 1974 respiration program; Hewlett-Packard 2100 A Mini Computer at the Kellogg Biological Station Computer Facility). 31 Figure 9. Experimental design for Gilson Respirometer estimates of community metabolism of particle-sized substrate. 32 PARTICLE SIZED COMMUNITY METABOLISM PLEXIGLAS CHAMBER BOTTOMS-------(21.5 cm diameter x 5 cm height) -FILLED WITH NATURAL SUBSTRATE BURIED IN RIFFLE SECTION £ COLONIZED l^MONTH OR MORE EXPERIMENTAL RUN D A Y : ' GENTLY REMOVE CHAMBER BOTTOM-TRANSPORT TO LAB | (ice chest) INORGANIC PARTICLES GENTLY RINSED AND PARTICLE SIZED------- ► ORGANIC MATTER + RINSE WATER PARTICLE SIZED: 250 ym MACROINVERTEBRATES REMOVED FROM SEDIMENTS (Wild M-5 Dissecting Microscope) FILTRATE RINSE WATER TO 10 1 VOLUME- 3-100 ml SUBSAMPLES FILTERED (0,45 ym Millipore filters) -------------AUTOCLAVED REACTION VESSELS 2 ml filtered stream water 0.4 ml 20% KOH in side arm + filter 3 REPLICATE VESSELS/PARTICLE SIZE: 15 ORGANIC + 12 INORGANIC VESSELS 6 or more CONTROL VESSELS (filtered stream water) 2 GILSON MODEL 20 DIFFERENTIAL RE|PIROMETERS (set at stream temperature) LIGHT RUN: ONE HOUR EQUILIBRATION PERIOD UNDER LIGHTS (15,000-20,000 lux - GE POWER GROOVE BULBS) t t CLOSE RESPIROMETER: DARK R U N : 3 hr RUN - READINGS/15 m in ONE HOUR EQUILIBRATION IN DARKNESS (opaque cloths c|ver respirometers + lights out) CLOSE RESPIROMET|R: 3 hr RUN - READINGS/15 min RUN TERMINATION: VESSELS FILTERED.ON PREWEIGHED 0.45 ym MILLIPORE FILTERS ▼ DRIED 52 C > 48 hrs t WEIGHED (mettler.Balance Model H-16) T ASHED (Thermolyne Model F-41730 Muffle furnace) ASH FREE DRY WEIGHT (AFDW) / RESULTS: Figure 9. yl 0 2 /mg- l A F D W *“G O2 /M" 2 33 RESULTS AND DISCUSSION Sunlight reflecting off Augusta Creek is jubilant in early spring wherein lies a diatomaceous carpet... M a r c h 1974 Physical Parameters The range and average stream temperatures and incident light based on half-hour interval measurements at each site during the experimental runs, were used to characterize the conditions of metabolic estimates (Appendix D, Tables D 1-5). Temperature Based on weekly maximum-minimum temperature data from Augusta Creek sites (1973-1977; King 1978, Mahan 1980) temperature patterns were similar throughout the drainage with maxima ranging from 24.6 C at B Avenue to 29.6 C at Upper 43rd (Table 1). During this study similar trends were evident with maxima ranging from 19.9 C at B Avenue and Smith Sites to 24.8 C at Upper 43rd Site with Nagel and Kellogg Forest Sites attaining maxima of 23.9 and 22.2 C respectively (Appendix D ) . Average daily temperatures exhibited typical temperate region seasonal trends at all sites (Figure 10). M e a n daily temperatures ranged from 0.4 C at the Upper 43rd Site in January to 22.6 C at the Nagel Site in August. Maxim u m daily averages at the other sites were: Smith 17.6 C, B Avenue 17.6 C, Upper 43rd 21.7 C, and Kellogg Forest 19 C. Although large differences in temperatures were not evident, the first order Smith and B Avenue Sites had the lowest temperatures, the open Upper 43rd (second order) and Nagel (third order) the 34 20 S MI T H 10 )T T E M P E R A T U R E oc 20 UPPER 20 4 3RD 20 10 MONTH Figure 10. Average daily temperature values for selected sites in the Augusta Creek watershed, Kalamazoo County, Michigan (1973-1975). 35 highest, while those of the downstream forested third order Kellogg Forest Site were intermediate (Figure 10, Appendix D). The greatest daily temperature fluctuations occurred in the first order sections during spring and summer (10.5 C) and were similar to fluctuations reported for an Arizona desert stream by Bush and Fisher (1981). However the greatest average daily fluctuations were at the Upper 43rd Site (4.0 C) followed by the Nagel Site (3.4 C) and indicated the influence of lack of canopy (Table 6). The lower Nagel average emphasized the presence of cold springs and perhaps increased volume at a third order reach (King 1978), while the stability of the Kellogg Forest temperatures probably represented increased discharge and shade effects. Since the metabolism experiments were run on a consecutive day basis such daily fluctuations were important. Consecutive day differences were significantly less than daily differences in range and average and varied from 27 to 50% of the average daily temperature fluctuations (Table 6). Light (Irradiance) Light (= irradiance) patterns at the five sites followed expected temperate region seasonal and site trends (Figures 11 and 12). Shading was indicated at the Smith, B Avenue, and Kellogg Forest Sites with maximum levels at the Upper 43rd and Nagel sites (Figures 11, 12; Table 7). The highest average daily light levels were recorded in May at the Upper 43rd Site (90,213 lux), and the Nagel Site (84,631 lux) and the lowest level was 2,391 lux during January at the B Avenue Site. The highest light levels were recorded at all sites during April, May and June, reflecting the approach of the summer solstice. 36 Table 6 . Temperature fluctuations at sites of Augusta Creek, Kalamazoo County, Michigan (1973-1975). Site Daily Range °C Daily Average °C Smith* 0.5-7.1 3.0 0.4-1.5 0.9 30 B Avenue 0.3-10.5 3.3 0.1-3.0 1.1 33 Upper 43rd* 0.5-7.2 4.0 0.5-2.7 1.7 42 Nagel 0 .0- 6.0 3.4 0.0-5.2 1.7 50 Kellogg Forest* 0.0-5.8 2.6 0 .1- 1.2 0.7 27 * Seasonal estimates only (1974-1975) Consecutive Day Range Average °C °C Percent of Daily 37 SMITH 20 40 20 CO o f" U P P E R 4 3 RD 5 80 IX 60 40 20 J F M A A M S O N 0 M O N T H Figure 11. Average dally light values for the Smith, B Avenue, and Upper 43 Rd sites of Augusta Creek, Kalamazoo County, M i c higan (1973-1975). 38 80 NAGEL 60 40 o x H K E L L O G G X 80 0 FOREST _i IX 60 40 20 F M A A S 0 N 0 M O N T H Figure 12. Average daily light values for Nagel and Kellogg Forest Sites of Augusta Creek, Kalamazoo County, Michigan (1973-1975). 39 Table 7. Light intensity fluctuations (irradiance) at selected sites of Augusta Creek, Kalamazoo County, Michigan (1973-1975). Annual Fluctuations Daily Average (Lux) Site Smith* 12,054 6,268-23,358 28 42 B Avenue 13,213 2,391-47,709 30 46 Upper 43rd* 43,622** 12,208-90,213 100 152 Nagel 36,959 2,824-84,631 85 128 Kellogg Forest* 28,787 6,404-71,840 66 100 JL Seasonal data only ** Maximum Percent Maximum Lux Percent Kellogg Forest Lux Daily Average (Lux) 40 Leaf emergence at the shaded sites significantly decreased light levels in June, while abscission Increased light in October when incident light is on a gradual decline in the temperate latitudes (Figures 11 and 12). Lower light values at Smith and B Avenue, even during periods when leaves were not present, indicated the shading effect by tree "skeletons". The annual patterns of average daily lux indicated positive relationships with stream cover and width. Narrow first order reaches had miraimum light, wider unforested reaches had maximum and the 7.0 m shaded Kellogg Forest reach Intermediate levels (Figures 3, 4; Table 7). Autotrophic zones, according to the continuum concept, generally will be limited in natural stream reaches of 10 m or less in width (Vannote et al. 1980). Because of canopy alteration at the second order Upper 43rd and third order Nagel Sites, significant light inputs resulted. As well, significant levels occurred at the forested Kellogg Forest Site (66% of the open sites; Table 7). The percentage of maximum daily light (Upper 43rd) received by each site ranged from 25 to 85% at the Smith and Nagel sites, respectively (Table 7). If the Kellogg ForeBt is considered a typical shaded third order reach the first order sites received 42 to 46% of maximum light in contrast to 28 to 30% of the open site (Table 7). The difference in light at the two open sites may be accounted for by the orientation of the channel (Upper 43rd E-W and Nagel N - S ) , as shown by Kobayasi (1961) for a canyon stream; variability in days monitored; or the increased observations at the Nagel Site (n = 8 Upper 43rd, n = 30 Nagel). The complexity of the light regime at the shaded Kellogg Forest 41 Site, represented in Figure 13, illustrated the predictable variation in potential autotrophic zones during leaf-out of riparian vegetation and reflected the heterogeneity with which one must contend in estimating in situ metabolic activity, hopefully, viewed as a challenge rather than a deterrent to in situ measurements. Community Metabolism The interrelationships of autotrophic and heterotrophic components within lotic communities has been of interest for some time. Only recently, concomitant with this investigation and the advent of field chamber techniques, have estimates of community metabolism been made on discrete communities (Bombowna 1972, Pfeifer and McDiffett 1975, Marker 1976b, Bott et al. Lavelle 1979, al. 1981). 1978, Pennak and Sumner and Fisher 1979, Bush and Fisher 1981, Hornick et Such studies allow analyses of the producing or consuming modes of the community and thus, the autotrophic-heterotrophic balance within. Community metabolism studies have followed the approach for studying lotic systems outlined by Shelford and Eddy (1929) with several excellent artificial stream and laboratory chamber studies completed prior to in situ work (Whitford and Schumacher 1961, 1964; Edwards and Rolley 1965; Kevern and Ball 1965; Mclntire and Phinney 1965; Mclntire 1966; Maki 1974; Pfeifer and McDiffett 1975). Methods and standardization of data presentation for studies of community metabolism are still in a state of flux. Many methods have been presented, however, often no mention is made of what was measured (i.e. gross versus net community productivity); standardization of units (oxygen, carbon, calories, gram calories, chlorophyll (various *SZ6 I ^Tnf 52 uo UBS-pqojw ‘^311003 oozemexBX ‘qaaaO BtjsnSny ‘asaao,* 33oxiax a® suaa^qBd a q S n ^IT®a nos n n d =o a a i d d v a = e a aa v h s= • (W)»Nva 1 h 01a ifl oud a o N v i s i a im 0 i v aa i s a i w 3 1 S £ 0003 • • • • • • • • e o e •• • •• • • • • • • • • 0081. •••OOOOOI«lt« •••••••••• 0091 ii«ooooii«iioeooo99S0oee ••••oooeeeeeeeeeeeeeeeee ••oeoeeoeeoeeeeeeeeeeeee oon 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0031- ooooeeeoooo»«*«««ooooooo O O O O O I M I f M M O O O O O O O O O O O O O O O O O I I M I t l t t O O O O O O O O oo#*##oooooooooooooooooo o o o o o o o o i 0 0 0 1- « * i o » « o o » « i o o o o ••••••••oooooooooo****** 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 oooooooooooooooooooo*«*« •••••••••••••••••••••••• XNV91H0IU XNv 8 1J31 008 (HOURS) • • 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 T I ME • • • o « « • • • o o « • • • •• • • • • • • I f O O I I * • • • • 0 « • • • • • •• • •• 43 pigments or total), glucose, dry weight, AFDW); or how the results are expressed (m“ 2, m ” 3, m g - 1chlorophyll, g“ l d ry weight, g” l AFDW, per chamber, etc.). Various estimates of PQ (photosynthetic quotient) and RQ (respiratory quotient) also complicate the situation of converting oxygen values to carbon and based on the few studies completed on periphyton communities, these quotients appear to be highly variable (PQ 1.45 to 2.6, RQ 0.85 to 1.1; 1978, Bott and Ritter 1981). Schindler et al. 1973, Bott et al Oxygen and estimates of community metabolism have been shown to measure similar magnitudes of productivity in coral reef communities (Littler 1973) and littoral zones of freshwater lakes (Hunding and Hargrave 1973); yet Schindler et al. (1973) compared four methods (oxygen, dissolved inorganic carbon, uptake and disappearance) and found little agreement other than general trends. They attributed the discrepancies to inappropriate PQ and RQ values for the gaseous methods. Ritter (1981) compared Bott and and oxygen estimates of community productivity in riffle communities and reported similar patterns of productivity, but lower estimates were obtained using the method; PQ values varied, which reinforced the necessity of measuring PQ, as values were generally higher than previously reported. Since the conversion of oxygen uptake to carbon fixation is not reliable, data from this investigation will be presented, as measured, as oxygen and compared with those studies reported in oxygen units. (Table 8 ). This study, based on intact riffle communities and not Isolated substrates, should yield a more realistic estimate of in situ metabolism of riffle reaches in Augusta Creek. Table 8. Literature estimates of in situ community m etabolism in streams (g O 2 m 2d- !). Date System Period NCP* CR GCP G 6‘thberg & Karlstrom 1975 Rlckle&n River-N,H Sweden Fall & Summer ----- 0.5-0.7 ---- Marker 1976 Bere Stream England Annual ----- 0.3-7.2 ---- White Clay Creek-N-3 Pennsylvania August 0.7-3.5 0.7-3.1 1.5-7.1 0.94-1.05 Chippewa River-N-4 Michigan Seasonal ----- 0.5-2.4 0 . 8 - 4 .6 1 .4-1.8 Fort River-N'3 M assachusetts Annual 1.5-7.0 0.1-6.5 0 .01- 0.68 Sycamore Creek-CM Ari zona Summer ----- 4.4-12.5 0.96-2.3 River Raba-N-2 Poland April-October ------ Danube River Annual Author/s P/R C H A MBER ESTIMATES Bott et al. Litke Sumner & Fisher Bush & Fisher 1978 1978 1979 1981 5.1 LIGHT BOTTLE-DARK BOTTLE Bombowna Ertl & Tomajka 1972 1973 0.96-1.32 0.9-0.55 Table 8. (cont'd.) Author/s Date System Period NCP* CR GCP P/R 0.13-11.34 ------ 0.04-14.0 0.1-1.1 L IGHT BOTTLE-DARK BOTTLE (Cont'd.) H e l a n et al. 1973 Brodskrf Brook-N C zecho slavakia June-March Itchen River-N England Nov.-March 4.2-20.2 Lark River-P England Nov.-May 35.0-53.0 0.53-0.39 0.01-1.1 White River-E Indiana Annual 5.8-18.6 0.6-1.1 UPSTREAM-DOWNSTREAM M ETHOD Butcher et al. Butcher et al. Hornuff Hornuff Hoskln 1930 1938 D enham Hornuff 1930 ** 1957 1957 1957 1959 5.5-14.0 Blue River Oklahoma 1.03 M o u n t a i n Fork Oklahoma 2.17 Honey Creek Oklahoma 2.66 Neuse River North Carolina 0. 2- 0.7 Table 8. (c o n t 'd .) Author/s Date System Period Pledomont Stream-N,T Georgia Aug.-Dec. NCP* CR GCP P/R UPSTREAM-DOWNSTREAM (Cont'd.) Nelson and Scott Noyce*** Baumgardner*** Duffer & Dorris Hannan**** Stockner Eley*** Flemer Flemer 1962 1965 1966 1966 1967 1968 1970 1970 1970 Cottonwood River Kansas 0.43-2.55 0.7-0.27 0.16-0.26 ------ Skeleton Creek-P Oklahoma 0 . 6 - 9 .6 1.5-7.0 0 . 6 - 9.6 1.5-7.0 ------ O' Blue River-N-2 Oklahoma June-August San Marcos River-N,M Texas ------------------ Ohanapecosh-N,H Washington ------ Skeleton Creek Pennsylvania Annual Roritan River-N N e w Jersey 4-5 Roritan River-E N e w Jersey 4-5 May-Sept. 6.1-19.9 1.5-48.0 ------ 4.1-19.9 2.5-27.4 0.33-8.64 0.56-2.7 0.89-11.3 ------- 1.1-5.0 33.5 13.6------ ------ ------ 3.7-5.4 2 . 0 - 8 .8 0 . 8 - 1 .6 Oct.-Dec.--------- ------ 0.9-1.9 2.5-16.7 0.9-1.9 Table 8. (cont'd.) Author/s Date Period {jQp* CR GCP P/R Roritan River-E 4-5 N e w Jersey Oct.-Dec. ------ 7.0-11.3 2.5-25.1 0.7-2.0 Ne w Hope-N N. Carolina 2 years----------- ------ 0.4-13.0 0.21-9.0 ------ Buffalo Creek-N Pennsylvania August 2.0-2.3 4.9-6.3 System UPSTREAM-DOWNSTREAM (Cont'd.) Flemer Hall 1970 1972 McDiffett, Carr & Young 1972 Cole 1973 Cole Profet & Ransom Profet & Ransom Profet & Ransom de la Cruz & Post ------ 2.6 .e- 1973 1974 1974 1974 1977 Spring Creek-N Pennsylvania Annual ------ 2.0-13.0 2.0-17.0 0.3-2.0 Spring Creek-E Pennsylvania Annual ------ 12.0-18.0 4.0-20.0 0.2-0.9 Kansas Summer------------ ------ 5.52-26.6 1.0-24.2 ------ Cottonwood River-N~4 Kansas Summer------------ ------ 19.9-26.4 9.9-32.3 ------ Walnut River-N Kansas Summer 7.2-12.0 ------ Catahoula River-N-2 M i ssissippi Seasonal---------- ------ Cedar Creek-N ------ 5.1-5.7 7.88-50.1 4.1-28.4 0.24-0.57 Table 8. (cont'd.) Author/s Date System Period NCP* CR GCP P/R UPSTREAM-DOWNSTREAM (Cont'd.) Gelroth & Marzolf G e lroth & M a r zolf 1978 1978 Lost Creek-N-2 Kansas Ju l y Lost Creek-C-2 Kansas Ju l y 0.54-1.88 0.51-1.21 2.11 * = All values g O 2 m - 2d"l except P/R. ** = Classification of system: N = N a t u r a l - O r d e r ,Order approximated **** Taken from Profet and Ransom, 1974. = Taken from Mathis, Taylor and Myers, from information in study; E = Effluent; M = Ma c r o p h y t e dominated; H = Hot Spring; = 4.16 All studies used oxygen techniques. p = Polluted; *** 0.64-0.95 1969. C = Channelized; T = Turbid; CM = Cladophora mat; H = Headwater. 1.97 49 Gross Community Productivity GCP, the most cited parameter in the productivity literature, indicates the maximum carbon producing capacity of the autotrophic community and is estimated by the addition of NCP plus CR for the photo-period. autotrophs. M u c h of the GCP m a y in turn be respired by the Verduin (1956) noted that the autotrophic community respired up to 80% of the GPP in phytoplankton studies while Hargrave (1969) and Hunding (1971) reported 13 to 15% of GCP respired by periphytic algae. In the present study NCP and CR were obtained from the same sediment on consecutive days, therefore it was assumed that similar heterotroph respiration rates were involved in both NCP and CR estimates. Intrasite seasonal and intersite variations in GCP occurred annually at riffle sites of Augusta Creek (Figure 14). values were recorded for the first order sections The lowest GCP (Smith 0.09 to 0.18 g O 2 m “ 2d- l and B Avenue 0.01 to 0.59 g O 2 m ” 2d“ l), the open Nagel and Upper 43rd were highest (0.53 to 5.35 and 0.34 to 3.25 g O2 m “ 2d_ l respectively), and the shaded third order Kellogg Forest reach was intermediate (0.53 to 2.16 g O 2 m " 2 d “ l; Figure 14; Appendices A, B). to highest: On an annual average basis they ranked from lowest Smith, B Avenue, Kellogg Forest, Upper 43rd, and Nagel (Table 9). Odum (1956) stated that streams may well be among the most productive environments on earth. However, his statement was based on standing crop estimates and ecosystem studies, using the upstreamdownstream oxygen method for estimating productivity, enriched by organic pollution. in streams often Literature GCP values using the 50 SM ITH 0. 5 B AVENUE 0- 5 U P P E R 4 3 RD NAGEL KELLOGG FOREST MONTH Figure 14. Average GCP ( + SE) estimates for selected riffle communities in the Augusta Creek watershed, Kalamazoo County, Michigan (1973-1975). Table 9. Average community metabolism parameters fop experimental runs completed at five riffle sites of Augusta Creek, Kalamazoo County, Mic h i g a n 1973-1975. X NCP X CR X GCP X NDM X EPILITHON X TOTAL g AFDW m -2 gA F D W m“ 2 gAFDW m " 2 g m- 2d~l g n T 2d~l g m - 2d"l g m " 2d“ * N S.E. C.V.% -0.16 4 0.05 56.3 0.56 4 0.10 35.3 0.13 4 0.02 30.7 -0.44 4 0.10 46.1 0.21 4 0.03 26.9 388.2 4 49.4 25.5 1763.2 4 161.2 18.3 2151.8 4 202.2 18.3 B Avenue N S.E. C.V.% -0.02 16 0.02 476.5 0.53 16 0.06 43.7 0.26 16 0.05 73.7 -0.26 16 0.03 50.9 0.45 16 0.06 53.1 290.4 14 36.7 47.4 2306.8 4 191.0 16.6 2625.5 4 204.1 15.5 Upper 43rd N S.E. C.V.% 1.24 4 0.40 64.0 1.79 4 0.49 54.3 2.18 4 0.63 58.1 0.45 4 0.31 134.9 1.20 4 0.17 27.9 240.1 4 31.5 26.3 3558.5 4 702.5 43.1 3503.5 4 417.9 41.0 Nagel 1.80 15 0.26 56.0 1.88 15 0.26 53.6 2.89 15 0.40 52.9 1.01 15 0.24 91.0 1.57 15 0.12 29.1 291.1 14 17.3 22.3 3507.5 4 658.5 37.6 3850.5 4 671.2 34.9 0.94 4 0.23 48.8 0.75 4 0.25 65.4 1.36 4 0.38 55.0 0.60 4 0.17 56.0 1.97 4 0.15 55.4 282.5 4 28.9 20.5 2675.0 4 378.0 28.3 2957.0 4 350.5 23.7 Site Smith N S.E. C.V.% Kellogg Forest N S.E. C.V.% X P/R X DETRITUS upstream-downstream method (Odum 1956) range from 0.51 g O 2 m - 2d -i in a natural second order creek of the prairie biome in Kansas (Gelroth and Marzolf 1978) to 48.0 g O 2 m _2d"l in Blue River Oklahoma (Duffer and Dorris 1966; Table 8 ). Chamber and light-dark bottle estimates ranged from 0.09 g O 2 m - 2d “ l on revetments of the Danube River (Ertl and Tomajka 1973) to 12.5 g O 2 m -2d"l in Cladophora mats of Sycamore Creek, Arizona (Bush and Fisher 1981). Chamber techniques have been developed to monitor rates of community metabolism in environments with low productivity and in riffle sections where more sensitive methods are essential. The rates of 6CP from riffle communities in Augusta Creek ranged from 0.09 to 0.59 g O 2 m - 2d"^ Smith Site to the open Nagel Site, respectively. at the shaded These rates are in the lower range of those reported for lotic communities (Table 8 ). Annual rates at the Smith and B Avenue sites are the lowest in situ measurements reported to date (Table 9; Appendix A). The Augusta Creek GCP maxima are similar to GPP rates of phytoplankton communities while the range is equivilent to terrestrial levels of GPP reported for deserts to temperate deciduous forests (Wetzel 1975a). The narrow, shaded Smith Site had the lowest seasonal GCP values (Figure 14; Table 9). Although, on a qualitative basis algae were never obvious at the Smith Site, maximum GCP rates were recorded in M a y prior to leaf emergence, which was during the period of visual diatom bloom in the rest of the watershed. The B Avenue site also exhibited low levels of GCP on an annual basis, but averaged twice those at Smith Site (Figure 14; Table 9). Maximum rates were measured in June, but within the range of variation rates were similar from April through October. Algal growth was never 53 obvious In the shaded reaches at B Avenue, but microscopic examination revealed a diverse array of pennate diatoms during the summer months. Annual rates of GCP at this site were similar to those obtained by Nelson and Scott in a 60-m wide piedmont stream in Georgia (1962); they attributed their low values to the presence of high silt loads limiting light penetration (Table 8 ). The differences in average daily light received by the two first order Augusta Creek study sites was probably insignificant, as both sites were heavily shaded (Figure 3). The averages for the June to October period were 9,055 lux (n=4) and 8,512 (n=15) at Smith and B Avenue, respectively. Estimates of light-saturated photosynthetic rates in periphyton communities range from 8,000 to 62,000 lux (Kobayasi 1961, Mclntire et al. 1974, Mclntire and Phinney 1965, Adams and Stone 1973, Lester et al. 1974). The averages for Smith and B Avenue Sites indicated potential shading effects with frequently less than light-saturated photoperiods (Appendix D). at both sites; but, O 2 m ” 2d~l) CR rates were similar Smith had slightly higher CR rates (0.56 g compared to B Avenue (0.53 g O 2 m - ^ " ! ) , estimates of GCP would tend to be at the Smith Site. therefore, higher Greater GCP at the B Avenue site was probably due to increased algal photosynthesis rather than an effect of CR rates. The Upper 43rd and Nagel Sites were approximately 13 times more productive than the first order reaches (Figure 14). They also received much higher average light inputs (43,622 lux Upper 43rd and 36,959 lux Nagel) and exhibited a definite succession of attached algae. During the spring, diatom blooms, often nearly pure populations of Meridion circulare Agardh, carpeted all available 54 substrates. Diatoms were followed in late spring by patches of epipsammic Vaucheria, followed by a patchy summer distribution of Cladophora and blue-green mats dominated by Oscillatoria. A second pulse of diatoms, a mixed association of pennate forms, developed in autumn. At Upper 43rd the rates of GCP increased in May, remained high throughout the summer and fall, and then decreased significantly during winter. Based on chamber estimates from larger order riffle communities (Table 8 ), the rates of GCP for Upper 43rd Site (0.34 to 3.25 g 0 £ m ” 2d _1 were higher than anticipated for a second order reach. By comparison, in a natural shaded second order section of Lost Creek, Kansas, Gelroth and Marzolf (1978) reported rates of 0.51 to 1.21 g O2 m ~ 2d~^, while in a cleared and channelized section rates of GCP rose to 4.16 g O 2 ra“ 2d"l, similar to maxima for the Upper 43rd reach. The third order Nagel Site followed the same seasonal trend as Upper 43rd, but maximum productivity was measured during August (Figure 14) and represented the highest GCP values recorded. These high Augusta Creek rates are similar to those reported by authors using the upstream-downstream technique in various systems (Table 8 ) and chamber techniques on an open third order stretch of White Clay Creek, Pennsylvania (Bott et al. 1978). The range of 0.66 to 5.35 g O2 m “2d ” l obtained for Nagel Site was similar to the chamber GCP estimates of Marker (1976b), Litke (1978), and Sumner and Fisher (1979) (Table 8 ). Based on these comparisons, canopy removal at the Nagel Site shifted the position of this site in the theoretical continuum (Vannote et al. 1980) to reflect the primary productivity of 55 a naturally more open fourth order reach. The shaded third order Kellogg Forest site averaged 47% the GCP of the cleared Nagel site (1.3 g O 2 n f ^ d - *)* An increase in spring plateaued to summer values, then decreased during the fall and winter (Figure 14; Appendix A). The range of 0.53 to 2.16 g O 2 m -2d- ^ over the annual period was within the range of studies cited in Table 8 . Both maximum and minimum values were lower than those obtained at the cleared Nagel Site and from a naturally shaded fourth order Chippewa River, Michigan, riffle section (Litke 1978). Average light at the Kellogg Forest Site was 66% that of the open reaches (Table 7) wi t h patchy algal development that was never as obvious as at the open Nagel Site. Similar differences in shaded versus unshaded reaches of the same order have been reported by Gelroth and Marzolf (1978), Gregory (1980), Hornick et al. (1981). M a x i m u m levels of GCP occurred during M ay and June at Smith, B Avenue and Upper 43rd Sites, while the Kellogg Forest maximum was in July, and Nagel in August (Figure 14). Seasonal trends such as these are typical of aquatic systems of the temperate region (Teal 1957, Cushing 1967, Stockner 1968, Hargrave 1969a, Flemer 1970, Gargas 1970, Hunding 1971, Hall 1972, Vannote and Ball 1972, Helan et al. 1973, Schindler 1973, Profet and Ransom 1974, Marker 1976b, de laCruz and Post 1977, Gelroth and Marzolf 1978, et al 1981). Sumner and Fisher 1979, Hornick The lowest GCP occurred during the winter at all sites of Augusta Creek, a pattern also characteristic of studies in which GCP was monitored on an annual basis (Cushing 1967, Ertl and Tomajka 1973, Marker 1976b, Stockner 1968, Sumner and Fisher 1979). Viewing natural streams as a continuum (Vannote et al. 1980), the hypothetical Augusta Creek site ranking based on levels of GCP would proceed from headwater downstream (Smith, B Avenue, Upper 43rd, Nagel, Kellogg Forest). were: 9). Based on increasing in situ estimates of GCP they Smith, B Avenue, Kellogg Forest, Upper 43rd, and Nagel (Table The open Upper 43rd and Nagel Sites illustrated the effects of perturbation by man, while the more natural state of a third order reach was shown by the Kellogg Forest results. The occurrence of open areas increases the complexity of community metabolism patterns in temperate woodland stream riffle sections. Natural and man-produced open areas are typical of present day stream systems and must be considered in applying the continuum concept. Such discontinuities in the continuum are treated in the intermediate disturbance hypothesis of Ward and Stanford (1981). Net Community Productivity Net community productivity (NCP), defined as the rate of carbon reduction minus usage in community respiration during the photoperiod, can be measured directly as oxygen gain or loss using the oxygen change technique. Radioactive carbon tracer uptake experiments (l^C) approximate NCP rates (Vollenweider 1969, APHA et al 1971, Schindler et al. 1973, Hunding and Hargrave 1973, Matheke and Horner 1974, Bott and Ritter 1981). NCP is of interest from an ecological standpoint since it represents the carbon available for use, storage, or export within a community. Likens (1975) points out that this is the commodity of interest to consumers and would therefore be important in community and ecosystem modeling. Pratt and Burkson (1959) stated that carbon produced in excess of metabolic need is equivalent to that 57 quantity that can be exported without change in community standing crop. The majority of work on primary productivity in lotic systems has been orientated towards quantitative or distributional studies (GCP) or laboratory experiments examining effects of physical parameters on rates of primary productivity. Few estimates of NCP exist in the literature (Grzenda et al. 1968, Stockner 1968, Hunding 1971, Adams and Stone 1973, Littler 1973, Cadee and Hegeman 1974, Bott et al. 1978, Bott and Ritter 1981) and not all of these cited are from lotic systems. Estimates of NCP at the five Augusta Creek sites ranged from a miximum of 4.02 g O 2 m “ 2d**l at the Nagel Site in August to a low of -0.25 g O 2 m “ 2d"l at the Smith Site in October (Figure 15; Appendices A, B ) . Ranked in increasing order of average NCP m ~ 2 d ” l the sites were in the same order as for GCP (Table 9; Figure 14). Although photosynthesis was taking place at the first order sites, as indicated by the positive annual GCP values, the NCP was zero or less with the strongest negative trend at the Smith Site (Figures 14, 15; Table 9). Dependence of headwater streams on detrltal input from the watershed with primary production often responsible for less than one percent of the annual carbon budget has been shown by Minshall (1967), Kaushik and Hynes al. (1971), Fisher and Likens et al. (1978). (1973), (1968), Coffman et Sedell et al. (1975), and Bell The low rates of NCP coupled with the higher detrital standing crops found at the Augusta Creek first order sites indicated a dependence on allochthonous carbon (Table 9). As discussed for GCP, average light inputs on an annual basis and especially during periods of leaf-out, were undoubtedly limiting the autotrophic potential at 58 SMITH 0.5 0 2 U P P E R 4 3 RD a NAGEL a o z IK J F M A M J J A S O N D MONTH Figure 15. Average rates (+ SE) of NCP at selected riffle sites in the Augusta Creek watershed, Kalamazoo County, Michigan (1973-1975. 59 these sites (Table 7; Appendix D). In addition to light limitation, grazing may also have accounted for the lowered levels of NCP at Smith and B Avenue Sites. Hargrave (1969a, 1972) reported low production in the littoral sediments of Lake Marlon and Lake Esrom which he attributed to grazing. Kesler (1981), documenting significant grazing effects on attached algae, reported grazing pressures by the midge Corynoneura scutellata to account for 3 to 70% of the NCP. Populations of the scraper caddlsfly Glossosoma nigor were abundant at the B Avenue site, and occurred at lesser densities at all other locations, therefore, one might speculate that they were influential in reducing the standing crop of algae and NCP. The open Upper 43rd and Nagel sites had the highest autotrophic potential of the Augusta Creek sites studied with no significant difference in NCP evident between sites (Figure 15; Table 9). Summer rates measured at Upper 43rd and Nagel sites were similar to those reported by Bott et al. (1978) for an open third order reach of White Clay Creek, Pennsylvania. N CP at the Kellogg Forest Site was significantly lower than the Nagel reach, but similar to Upper 43rd in average NCP (Table 9). The major divergence was notable during the summer leaf-out period (Figure 15). During this period the average light at the Kellogg Forest was 3,493 lux d“ l, similar to January levels at this site (Appendix D), and well below light saturation levels of periphyton communities (8,000 to 62,000 lux). It has been estimated that in aquatic communities 15 to 80% of the GCP m a y be respired by the in place community (Verduin 1956, 60 Hargrave 1969a, Hunding 1971), which leaves a net gain of 20 to 85% as NCP. The average NCP as a percent of average GCP for Augusta Creek sites was highly variable: -128.5% Smith, -7.6% B Avenue, 57% Upper 43rd, 60% Nagel, 69% Kellogg Forest. The shaded first order sites wi t h negative average deficits are unique to the primary productivity literature, while the open sites (Upper 43rd and Nagel) and shaded third order (Kellogg Forest) site, averaging 62% NCP, were similar to the estimated range in the literature. This again illustrates the dependence on allochthonous carbon sources and the heterotrophic nature of shaded first order Augusta Creek sites and obvious shifts in the magnitude of NCP due to clearing illustrated by the comparison of the Nagel and Kellogg Forest Sites. Community Respiration Community respiration (CR) is the rate of oxidation of organic matter to inorganic carbon dioxide by the autotrophlc and aerobic heterotrophic components combined. In this study CR was measured on intact riffle communities including the smaller particle-sizes of sediment and detritus versus selected stream substrates placed in chambers. Intact in situ community simulation must be kept in mind when comparing CR rates of this study with other studies, because inclusion of all partlcle-slzes may increase CR estimates, since small particles may be heavily colonized by bacteria and to some extent fungi and algae (Suberkropp and Klug 1974, Madsen 1975). However, rates may also be lower due to lack of community disturbance and less exposure of surfaces to higher oxygen levels. No attempt was made in the present study to partition CR. 61 Hargrave (1969a) attributed 33% of CR to macroinvertebrates, 45% to bacteria, and 15% to algae and Edwards and Rolley (1965) estimated 40% respired by macroinvertebrates in soft sediments. studies were from lentic communities. Both of these On a biomass basis, the macroinvertebrates were a small percentage of the total organic content of the Augusta Creek substrates and therefore might be expected to account for less than the estimated 40 to 45%. W i t h early interest in the effects of pollution on oxygen uptake, estimates of CR have been made in lotic systems since the 1930's (i.e. Butcher et al. 1930, Calvert 1933). Since Odum's (1956) introduction of the upstream-downstream procedure for estimating community metabolism in streams, many estimates of CR have appeared in the literature, but few in situ chamber estimates have been made (Table 8 ). Rates of CR ranged from 0.24 to 3.67 g O2 m " * ^ - ! during January at the B Avenue Site and July at the Nagel Site, respectively (Appendices A, B; Figure 16). The annual averages (g O 2 m ” 2d“ l) for each site ranked in increasing order were: B Avenue 0.52, Smith 0.56, Kellogg Forest 0.75, Upper 43rd 1.79, and Nagel 1.88; this pattern was similar to that of ranked sites for GCP and NCP (Table 9). Smith and B Avenue CR values were very similar, with the lowest rates observed in the winter, spring, followed by significant Increases in stable summer values, and decreasing rates from late fall to low winter values (Figure 16). The Smith Site had an autumn maximum in CR during leaf-input, but was only based on four observations. Further, a more comprehensive data set from B Avenue did not show such a maximum (Figure 16). Autumnal peaks in CR have been reported and 62 1 SMITH 0 1 0 2 1 0 NAGEL 3 2 1 0 KELLOGG FOREST 1 0 MONTH Figure 16. Rates of average CR (+ SE) for selected riffle communities in the Augusta Creek watershed, Kalamazoo County, Michigan (1973-1975). 63 related to Increased detrltal Inputs In various other systems (Teal 1957, Nelson and Scott 1962, Flemer 1970, Hargrave 1972, Maki 1974). Significantly higher rates of CR were observed at the Upper 43rd and Nagel sites than the first order reaches (Table 9). observations, Based on four the Upper 43rd Site had lowest CR during the winter, but steadily increased to a July maximum and remained high through the fall (Figure 16). Although the Nagel Site illustrated a more irratic pattern, maximum and minimum rates occurred during the same periods as at Upper 43rd. The highest CR for all sites was observed during July at the Nagel site (Appendices A, B; Figure 16), which was also during the period of maximum light, day-length, development (Appendix D). temperature and Cladophora Rates at both the Nagel and Upper 43rd Sites declined in August, possibly due to the decline of the green and blue-green community. Other studies noting summer maxima include Butcher et al. (1930), Stockner (1968), Flemer (1970) Vannote and Ball (1972), de laCruz and Post (1977), and Sumner and Fisher (1979). The average CR at the Kellogg Forest was intermediate among the sites (Table 9), but most similar to the rates at Smith and B Avenue sites. Only during the summer were CR values for the three sites significantly different (Figure 16; Table 9). Summer increases at the Kellogg Forest were attributed to increased light availability during this period (Appendix D, Figures 11, 12). The similarity between the Kellogg Forest third order reach and the first order sections is the relationship predicted by the continuum concept (Vannote et al. 1980). In the open reaches increases in CR as well as primary productivity were observed with rates of average CR (1.8 g O 2 m “ 2d“ l Upper 43rd, 1.9 g 0£ m ” 2d“ l Nagel) similar to the open third order 64 White Clay Creek, Pennsylvania (Bott et al. 1978; Tables 8 , 9). CR values obtained for riffle communities of Augusta Creek were generally in the low range of upstream-downstream studies reported in the literature and within the ranges reported for chamber estimates (Table 8 ). Thus, patterns of CR observed in Augusta Creek were not unusual, but reinforced, by in situ m e a s u rements, the general magnitude of respiratory activity in natural and perturbed reaches of first through third order riffle sections of temperate streams and specifically illustrated the effects of riparian zone clearing. Community respiration, ranked as a percentage of GCP (annual average) for Augusta Creek sites was: Smith 445%, B Avenue 200%, Upper 43rd 82%, Nage 60%, and Kellogg Forest 55%. Literature values * of CR/GCP as a percent typically range from 13 to 68% (Kobayasi 1961, Pamatmat 1968, Hunding 1971, Gallagher and Daiber 1973); Likens 0 to 95%. (1975) The first order sites did not fit within the latter range, apparently due to low NCP. Such an analysis further indicates the dependency of these first order reaches on allochthonous carbon inputs. A definite and inverse relationship, especially at higher temperatures, was seen between NCP and CR at these two first order sites, and to a lesser extent at Upper 43rd Site. P/R Ratio The ratio of GCP to CR (P/R) over 24 hour periods is a valuable integrator of community metabolism, which allows comparisons and rankings of communities. As Humphrey (1975) noted, Ryther (1954) and Steele (1965) proposed P / R ratios as the main measure of community physiological state and, therefore, useful in studying marine and 65 lentic freshwater systems. Odum, comparing the magnitude of in-stream photosynthetic production (autotrophy) to the import of organic matter from the landscape (heterotrophy) as energy sources, ratios in flowing water systems. first applied P/R The use of this parameter in stream classification was further reinforced by Hynes (1970) and Pieczyn/ka (1970). Fisher and Likens (1973) proposed that the P/R ratio should be used in conjunction with the import/export ratio in characterizing a stream reach. Vannote et al. (1980) stated that estimates of P/R allow characterization of the biological processes of a stream reach as to the productive (autotrophic) or consumptive (heterotrophic) mode, thus the present community usage of various inputs is indicated. However, caution must be exercised; if the P/R ratio is <1, the source of carbon (i.e. Imported, or stored from earlier autochthonous production) cannot be delineated from the P/R alone. Pavetid" et al. (1976) suggested monitoring P/R in running waters in the spring, when light and dark periods are equal, to characterize a reach physiologically, but unfortunately their usage of Pg/CR night caused confusion with the conventional terminology (Pq/CR 2 4 )* Each Augusta Creek site displayed a constant pattern in P/R (Figure 17). The ratios measured at the Smith and B Avenue sites were consistently below one, but above one at Upper 43rd, Nagel and Kellogg Forest. (Figure 17; Appendices A, B). Such consistency has been reported by other lotic investigators (Stockner 1968, Cole 1973, Sumner and Fisher 1979) and by Ganning and Wulff (1970) in brackish rock pools. Hall Others such as Flemer (1970), Vannote and Ball (1972), (1972), and Bott et al. (1978) reported fluctuations in P/R from greater than to less than one in stream reaches and Mathis et al. 66 1 SMITH 0 1 O UPPER 43 RO 1 K .250 < .075 a .45 >m 16 4 1 .250 .075 .45 jim 16 .250 •075 .45 jum k i l l H k t k k k W 50 g Figure 20. KELLOGG J ^ ftjrest SP SU SEAS ON Seasonal distribution of particle-sized detritus from selected riffle sections of Augusta Creek, Kalamazoo County, Michigan (1973-1975). 79 low stable values winter and summer (Figure 19). Although variance was high these data suggested a spring input (Figure 19) of CPOM. Mahan (1980) found the highest CPOM inputs at Smith followed by Kellogg Forest, Upper 43rd (wooded section) and Nagel Sites. sites leaves were the dominant form of CPOM. At all The CPOM inputs in Smith Site were significantly higher during autumn and spring, with spring CPOM averaging 56% of the fall input. The quality, however, shifted from a dominance of leaves in fall to wood in spring. showed no marked fluctuation in inputs. The other sites The disturbed Nagel site was by far the lowest in CPOM inputs (Mahan 1980). The consistency noted in second and third order tributary riffle sections in this study was also reported for a fourth order riffle of the Chippewa River, Michigan (Litke 1978). Such stability probably resulted from more continual autochthonous Inputs mediating allochthonous inputs over an annual period. The stability of the community and carbon inputs were indicated by the P/R ratios and NDM of these sites (Figures 17, 18). Naiman and Sedell (1979b) suggested that this stability indicated a balance in processing, storage, export and import. If this is so, changes in P /R would serve an an indication of changes within the communities. The relatively stable patterns observed for total detrital standing crop (Figure 19) are, however, not indicative of processing, changes in particle-*size composition on a seasonal basis, or quality (i.e. labile-refractory nature) or turnover time of particles. Distribution of particle-size on a seasonal basis (Figure 20) illustrated changes occurred within the detrital pool. Although total detrital standing crop levels may be consistent at a site, shifts in 80 particle-size composition may illustrate dynamic changes within the riffles. All sites were dominated by FPOM with Smith Site having the highest annual average and Upper 43rd the lowest (Figure 20; Table 11). Greater fluctuations were found in the CPOM over the year than in the FPOM standing crop as indicated by the C.V.% (x CPOM 50.6 and FPOM 30.8; Table 11). The most stable and also the lowest CPOM standing crop was found at Upper 43rd, while the FPOM was most consistent at the Nagel site. Shifts in particle size at Smith were not large, but by spring 16 and 4 m m particles were not observed and the 250 y m size decreased, either by processing or export, which were not monitored in this study. A more marked pattern was seen at B Avenue where the large autumn standing crops were reduced annually, which indicated significant export. However, a study of retention of basswood (Tilia) leaves at this site indicated that CPOM traversed a mean distance of 10 to 30 m before entrainment, and therefore storage and processing would occur within the reach with possible export of finer particles (Cummins et al. 1981). The distribution of FPOM was similar during all seasons with the shifts occurring in the CPOM range where lowest levels occurred during the spring, possibly the result of flooding, since corresponding increases in FPOM did not occur (Figures 19, 20). Spring allochthonous CPOM inputs may have accounted for the observed higher summer values (Figure 20). The Upper 43rd particle-size specific distribution was more constant than the first order sites with algal inputs probably occurring during the summer as indicated by increases in the FPOM pool (Figure 20). Litke (1978) observed an increase in the detrital pool 81 Table 11. CPOM and FPOM detrital standing crops and CPOM/FPOM ratios from selected sites in Augusta Creek, Kalamazoo County, Michigan (1974-1975). Month Site Oct. Jan. May July Smith Smith Smith Smith Oct. Jan. April July Oct. Jan. May July Nov. Jan. May July Nov. Feb. May July B B B B CPOM gm" 2 X S.B. C.V.% 406.7 327.4 279.4 274.5 322.0 30.7 19.0 0.18 0.34 0.13 0.16 0.20 0.02 46.3 X S.G. C.V.% 172.0 42.6 27.5 83.0 81.3 32.4 79.8 419.9 134.4 98.4 228.7 220.4 72.0 65.3 0.41 0.32 0.28 0.36 0.34 0.02 16.2 X S.E. C.V.% 50.3 42.4 53.6 22.5 42.2 7.0 33.1 168.0 192.6 123.9 281.2 191.4 33.1 34.6 0.30 0.22 0.43 0.08 0.23 0.07 56.9 X S.G. C.V.% 100.5 159.4 77.8 42.9 95.2 24.5 51.4 287.7 233.9 194.9 258.2 243.7 19.6 16.1 0.35 0.68 0.40 0.17 0.40 0.11 52.8 X S.G. C.V.% 29.5 29.5 46.6 18.3 30.5 5.9 38.7 185.3 259.5 296.7 267.4 252.2 23.7 18.8 0.16 0.11 0.16 0.06 0.12 0.02 39.1 43rd 43rd 43rd 43rd Nagel Nagel Nagel Nagel Kellogg Kellogg Kellogg Kellogg CPOM/FPOM 73.0 112.3 37.2 46.4 67.2 16.8 50.1 Avenue Avenue Avenue Avenue Upper Upper Upper Upper FPOM gm-2 Forest Forest Forest Forest 82 due to Cladophora inputs in a fourth order riffle during the summers. Such an increase was also observed at the Nagel Site, where CPOM levels were highest during the fall-winter period. The Kellogg Forest had little change in detrital particle-size distribution over the year, which indicated the influence of mainly non-filamentous autochthonous production at this site. The CPOM/FPOM ratios ranged from 0.06 during the summer at Kellogg Forest to 0.68 in the winter at the Nagel Site. The lowest ratios, indicating the autochthonous energy base, were at the higher order sites during the summer and at6 Avenue and Smith during the spring at the time of increased light and GCP "maxima". The highest ratios varied with site, but correlated with depressed P / R ratios (Figure 17; Table 11). The winter-early spring dependence on CPOM at the autotrophlc sites correlated with low levels of GCP and NCP, while the .autumn-winter inputs of CPOM and subsequent processing within the riffles coincided with the high CPOM/FPOM at the heterotrophic sites (Figures 14, 15; Table 11). Therefore, the P/R and CPOM/FPOM ratios can be used in conjunction to elucidate the autotrophic-heterotrophic relationships of stream riffles. The annual average CPOM/FPOM ratios were highest in the shaded first order reaches (0.2 Smith and 0.34 B Avenue) and lower in the third order reach (0.12 Kellogg Forest). This pattern was in agreement with the continuum concept and reflected a greater dependence on autotrophlc production as stream order increased (Vannote et al. 1980). However, the cleared Nagel section had the greatest average CPOM/FPOM ration (0.4) and the highest average CPOM standing crop. When the S.E.'s are considered (Table 11), the Nagel 83 CPOM standing crop was comparable to those from Smith and B Avenue. Based on their overall productivity differences, the Nagel CPOM resulted from inputs of filamentous algae and aquatic macrophytes and thus, had been autochthonously produced as documented by Minshall (1978) in desert streams. Although the actual origins of allochthonous CPOM were not demonstrated, Mahan (1980) ranked Nagel the lowest in CPOM inputs of the four Augusta Creek sites studied, which supports the autotrophlc input hypothesis. The meadow reach, Upper 43rd, average 0.23 CPOM/FPOM ratio, which was not significantly different from that at the Smith Site. Therefore, the generalization of decreasing CPOM/FPOM ratio with increasing order did not hold for open reaches of first through third order streams. Detrital (AFDW) Community Metabolism Little relationship between detrital AFDW and CR has been noted in the literature (Wetzel 1963, Edwards and Rolley 1965, Hargrave 1969b, Pennak and Lavelle 1979). Odum (1956) reported CR to be dependent on the concentrations of organic matter in the sediment, Edberg and Hofsten (1973) found the nature and quality of detritus had a decided effect on oxygen uptake in soft sediments, and Gothberg and Karlstrom (1975) found CR to be dependent on the amount of detritus present in riffle sections of the Ricklean River, Sweden. These findings contrasted to those of Hargrave (1969b) that organic content and composition appeared to have minor significance in oxygen consumption of littoral sediments. In a later study comparing mud, sand, and detritus in short term experiments Hargrave (1972) found detritus consumed three times more oxygen that s a n d , and that log 84 oxygen uptake was inversly related to organic content of sediment. Intuitively, CR would be linked most closely to detrital activity, and most studies have emphasized this relationship. Gdberg and Hofsten (1973) indicated that CR of algal detritus was much higher than allochthonous forms, which may account for increases of CR during periods of algal growth and senescence. Detritus (especially CPOM) can also be a substrate for algal colonization and a relationship may exist between AFDW and productivity. Community metabolism estimates expressed on an AFDW basis may reflect changes in the composition of the detrital standing crop; Increased productivity g“ l m a y indicate algal colonization of detrital particles or contribution to the detrital pool through sloughing from inorganic substrates, and increasing CR rates during spring or fall may indicate more labile algal contributions to the detrital Community metabolism expressed pool. on a biomass basis (g~^AFDW; Appendix A; Table 12) indicated the same trends (Figures 21, 22) as measured on an areal basis with the exception of CR (Figures 16, 18; Table 9). 14, 15, Smith and B Avenue were reversed in CR ranking by expression on a weight basis, but there was no significant difference between these values. Since the mean total standing crop estimates at all sites, excepting Smith, were so close one would not anticipate large changes in seasonal trends (Table 11). Periods of maximum and minimum values on an areal or biomass basis for these parameters GCP, CR, NDM) were generally similar at all sites. (NCP, Values at the Smith and B Avenue sites were most stable with only small annual changes. CR varied most of any parameter: summer and spring maxium rates occurred when expressed on an areal basis, while on a biomass 85 Table 12. Annual average estimates of detrital community metabolism (mg O 2 g ” ^AFDW) for selected sites of Augusta Creek, Kalamazoo County, Michigan (1974-1975). Site Smith B Avenue Upper 43rd Nagel Kellogg Forest NCP CR GCP NDM X S.E. C.V.% -0.7 1.8 0.3 -1.5 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 X S.E. C.V.% 0.0 0.0 0.0 2.4 1.2 -1.3 0.0 0.0 0.0 0.0 0.0 0.0 X S.E. C.V.% 5.8 0.0 8.0 0.0 55.0 39.5 9.8 3.2 64.8 0.0 0.0 6.7 7.2 10.6 0.0 0.0 0.0 62.0 66.4 2.3 X S.E. C.V.% 3.4 47.1 X S.E. C.V.% 3.8 2.5 5.0 2.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 92.4 86 SMITH I 3e 10 o u. < 0> B 0 o E Ix NAGEL 20 10 KELLOGG FOR E S T 10 J F M A M J J A S 0 N D OsNCP • CR Figure 21. Estimates of NCP and CR on a weight basis from selected Augusta Creek riffle sections, Kalamazoo County, Michigan (1973-1975). 87 SMITH 10 U P P E R 4 3 R0 cl 30 O NAGEL (M o 20 K ELLOGG F OR E S T J F M A M J J A 0 N D MONTH 0 ‘ GC P • =N D M Figure 22. Estimates of GCP and NDM on a weight basis from selected Augusta Creek riffle sections, Kalamazoo County, Michigan (1973-1975). 88 basis the maximum was in the autumn (Appendix A; Figures 16, 21). At the Upper 43rd and Nagel Sites these increases in CR ma y have been the result of algal inputs to the detrital pool (Edberg and Hofsten 1973). Calow (1975) reported epilithic detritus was higher in organic matter and protein in comparison to adjacent detritus which was higher in lignin and cellulose. Thus, when periphyton assemblages slough into the detrital pool, higher rates of CR would be expected, from the addition of a more labile substrate. It should be noted that in all cases values were low on a biomass basis and there was little difference in seasonal ranges (Table 12; Figures 21, 22). The autotrophic Nagel Site had high levels of detrital standing crop and also had the highest values of most parameters on a biomass basis (Table 12). The high state of activity g“ * detritus at the Nagel site was also characteristic at Upper 43rd. Based on average detrital standing crop, Smith was highest followed by Nagel, B Avenue, Kellogg Forest, and Upper 43rd. If metabolic parameters were directly correlated with total detritus the highest activity would have been at Smith Site, which had the lowest activity followed by B Avenue, Kellogg Forest, Upper 43rd and Nagel. Therefore, other factors such as light temperature or epllithon development and composition must be mediating metabolic activity. Epilithon Epilithon, is here defined as that organic assemblage of algae, bacteria, fungi, protozoans, detritus, and even small macroinvertebrates which colonize this mass, attached to inorganic substrates. This definition is often lumped with the category of 89 periphyton or Aufwuchs. Round (1965) has long campaigned to characterize the assembladge based on substrate colonized (i.e. epilithon - rocks and s tones , epipsammon - s a n d , epipelon - m u d , epiphyton - plants). The term epilithon has been used by Marker (1976a) to describe the biomass of benthic algae on rocks and Calow (1975) has studied epllithic detritus. Epilithon biomass values (Appendix C) are based on the 550 AFDW of particle-sized inorganic substrates. Nelson and Scott (1962) ashed, rewetted and redried at 63C to attempt rehydration of clay particles. They estimated error due to loss of hydration at 5%. Losses of this type in Augusta Creek sediments to 75 p m in size would be small and no corrections were made. The method developed showed comparative differences between sites in Augusta Creek epilithon development. Sediments throughout the Augusta Creek drainage were similar at all sites; they consisted of granite, basalt and limestone. However, visual differences of sediment surfaces were obvious between autotrophlc and heterotrophic sites. Sediments from Nagel and Upper 43rd had obvious travertine deposits, Kellogg Forest intermediate in travertine build up, and Smith and B Avenue were free of visible carbonate deposits. equilibrium, In hardwater streams photosynthesis, shifting the can be responsible for deposition of CaC03 on sediments or aquatic macrophytes and subsequent travertine development 1970, Wetzel 1975a). (Hynes Photosynthetic activity was largely responsible for both visual and biomass differences between sites and epilithon development was a more reliable indicator of autotrophic-heterotrophic relationships of riffle sections than was detrital standing crop. 90 Sites ranked according to Increasing annual average epilithon development were: Smith, B Avenue, Kellogg Forest, Nagel and Upper 43rd (Table 9; Figure 23). Using overlap of standard errors as a criterion, Nagel and Upper 43rd were essentially the same, Kellogg Forest intermediate, and Smith significantly lower than all other sites in epilithon development. These trends reflected the same pattern as NCP and GCP in Augusta Creek (Table 9). Epilithon estimates ranged from 1480 g m “ 2 at Smith to 5038 g m~2 at Nagel Site during the spring. The maximum at the Nagel Site undoubtedly resulted from the diatom bloom which carpeted the substrate, while the low at Smith Site was attributed to spring scouring effects. These epilithon values from the alkaline Augusta Creek were higher than those obtained using the same method in soft water, turbid riffle sections of the Buttahatchie River Mississippi (Miller and King 1981). Their total AFDW estimates of sediments, characteristic of epilithon weights (Table 9), ranged from 260 to 363 g m” 2 in August. Development of the autotrophlc epilithon component in this system was limited by turbidity, thus, these levels characterize the contribution of the detrital component to the biomass. Epilithon values from six replicate chambers placed in a second order riffle of the west branch of the Clark's Fork of the Yellowstone River, Montana, where primary productivity levels were very low, ranged from 999 to 1892 g ra“ 2. The average (1338 g m” 2) was similar to that of the heterotrophic Smith Site of Augusta Creek (1763 g m “ 2) as were the P/R ratios of approximately 0.3 (Table 9). The first order detrital based systems of Augusta Creek ranged from 1480 to 2811 g m~2 epilithon. Both first order sites indicated 91 SMITH 2 0 2 0 4 U P P E R 4 3 RD 2 0 NAGEL 4 2 0 4 2 KELLOGG f o r e s t 0 SP Figure 23. SU SEASON Estimates of eiplithon development, obtained on a seasonal basis from selected riffle sites of Augusta Creek, Kalamazoo County, Michigan (1974-1975). 92 less variance In epilithon estimates over the year with the C.V.% averaging only 18.3% at Smith and 16.6% at B Avenue in contrast to the overall average of 28.7% (Table 9). This consistency, indicating a balance within the riffle sections, was typical of most parameters at these sites. The highest values of epilithon at Smith Site were during autumn (Figure 23) at the time of significant detrital inputs. The nature and consistency of epilithic detritus has been investigated by Madsen (1972). Using SEM techniques, detritus was the major component, bacteria and fungi were prevalent, with few algae evident. This material may be a rich source of organic matter (protein) to consumer organisms (Calow 1975). Colonization of stones by heterotrophs thus leads to organic accumulation and further entrapment of detritus all of which would be a component of the epilithon as determined by ashing. These levels at Smith Site were significantly different from the autotrophlc based sites (Figure 23; Table 9). Helan et al. (1973) ashed inorganic substrates from mountain streams in Czechoslavakia and found highest amounts of epilithon in the upper reaches during April and June, similar to patterns observed at B Avenue, Augusta Creek (Figure 23). The Kellogg Forest Site, intermediate in epilithon development, had little change in quantity of epilithon, and was consistently higher in values than the first order sites (Table 9: Figure 23). The open sites illustrated more significant seasonal changes, with the Upper 43rd Site maximum attained during the autumn after the autotrophlc build up from a spring low (Figure 23). The Nagel site exhibited spring and autumn increases (diatom blooms) much like that reported for the middle 93 sections by Helan et al. (1973). The lower summer values at Nagel (Figure 23) reflected shading effects of extensive aquatic macrophyte growth during mid to late summer. On a particle-size specific basis (Figure 24), changes occurred m a i n l y in the 16 and 4 m m size fractions. McConnell and Sigler (1959) found larger substrates (12 cm and greater) to have more chlorophyll a content than smaller s e d i men t s , with less than 6% of the chlorophyll a concentration in sediments smaller than 2.5 cm in diameter. English streams Marker In (1976a) found 85% of the algal biomass on substrates greater than 6 cm. This pattern of highest autotrophlc colonization on larger substrates was consistent with that in Augusta Creek. Based on productivity levels and visual observations, this assemblage at the autotrophlc sites was due to algal development. Since the epilithon standing crops in Augusta Creek riffles were so high compared to the detrital standing crop, total organic weight of the sediments followed the trends of the epilithon community. Detritus averaged 12% of the total particulate organic matter (TPOM), w i t h the percentages ranging from an average of 18% at Smith to 7.5% at Upper 43rd (Table 13). Detrital percentages indicated a greater importance of detritus at Smith and the autotrophlc nature of the Upper 43rd and Nagel riffles. This relationship was consistent over the annual period at all sites, although the magnitude of epilithon and detritus estimates varied. The heterotrophic riffle of the west branch of the C l a r k ’s Fork of the Yellowstone River, Montana averaged 4.2% non-attached detritus, which indicated the importance of non-photosynthetic epilithon in this system. Epilithon/detritus ratios (E/D) establish an index of the 94 E PI LI T H O M Bm -2* r r r r SM ITH 16 4 1 .250 .075 B 16 E E 4 1 Ut2&0 £.075 V) AVENUE r r r r UPPER 4 3 RD NAG E L KELLOGG FO RES T 16 .250 .075 rr r r W * ■ SP = 400 g Figure 24. su SEA SO N Seasonal distribution of particle-sized epilithon from selected riffle sections of Augusta Creek, Kalamazoo County, Mich i g a n (1974-1975). Table 13. Percentage of detritus and epilithon and epilithon/detritus (E/D) ratios of sediments from selected sites of Augusta Creek, Kalamazoo County, Michigan. WINTER SPRING Site % E % D % E Smith 79 21 85 B Avenue 92 8 Upper 43rd 91 Nagel Kellogg Forest % D SUMMER MEAN ANNUAL FALL ANNU A L % E % D % E % D % E % D range E/D 5 82 18 82 18 82 18 3 . 7 - 4 .8 4.5 94 5 89 11 79 21 89 11 3.46-16.5 7.9 9 91 9 92 8 96 4 93 8 10.2-15.0 14.8 91 9 94 6 88 12 89 11 91 10 7.5-15.4 12.0 90 10 82 18 92 8 94 6 90 11 4.6-15.8 9.5 X E/D 96 relative importance of the two components within riffles. Based on seasonal data, the average E/D for Smith Site was 4.5, B Avenue 7.9, Upper 43rd 14.8, Nagel 12.0, and Kellogg Forest 9.5 (Table 13). The closer the ratio is to one (E « D ) , the higher the dependence of the riffle section on non-attached detritus versus an epilithon assemblage. Conversely, the larger the ratio, the greater the dependence on the epilithon organic assemblage. The Smith and B Avenue Sites were more dependent on non-attached detritus than the other reaches. Annual average E/D ratios from Augusta Creek sites followed the patterns of average NCP and GCP (Tables 9, 13), which indicated an algal dominance in the epilithon at the autotrophlc sites. The summer data from the west branch of the Clark's Fork of the Yellowstone River, Montana averaged 23.6 E/D; this value, higher than the autotrophlc Augusta Creek sites, indicated an epilithon based community, although detrital, since levels of NCP and P/ R were low. Rocks from this riffle were slippery to the touch during the study period, but not high in algal content. Therefore, the magnitude of this ratio Indicates the importance of epilithon versus non-attached detritus within a system, but not the autotrophic-heterotrophic balance within the assemblage. Seasonal patterns of E/D ratio exhibited no constant pattern except at the Smith Site (Figure 25). The values of approximately five measured at the Smith Site were indicative of consistent dependence on non-attached detritus as an energy base (heterotrophic). The B Avenue Site, indicating a more complex pattern of E/D, was also l ow in E/D average, similar to Smith Site. A spring Increase in epilithon development concomitant with the spring diatom bloom in 97 20 20 SMITH 10 10 UPPER 43R D 20 10 NAGEL 20 20 10 10 KELLOGG FO R E S T SEASON Figure 25. Epilithon/Detritus ratios over an annual period for selected riffle sites of Augusta Creek, Kalamazoo County, Michigan (1974-1975). 98 Augusta Creek was reflected by an Increase In E/D (Figure 25) at the B Avenue Site. The P/R ratio and values of NCP Increased at this time as well, which indicated the increase in E/D was of algal origin (Figures 15, 17, 25). The autotrophlc sites had an inverse relationship between GCP, NCP, and/or P /R with the E/D ratio. Autotrophlc sites remained productive, but as epilithon increased, rates of primary productivity decreased or remained stable at all three sites. This pattern follows the generally accepted trend of decreased rates of primary productivity with periphyton assemblage build-up caused by self-shading and competition for light, carbon dioxide, and nutrients (Goldman 1964, Hynes 1970, Wetzel 1975a). When one observes riffles within the deciduous biome it is generally the inorganic sediments which predominate and are the major substrates available for colonization, not detritus. Although the scientific literature focuses on water percolation through leaf packs in riffles, a major source of POM might well be the epilithon in many stream riffles. Winterborn et al. (1981) indicated that in New Zealand streams the organic component from inorganic substrates was important to the support of macroinvertebrates and suggested attention be refocused on the organic layer of stones. Therefore, the E/D ratio used with the P/R ratio should be a useful relationship to monitor in ecological investigations involving stream structure and function. Factors Affecting Rates of Community Metabolism Although reseachers engaged in monitoring community metabolism may disagree, Lieth (1975) stated that measurement of primary productivity is not in itself a sufficient goal, but should be used as a tool for characterization and interpretation of communities and their relationships to the environment. One needs only to scan the periphyton primary productivity literature to note investigations of regulatory factors on rates of community metabolism and contributions of the autotrophs to community structure and function. Most work has been completed in marine and freshwater lentic systems and includes investigations relating physical and chemical parameters to community metabolism of benthic communities from tidal flats (Pamatmat 1968, Gannlng and Wulff 1970, Gargas 1970, Gallagher and Daiber 1973, Littler 1973, Cadee and Hegeman 1974), freshwater littoral zones (Felfoldy 1961, Wetzel 1963, Hargrave 1969a, Hunding 1971, Hunding and Hargrave 1973, Schindler et al. 1973), experimental systems (Odum and Hoskin 1957, Felfoldy 1961, Whitford and Schumacher 1961, 1964, Beyers 1962, Mclntire et al. 1964, Mclntire and Phinney 1965, Kevern and Ball 1965, Lester et al. 1974, Edberg and Hofsten 1973, Pfeifer and McDiffett 1975) and in lotic systems (McConnell and Sigler 1959, Kobayasi 1961, Cushing 1967, Stockner 1968, Brehmer et al. 1969, Hall 1972, Cole 1973, Ertl and Tomajka 1973, Profet and Ransom 1974, Marker 1976b, Pennak and Lavelle 1979, Sumner and Fisher 1979, Gregory 1980). Upstream-downstream techniques were mainly used prior to the collection of data for the present study (1973-1975). The most frequently examined parameters were light and temperature, but included chlorophyll content or AFDW of periphyton, current, organic content of sediments, nutrients, and recently grazing (Hunter 1980, Gregory 1980, Kesler 1981). In the present study, parameters examined in relation to NCP, GCP, CR, and NDM included stream order, temperature, light, and AFDW 100 of the sediments (detrital, epilithon and total). Cusing (1967) studied many parameters over an annual period from periphyton communitites developed on artificial substrates, but none of the above studies reported in situ data for as extensive a set of parameters over an extended period. Regression analyses were completed for each parameter listed above against each dependent variable NDM) on an annual average basis, as well as by site. (NCP, GCP, CR, Individual half-hour rates of NCP and CR (parameters measured directly in the field) were matched with corresponding light and temperature intervals and run individually and together in multiple regression. The independent variables are discussed separately, but a summary of the regressions is available in Appendix G. Stream Order The order system, as defined by Strahler (1957) classifies streams on a physical basis and is convenient for comparison of data particularly within a given watershed. Hynes (1970) suggested heavier usage of this format for characterization of streams. have begun to relate results on an order basis. Investigators Cummins (1974, 1975b, 1977) and Vannote et al. (1980) related various components of stream community structure and function to an order scale. M a h a n (1980) found a relationship between amounts of CPOM input to woodland streams, Naiman and Sedell (1979a) investigated benthic organic matter on an order basis and Seyfer and Wilhm (1977) found relationships between periphyton species composition, diversity, and biomass to be a function of stream order. Regressions with stream order as the independent variable 101 indicated a significant (P < 0.01) positive relationship for all parameters with n = '100 for each case (Appendix E; Table 14). Multiple correlation coefficients (R) ranged from 0.52 for CR to 0.74 for NCP, and F ratios (test for signlficances l; higher, more significant) were high especially for GCP and NCP (Table 14; Appendix E). The correlation between stream order and each dependent variable was always highest, which indicates the importance of geomorphology on the functional aspects of stream communities. It is intuitive that as streams widen they receive more light and higher rates of primary productivity would result unless factors such as turbidity, or chemical/physical perturbations were involved. NCP and GCP had the highest F ratios (Table 14), which would be most directly related to light. Rates of CR, although significant, were least affected by stream order, and illustrated a lesser dependence on location (i.e. light availability) than the photosynthetic processes and a heterotroph dominated CR. NDM, influenced by rates of CR, had a lower F ratio, but still indicated a strong relationship with order (F = 74; Table 14). Light Various authors have eluded to the importance of light on aquatic primary productivity in the field (Butcher et al. 1930, Calvert 1933, McConnell and Sigler 1959, Profet and Ransom 1974, Sumner and Fisher 1979, Hornick et al. 1981), but few studies have carefully examined the relationship (Kobayasi 1961, Stockner 1968, Hunding 1971, Caded and Hegeman 1974). Marker (1976b), using Tygon cloth to shade chambers (9,000 lux 102 Table 14. Summary of regression data for various dependent variables and the independent variable stream order for all sites Augusta Creek, Kalamazoo County, Michigan. p*** Dependent Variable N* R** F Ratio X GCP 101 0.72 108.1 <0.001 X NCP 103 0.74 121.9 <0.001 X CR 104 0.52 37.5 <0.001 X NDM 101 0.66 74.2 <0.001 * Number of values ** = Multiple Correlation Coefficient *** = Level of Significance 103 versus 27,000 lux lighted) illustrated a 30% decrease in primary productivity. Gregory (1980) artificially lighted a section of a wooded stream (10,000 lux) and significantly increased productivity of a riffle section. The effects of clear cutting and channelization on photosynthesis have been shown by Gelroth and Marzolf Gregory (1980). (1978) and Pennak and Lavelle (1979), studying in situ NCP in mountain stream riffles, found no correlation between NCP and light intensity, but the results were probably Influenced by very low periphyton standing crops. Regression analyses from Augusta Creek communities indicated light was the second most important parameter affecting individual NCP and average GCP, NCP, and NDM; outranked only by stream order (Table 15; Appendix E). F ratios and R values were high and significant in all cases ( P < 0.05; Table 15). Average data (n = ~ 100) with correlation coefficients ranging from 0.52 to 0.57 and F ratios ranging from 36 to 46, indicated the importance of light on levels of GCP, NCP, and NDM within the Augusta Creek drainage, GCP had the highest R (0.57) and F ratio (46.0), but all were similar (Appendix E; Table 15). When individual measures of NCP were used in regression analyses (n - 666 and R - 0.6 with a high level of significance; Table 15) and when multiple regression with temperature and light was completed (Table 16) R = 0.6 with light R value = 0.597 and temperature 0.005. The Augusta Creek correlation coefficient for light and NCP, although lower than those of Cushing (1967) and Stockner (1968) for lotic periphyton communities (R=0.8), included values from heterotrophlc as well as autotrophic sites. Pamatmat (1968) and Gargas (1970) cited light as the most important factor 104 Table 15. Summary of regression analyses involving various dependent variables and the Independent variable light at five sites from Augusta Creek, Kalamazoo County, Michigan. p*** Dependent Variable N* R** Individual - NCP 666 0.60 1089.7 <0.001 X GCP 100 0.57 46.0 <0.001 X NCP 101 0.52 36.0 <0.001 X NDM 100 0.53 39.2 < 0.001 12 12 12 0.26 0.27 0.23 0.695 0.786 0.573 0.424 0.396 0.467 33 33 33 0.07 0.16 0.23 0.162 0.812 0.573 0.690 0.374 0.467 10 10 10 0.48 0.41 0.61 2.444 1.569 4.713 0.157 0.246 0.062 34 35 34 0.38 0.24 0.26 5.496 2.035 2.373 0.025 0.163 0.133 11 11 11 0.53 0.42 0.64 3.533 1.984 6.146 0.093 0.193 0.035 F Ratio SMITH X GCP X NCP X NDM B AVENUE X GCP X NCP X NDM UPPER 43RD X GCP X NCP X NDM NAGEL X GCP X NCP X NDM KELLOGG FOREST X GCP X NCP X NDM * N * number of va l u e s , ** R = Multiple Correlation Coefficient *** P - Level of Significance 105 Table 16. Summary of multiple regressions involving the effects of temperature and light on individual rates of NCP and CR from all sites of Augusta Creek, Kalamazoo County, Michigan. Dependent Variable p*** N* R** F Ratio T + L 1957 0.598 544.6 <0.001 L only 1957 0.597 T only 1957 0.005 T + L 1189 0.523 447.1 < 0.001 L only 1189 0.241 T only 1189 0.425 Individual NCP Individual CR N = number of values ** R = Multiple Correlation Coefficient *** P = level of Significance 106 influencing community metabolism of marine microbenthos (F ratios =• 72-98), as did Littler (1973) in coral reef communities, and Hunding and Hargrave (1973) in freshwater littoral zones. The sites trend in F ratios reflected productive capacity. Lower values were from Smith and B Avenue, while the Upper 43rd, Nagel and Kellogg Forest F ratios were greater than one, however, on a site basis values were not significant in most casts (Table 15). At autotrophic based sites the effects of light were greater on GCP and NDM than NCP (Table 15). Values of GCP and NDM were calculated using estimates of CR; one is a positive influence negative (GCP) while the other is (NDM) and both increased, which indicates some unmeasured parameter such as grazing losses may have been involved. (1981) Kesler reported Chlronomus Scutellaria may consume 3 to 70% of the NCP, while Hunt (1980) and Gregory (1980) found that snails could effectively reduce periphyton communities. organisms, Populations of grazer including H y d r o p t l l a , N e o p h y l a x , G l o s s o s o m a , Helicopsyche (Trichopterarcaddisflies), Stenonema (Ephemeroptera:mayflies), Psephenus (Coleoptera:beetles) and Ferrlssia (Gastropoda:snails) were common, especially at the Nagel Site and observations of grazing rates by D. Hart (personal communication, Kellogg Biological Station) indicated active cropping of periphyton at the Nagel Site. The relationship between light intensity and rates of photosynthesis is not linear over a large range of light values. Light saturation and photoinhibition often occur at higher light intensities in phytoplankton communities (Ryther 1956a, Verduin 1956, Pamatmat 1968, Marker 1976b, Ganning and Wulff 1970. Light saturation occurred at approximately 20,000 lux for combined sites of Augusta 107 Creek (Appendix F; Figure F - 1). ranges were very similar: Except for Smith Site, saturation B Avenue (n=677) 20,000 lux, Upper 43rd (n=220) 24,000 lux, Nagel (n=571) 15,000 to 20,000 lux, and Kellogg Forest (n=242) 18,000 to 20,000 lux (Figures F 1-6). Rates of NCP, and likely algal standing crops, were very low at the Smith Site and few measurements of NCP were made above 20,000 lux (Figure F - 2), therefore, resolution of light saturated photosynthesis was not possible. These saturation levels are within the range reported for marine phytoplankton (5,000 to 30,000 lux; Ryther 1956a), and lentic benthic algae (10,000 to 12,000 lux; Hunding 1971, Lester et al. 1974, Colljn and Buurt 1975), and natural and artificial streams (4,000 to 62,000 lux; Kobayasi 1961, Mclntire et al. 1964). Photoinhibition was not observed at any of the sites (Figures F 2- 6 ), which was not surprising, as it has not been reported in lentic periphyton communities (Pamatmat 1968, Ganning and Wulff 1970, Hunding 1971, Gallagher and Daiber 1973, Littler 1973, Caded and Hegeman 1974, Colijn and Buurt 1975). In artificial stream developed periphyton communities, Mclntire et al. (1964) and Mclntire and Phinney (1965) found no photoinhibition and Marker (1976b) noted no photoinhibition from an in situ study. The large number of in situ cases examined from autotrophic and heterotrophic sites in the Augusta Creek watershed reinforced these noted examples. The effects of temperature on light saturated photosynthesis are illustrated in Appendix F (Figures F 7-11). The most linear of these plots (0 - 5 C) indicated light saturation at 18,000 lux (Figure F 7, n=414). At temperature intervals 5 to 10 C, 10 to 15 C, 15 to 20 C, and 20 to 25 C light saturation occurred between 20,000 to 25,000 108 lux (Figures F 8-11). Kobayasi (1961) found seasonal shifts in light saturation from 8000 lux in a winter diatom community to 15,000 lux in the summer Cladophora dominated community. As indicated by Ryther (1956a) light saturation levels vary for various algal taxa, and the shift observed by Kobayasi (1961) and the slight increase at 15 - 20 C (Figure F-10) at Augusta Creek sites may have been due to changes in species composition rather than temperature. temperature on photosynthetic activity, The influence of studied by Rabinovich (1951) and Jprgensen and Steemann Nielsen (1965, 1968), discussed in Mclntire et al. (1964) and Hunding (1971), indicated the rapid increase portion of the activity curve to be due to photochemical processes independent of temperature, while the saturated portion of the curve fluctuated depending on temperature effects on the photosynthetic enzymes. Although rates of NCP increased with temperature, similar levels of light saturation were indicated over the temperature range naturally encountered by Augusta Creek communities (0 - 25 C) and illustrated the complexity of photosynthetic responses to more than one variable. Light was the primary factor regulating primary productivity in woodland streams and based on light-saturated photosynthesis levels and average daily light of 12,000 to 13,000 lux, amounts of NCP at Smith and B Avenue would be limited. Reduced levels of NCP were documented by in situ measurements at these two sites compared to the open and shaded larger order study reaches (Figure 15). Temperature The effects of temperature on organisms and metabolic processes have been extensively investigated, and Qjg values have been calculated 109 for process of organisms from bacteria to mammals. The relationship between temperature and community metabolism in streams, however, has not received much attention. In aquatic systems the effects of temperature on CR was investigated in marine tidal flats (Pamatmat 1968, Gallagher and Daiber 1973, Cadee' and Hegeman 1974), and freshwater littoral zones (Hargrave 1969a, 1969b; Edberg and Hofsten 1973) with all investigators reporting temperature to be an important controlling factor. The effects of temperature on community productivity have been studied, but mainly in lentic systems where a positive correlation has been indicated, but of lesser importance than the effects of light (Gargas 1970, Hunding 1971, Adams and Stone 1973, Ertl and Tomajka 1973, Gallagher and Daiber 1973, Caded 1974, Colijn and Buurt 1975). and Hegeman The average data (n=100) indicated a positive relationship between temperature and all parameters with R values ranging from 0.64 for CR to 0.33 for NCP and F ratios of 12 to 70 (Appendix E; Table 17). These values were generally lower than the values for light, but still significant at the P=0.01 level (Tables 15, 17). While CR had the lowest correlation with light, it was most strongly correlated wi t h temperature. Hargrave (1969b), Ertl and Tomajka (1973), and Cadee and Hegeman (1974) reported positive correlations between levels of GPP and temperature in lentic systems. temperature and GPP from intertidal flats The R of 0.5 for of the Wadden Sea was similar to the 0.41 R for GCP and temperature from Augusta Creek sites (Table 17). Individual rates of NCP (n=1957) indicated a significant correlation with temperature (R = 0.18; F ratio = 67.1). However, 110 Table 17* Summary of regressions Involving the effects of temperature on NCP, GCP, CR, and NDM from various sites of Augusta Creek, Kalamazoo County, Michigan. Dependent Variable N* R** F Ratio p*** Individual - NCP Individual - CR 1957 1189 0.182 0.523 67.1 447.1 <0.001 <0.001 GCP NCP CR NDM 100 102 103 100 0.414 0.328 0.640 0.339 20.3 12.0 70.0 12.7 <0.001 0.001 <0.001 0.001 GCP NCP CR NDM 12 12 12 12 0.022 0.0003 0.013 0.133 0.005 0.000 0.002 0.180 0.946 0.999 0.968 0.680 34 35 34 34 0.600 0.047 0.785 0.368 17.8 0.07 51.2 5.00 <0.001 0.790 <0.001 0.032 GCP NCP CR NDM 10 10 11 10 0.677 0.547 0.898 0.519 6.7 3.4 37.6 2.9 0.032 0.102 <0.001 0.125 GCP NCP CR NDM 34 35 34 34 0.529 0.377 0.742 0.263 12.4 5.5 39.3 2.3 0.001 0.026 < 0.001 0.133 10 10 12 0.660 0.361 0.765 6.2 1.2 14.1 0.038 0.305 0.004 X X X X SMITH X X X X B AVENUE X X X X GCP NCP CR NDM UPPER 43RD X X X X NAGEL X X X X KELLOGG FOREST X GCP X NCP X CR N = number of values ** R ■ Multiple Correlation Coefficient *** P ° Level of Significance____________ Ill this was much less than the regression with light (R ■ 0.06; F ratio = 4089.7). Individual rates of CR (n=1189) had an R value of 0.52 and F ratio of 447,1, which indicated a highly significant relationship (Table 17). When run with light in multiple regression, temperature had an R > 0.43 in contrast to R ■ 0.24 for light (Table 16). Together (n«1968), or on a site basis, there was no evident relationship between NCP and temperature (Figures G 7-12). Considering the 1189 points in Figure G - 1, rates of CR generally increased linearly with increasing temperature, especially if data from Smith and B Avenue Sites are omitted. Both of these sites showed constant CR regardless of temperature (Figures G-2, G-3). The lack of correlation between temperature and CR at Smith Site was evident from the regression analyses (Table 17). Assuming that these communities are heterotrophic, one would expect CR to increase with temperature as reported for bacteria (Brock 1966). The consistency noted at the first order sites was related to the comparatively low epilithon standing crops. Beyers (1962) recorded that in experimental microcosms temperature affected CR until a balanced ecosystem had developed. He related this to the multlclpllcity of metabolic pathways at work within the community, which easily adapted to changing conditions. Therefore, the consistency may be related to establishment of a dynamic equilibrium as rates of CR varied little over the seasons. At the open Upper 43rd and Nagel sites with well developed epilithic communities, a direct relationship existed with temperature (Figures G-4, G-5). development, Levels of CR correlated with epilithon ranking from low to highest rates and development: 112 Smith, B Avenue, Kellogg Forest and Nagel (Kellogg Forest = Nagel) (Table 9). Although temperature was important in controlling rates of CR, a community must be present to respire, and at Smith Site low CR indicated low microbial development. O n a site basis a significant relationship with temperature was not evident for any parameter measured at Smith Site, while at B Avenue CR, GCP and NDM were correlated with temperature Table 17). (Appendix E, Cooler temperatures probably influenced algal development at these sites (Smith x 9.5 C, B Avenue x 8.7 C ) , but temperature ranges (0 - 25 C) included periods in which diatom growth abounded (Table 10). cover, The higher temperatures occurred during maximum canopy therefore, complicating interpretation of development patterns. Diatoms were visually present at B Avenue, and GCP and NDM were correlated with temperature at this site. At the Nagel, Upper 43rd, and Kellogg Forest sites, CR was again the most obviously correlated parameter with temperature; GCP was the only other significantly correlated parameter (P = 0.05; Appendix E; Table 17). NCP levels were more positively influenced by temperature at these open sites (Figures G 10 - 12), as indicated by increased R values and F ratios (Table 17). This relationship indicated that temperature may have been limiting epilithon development at the shaded first order sites when light was available for increased levels of primary productivity. Adams and Stone (1973) reported seasonal differences in NCP of Cladophora to be closely related to temperatures. NDM, The importance of temperature on rates of CR, GCP, and possibly through CR mediation on GCP and NDM, or some unknown influence such as community composition and species temperature 113 optima, or total epilithon development and stability was evident at sites of Augusta Creek. Detrltal-Ash-Free Dry Weight The question of the effects of detrltal standing crops on CR of communities has been addressed In lentic and lotic systems (Nelson and Scott 1962, Hargrave 1969b, Edberg and Hofsten 1973, Gtfthberg and KarlstrOm 1975). Studies relating productivity to AFDW of detritus have not been reported. The high values of P make it evident that detritus had little effect on community metabolism at sites of Augusta Creek (Table 18). Few of the F ratios were greater than one, which also indicated there was little influence on average rates or on an individual site basis (Table 18; Appendix G). These ratios ranged from 0.01 for average GCP to 1.3 for average NDM and on a site basis from 0.02 for NCP at B Avenue to 3.24 for CR at the Nagel Site. There were no clear patterns established as values greater than one were for different parameters. The correlations between parameters and detritus were weak, and measurement of detrltal standing crop was not of value in predicting rates of community metabolism at riffle sites of Augusta Creek. Epilithon Epilithon has been characterized in the literature mainly by estimates of biomass obtained from chlorophyll extraction (McConnell and Sigler 1956, Wetzel 1963, Duffer and Dorris 1966, Grzenda et al. 1968, Bowbdwna 1972, Cadee and Hegeman 1974, Helan et al. 1977, Seyfer and Wilhm 1977, Sumner and Fisher 1979, Bush and Fisher 1981). Dry weight and AFDW determinations on artificial substrates have also been 114 Table 18. Regression summary for the relationship between GCP, NCP, CR, and NDM and the independent variable detrltal standing crop (AFDW) for five sites of Augusta Creek, Kalamazoo County, Michigan. Dependent Variable X X X X N* R** F Ratio GCP NCP CR NDM 94 96 96 94 0.10 0.11 0.08 0.12 0.012 1.300 0.600 1.400 0.900 0.255 0.435 0.244 GCP NCP CR NDM 12 12 12 12 0.32 0.13 0.35 0.32 1.112 0.175 1.434 1.126 0.316 0.694 0.259 0.314 29 30 29 29 0.12 0.02 0.17 0.18 0.372 0.016 0.826 0.917 0.547 0.900 0.372 0.347 GCP NCP CR NDM 10 10 11 10 0.14 0.28 0.19 0.34 0.149 0.687 0.351 1.045 0.710 0.431 0.568 0.333 GCP NCP CR NDM 32 33 33 32 0.25 0.18 0.31 0.13 2.062 1.031 3.243 0.524 0.161 0.318 0.081 0.475 11 11 11 11 0.23 0.27 0.06 0.47 0.478 0.711 0.029 2.598 0.507 0.421 0.869 0.141 SMITH X X X X B AVENUE X X X X GCP NCP CR NDM UPPER 43RD X X X X NAGEL X X X X KELLOGG FOREST X X X X GCP NCP CR NDM * N = number of values ** R - Multiple Correlation Coefficient *** P = Level of Significance 115 used (Cushing 1967, Edberg and Hofsten 1973, Helan et al. 1973, Calow 1975, Naiman 1976, Seyfer and Wilhm 1977). These above studies indicated variable correlation between biomass and community metabolism. However, these studies were aimed at determining autotrophic colonization rather than the entire assemblage including detritus. M adsen (1972) found the epilithon of shaded Denmark streams to be detrital dominated, and Calow (1975) found epilithic detritus to be a high quality food for macroinvertebrates. Perkins and Kaplen (1978), reported epilithon to be 76% detritus and further characterized this segment to be accumulated autochthonously produced diatom stalks. Clearly, detritus is a portion of the epilithon and will be involved in any in situ measurements of community metabolism and the impact of the intact assemblage is of importance in the function of stream ecosystems regardless of the composition. On an annual average basis, a significant positive relationship existed between epilithon development and all dependent variables (Table 19). 19). F ratios ranged from 7.8 for NDM to 17.6 for NCP (Table The F ratios for epilithon followed stream order and light in order of magnitude (Appendix E ) ; these values were much more significant than estimates of detrital AFDW. Correlations ranging from R = 0.28 to 0.8 have been reported for biomass and primary productivity (Cadee and Hegeman 1974, Ertl and Tomajka 1973). Augusta Creek values of R = 0.42 NCP and R = 0.39 GCP (Table 19) are within this range. Various relationships have been demonstrated using chlorophyll a extractions as an estimator of biomass (Wetzel 1963, Schindler et al. 1973, Sumner and Fisher 1975, Marker 1976b, de laCruz and Post 1977, Bush and Fisher 1981). While chlorophyll estimates may 116 Table 19. Regression summary for the relationship between NCP, GCP, CR, and NDM with the independent variable epilithon for five sites of Augusta Creek, Kalamazoo County, Michigan. Dependent Variable X X X X p*** N* R** F Ratio GCP NCP CR NDM 83 85 85 83 0.39 0.42 0.31 0.30 14.8 17.6 8.6 7.8 0.002 0.000 0.004 0.007 GCP NCP CR NDM 12 12 12 12 0.41 0.76 0.38 0.66 1.966 13.601 1.699 7.696 0.191 0.004 0.222 0.020 23 24 23 23 0.24 0.21 0.19 0.10 1.270 0.979 0.796 0.215 0.273 0.333 0.382 0.648 GCP NCP CR NDM 9 9 10 9 0.00 0.03 0.16 0.37 0.000 0.007 0.206 1.126 0.992 0.935 0.662 0.324 GCP NCP CR NDM 28 29 29 28 0.07 0.08 0.20 0.18 0.113 0.189 1.104 0.881 0.740 0.667 0.303 0.357 11 11 11 11 0.19 0.38 0.17 0.63 0.321 1.523 0.260 6.254 0.585 0.248 0.622 0.034 SMITH X X X X B AVENUE X X X X GCP NCP CR NDM UPPER 43RD X X X X NAGEL X X X X KELLOGG FOREST X X X X GCP NCP CR NDM N = number of values ** R = Multiple Correlation Coefficient *** p = Level of Significance 117 reflect productivity changes, using eplllton estimates the entire community is included and integrated. The effects of epilithon development on CR and NDM have not been examined by many investigators. However, Bush and Fisher (1981) reported increased CR with increased biomass in a desert stream (Arizona). The relationships Indicated by regressions from Augusta Creek riffles were not high, but ranked above detrital standing crop estimates (Table 19). For individual sites several F values were greater than one, but few regressions were significant (P ■ 0.05). This indicates a stabilization in epilithon development at a site, with site differences reflected in the average epilithon and metabolic capacities. Dynamic changes in biomass per day would not be expected dealing with large initial epilithon weights (compared to cell or detrltal weights). physical factors, Rates of metabolism oscillated with changes in therefore, only when averaged over the annual period would these correlations become obvious. Average integration is further supported by the higher correlations at Smith site where epilithon levels were lower and changes were more readily detected. Sites Examining the regression data (Appendix E) for each dependent variable stream order was the most Important factor influencing community metabolism. Of the variable parameters light was by far the most Important parameter for all facets of community metabolism except CR where temperature was most important. At the sites controlling factors varied (Table E - 3). The highest and only significant correlation and F ratio was epilithon at 118 the Smith Site, which illustrates the Importance of even small amounts of detrital or algal accumulation. At the B Avenue Site the only significant relationships were with temperature for CR, GCP, and NDM, w h i c h indicated a bacterial dominated community. Upper 43rd Site had significant correlations between GCP and CR with temperature, Nagel had temperature correlating with NCP, CR, GCP and NDM and light with GCP, while the Kellogg Forest had significant regressions for GCP, CR, and temperature and NDM with light and epilithon. Temperature on a site basis was the influencing factor on average rates of community metabo l i s m and only when Individual rates were examined was the overwhelming importance of light evident. Particle-Size Community Metabolism The impact of various inorganic and organic particle-size distributions within sediments on community metabolism in lotic systems has received little attention (Minshall et al. 1982). M cConnell and Sigler (1959) and Marker (1976b) indicated that the larger inorganic particles contained approximately 95% of the algal biomass based on chlorophyll extr a c t i o n s , which indicated larger particles more suitable for algal colonization. Nelson and Scott (1962) studying CR rates in various sections of the Oconee River, Georgia, using the upstream-downstream technique, found CR to be insignificant on shifting sand substrates. This technique does not allow separation of discrete particle-sizes or detritus from inorganic sediments. Electron and epiflouresence microscopy of epilithon of small stones Indicated a prevalence of detritus (Madsen 1972) and Suberkropp 119 and Klug (1974) Illustrated that as detrltal particles decreased In size the major colonization shifts from fungal hyphae to bacteria. their micrographs algae (diatoms) were also present. In It Is of both theoretical and applied interest to examine the metabolism of various sediment partlcle-sizes and their contribution to total community metabolism. The activity on an AFDW basis (yl02 g~*) of various particle sizes m a y be compared and when converted to g O 2 m -2d"l the relative contribution of particles to overall community metabolism can be partitioned and the trends compared to in Bitu measurememnts H). (Appendix Comparisons of Gilson metabolism estimates to in situ rates of NCP, GCP and C R m “ 2 indicated parameters to generally be greater when estimated by the former method (Table 20). This trend, especially for macroinvertebrate-free sediments, m a y indicate the inadequacy of short term estimates, extrapolation to an areal basis assuming all layers of epilithon metabolize at equal rates, particle-sizing and assuming all particles behave the same in situ where depth of the particle would be of importance, or exposure through particle-sizing of surfaces to higher oxygen concentrations with subsequent increased activity of the flora, especially facultative anaerobes. However, the Gilson estimates are of value for comparative purposes and particle-size analyses. Particle-sized estimates of community metabolism (AFDW basis) NCP for detritus ranged from -319.7 y l 02 g"^h” l at B Avenue in August to +1687.6 y I O 2 g “ ^h” l at Upper 43rd in June; were always most active (Table H — 1; Figures 26, 27). 0.45 ym particles Annual averages 120 Table 20. Comparison of community metabolism parameters from in situ and Gilson Respirometer experiments from five riffle communities of Augusta Creek, Kalamazoo County, Michigan. Annual X Gilson g O 2 m " 2d ” l Annual X in situ g 0 2 m " 2d"l Parameter Site NCP NCP NCP NCP NCP Smith B Avenue Upper 43rd Nagel Kellogg Forest -0.79 -1.43 5.46 2.77 1.39 -0.16 - 0.02 1.24 1.80 0.94 GCP GCP GCP GCP GCP Smith B Avenue Upper 43rd Nagel Kellogg Forest -0.08 0.28 6.83 4.53 3.01 0.12 0.26 2.18 2.89 1.36 CR CR CR CR CR Smith B Avenue Upper 43rd Nagel Kellogg Forest 1.24 2.87 2.27 2.93 2.84 0.56 0.53 1.79 1.88 0.75 NDM NDM NDM NDM NDM Smith B Avenue Upper 43rd Nagel Kellogg Forest -1.32 -2.59 4.55 6.60 0.17 -0.44 -0.26 0.45 1.01 0.60 121 DETRITUS EPILITHON 0 SMITH -100 -500 W SP SU W SP SU 0 B AVENUE -100 r -500 W SP SU W SP SU CM a 5000 k U P P E R 43 rd r 1000 I W SP SU • 4mm o imm W S P SU SEASON Figure 26. ■ 250jum □ 75 urn a 0 - 45*101 Estimates of NCP from particle sized detritus and epilithon communities from Smith. B Avenue and Upper 43rd riffle sites of Augusta Creek, Kalamazoo County, Michigan (1974-1975). 122 EPI LI THON DETRITUS NAGEL 0 W -500 M " CL W SP SP SU m — SU 1000 O z 100 KELLOGG F OREST ■100 '5 0 0 W SP SEASON Figure 27. W SU • o ■ o A SP SU 4m m 1 mm 2 50>um 75jjm 0 • 45jum Estimates o£ NCP from particle-sized detritus and epilithon communities from Nagel and Kellogg Forest riffle sites of Augusta Creek, Kalamazoo County, Michigan (1974-1975). 123 for particle-sizes ranged from -213.3 to +485.3 Avenue and Upper 43rd, respectively (Table 21). pi02 at B The striking featurs of detrital NCP included negative values for the majority of particle sizes during all seasons, with least negative values of NCP generally during the February diatom bloom. June at the Upper 43rd Site. The only positive value was during Similar seasonal trends existed for all sizes within sites (Figures 26, 27). Comparing average rates of NCP between sites, B Avenue and Smith values were consistently more negative than the autotrophic sites (Table 21). Ranking the particles on increasing levels of NCP at each site (Table 22) there were no clear patterns within particle-sizes, but ranked by total average NCP, the 0.45 pin detritus had the only positive value, while the others were similar with the 75 p m , which were lowest in autotrophic colonization (Tables 21, 22). Estimates of epilithon N CP ranged from -321.0 to +2174.2 p l 0 2 g - 1h -1 for 75 p m mineral sediments in August at Smith and 4 mm sediments in M a r c h at the Kellogg Forest Sites, respectively (Appendix H; Figures 26, 27). Averages ranged from -155 particles to +752.9 p l 0 2 g “ *h“ l for 4 m m gravel at Kellogg Forest (Table 21). p l02 g ” ^h” ^ for 75 ym Based on rates of positive NCP, epilithon of all inorganic fractions at Upper 43rd were colonized by autotrophic organisms during all seasons, as well as the majority of the Nagel and Kellogg Forest particles (Figures 26, 27). Lowest rates of NCP at these sites occurred in August, while maxima were scattered in occurrence (Figures 26, 27). colonization, The heterotrophic sites had little algal but greater NCP occurred on these sediments than the detrital segements, especially in the 4 and 1 m m inorganic fractions. 124 Table 21. Average particle-sized community metabolism estimates (weight basis) from Gilson Respirometery for five riffle sites of Augusta Creek, Kalamazoo County, Michigan. Site X NCP 4mm X X CR -133.9 - 65.2 - 88.6 - 23.1 -107.4 - 83.6 DETRITUS X CR EPILITHON Smith B Avenue Upper 43rd Nagel Kellogg Forest X x /g 95.2 91.1 83.5 40.3 60.1 - 79.0 -133.0 - 50.0 - 54.2 - 62.6 -116.3 -130.1 - 25.6 - 94.8 - 86.4 -159.2 -213.3 +485.3 - 83.9 -114.9 -116.7 -124.5 + 47.5 - 59.3 - 86.3 - 74.0 - 75.8 - 90.6 17.2 - 67.2 64.0 111.9 34.1 124.1 42.1 94.9 125.7 42.0 64.3 63.0 78.5 129.7 71.5 91.8 93.9 160.6 351.0 157.6 262.0 104.5 80.3 73.8 86.9 205.0 8.3 116.3 35.8 752.9 - 18.7 - 25.5 75.1 20.0 57.2 -123.4 - 61.4 215.1 106.6 124.2 -155.0 - 96.3 386.0 205.8 214.6 178.2 21.6 52.2 111.0 68.5 90.6 171.7 67.9 151.7 ylo2 g- 1h -1 EPILITHON X 0.45pm 76.5 55.1 140.4 34.3 216.1 X Smith B Avenue Upper 43rd Nagel Kellogg Forest 75pm y l 02 g- 1h -1 Smith B Avenue Upper 43rd Nagel Kellogg Forest X NCP 250pm p l 02 g- 1h _1 DETRITUS Smith B Avenue Upper 43rd Nagel Kellogg Forest 1mm - 5.5 ----- - 75.7 - 47.9 198.1 92.1 287.2 ylo2 g - 1h-1 12.1 10.2 19.3 29.0 18.4 39.3 26.8 43.0 24.3 35.8 61.8 74.9 83.3 48.8 135.3 38.9 104.0 110.3 126.9 103.9 38.6 28.4 80.8 90.8 43.3 56.3 66.4 54.6 78.6 125 Table 22. Ranked particle-size contributions to various parameters of community metabolism (weight basis) from five riffle communities of Augusta Creek, Kalamazoo County, Michigan. Site X NCP Rank Particle size rank - Highest to lowest 1 2 3 4 5 DETRITUS Smith B Avenue Upper 43rd Nagel Kellogg Forest 250pm 4mm 0.45pm 4mm 1m m 1m m 1m m 75pm 1m m 250pm 75pm 75pm 250pm 250pm 75pm 4mm 250pm 1mm 0.45pm 4mm 0.45pm 0.45pm 4mm 75pm 0.45pm Sites Average 0.45pm 1m m 250pm 4mm 75pm Smith B Avenue Upper 43rd Nagel Kellogg Forest 0.45pm 0.45pm 0.45pm 0.45pm 0.45pm 4mm 250 pm 4mm 75pm 4mm 1mm 75pm 75pm 250pm 1mm 75pm 1mm 250pm 4mm 75pm 250pm 4mm 1mm 1mm 250pm Sites Average 0.45pm 4mm 75pm 1m m 250 pm 4mm 4mm 75pm 75pm 4mm 1m m 1m m 250pm 250pm 75pm 250pm 250pm 4mm 4mm 250pm 75pm 75pm 1m m 1m m 1m m 4mm 75pm 250pm 1m m 75pm 75pm 75pm 75pm 250pm 250pm 250pm 250pm 250pm 75pm 1m m 1mm 1mm 1mm 1mm 4mm 4mm 4mm 75pra 250pm 4mm 1m m X CR X NCR DETRITUS EPILITHON Smith B Avenue Upper 43rd Nagel Kellogg Forest Sites Average X CR EPILITHON Smith B Avenue Upper 43rd Nagel Kellogg Forest Sites Average 4mm 4m m 126 Sites ranked by average epilithon NCP in Increasing order were: Smith, B Avenue, Nagel, Upper 43rd, and Kellogg Forest (Table 21). Obvious differences were evident between the heterotrophic headwater sites and the open sites of Augusta Creek. Ranked particle-sized epilithon NCP (Table 22) showed variable results with larger sizes autotrophically colonized most heavily at Smith, B Avenue, and Kellogg Forest, while smaller particles at Nagel and Upper 43rd had the highest values of NCP. Averaging sites, the two extremes in particle-size had the highest rates of NCP with 1 m m particles lowest in activity (Table 22). Heaviest autotrophic colonization of larger particles has been shown by other investigators 1959, Duffer and Dorris 1966, Marker 1976b). (McConnell and Sigler Although primary productivity has not been reported on discrete particle sizes in the literature, Gargas (1970) reported the epipsammon to contribute 60 to 83% of the primary production of marine littoral sediments as compared to detritus and noted Gronsted (1960, 1962) found similar results in sediments of the Wadden Sea and Dani s h fjords. autotrophic colonization on fine particles These studies, with (sands) are similar to the increased levels of NCP o n the 75 urn inorganic sediments reported in this study. Partitioned CR (yl02 g ” ^h“ l) results for sediments indicated the detrital particles had higher rates of CR than the inorganic sediments. Higher (40 to 90%) detrital than inorganic sediment CR has been reported by Teal (1962) and Hargrave (1972) from lentic systems. The 4 m m and 1 m m inorganic particles from Augusta Creek approximated 35% of the detrital CR, but that of smaller particles (250 andp75 was approximately equal As particle-size (104 and 109% respectively). m) 127 decreases, surface area to volume Increases and floral shifts from fungi to bacteria were noted by Suberkropp and Klug (1974), which may have accounted for Increased rates of metabolic activity on the finer particles at Augusta Creek sites. However, Nelson and Scott (1962) found CR to be insignificant on shifting sand bottoms, which indicates a complex relationship between particle-size and CR, dependent on the location and stability of small particles within the sediments. Total inorganic CR averaged 70% of total CR at Augusta Creek riffles. Detrital CR ranged from 15.1 ylOj g “ ^h“ ^ for 4 m m fractions from Nagel Site in March to 738.8 yl02 g"^h“ l for 0.45 y m Upper 43rd detritus in June, while particle-size specific CR averaged over the early spring to late summer period ranged from 34 yl02 g - ^h- ^ for 4 and 1 m m detritus to 351 y l 02 g ” lh” l for 0.45 y m components, both at Upper 43rd (Appendix H; Table 21). The lowest rates at all sites were during the early spring; they increased during early summer and generally attained maxima in late summer (Figure 28). particulate organic matter The ultrafine (0.45 ym category = UPOM) rates of CR were consistently highest at all sites with most other particles similar and lower in rates of CR (Figure 28). The highest average rates were from Upper 43rd detritus followed by Kellogg Forest, B Avenue, Smith and Nagel (Table 21). The average activity by particle size (Table 22) indicated 0.45 ym detrital particles had the highest rates of CR followed by 4 m m particles with 75 ym, 1 mm, and 250 ym detrital particles similar. These data are unlike those of Fenchel (1970) and Hargrave (1972) who found detritus CR rates inversely related to particle size. Based on surface to volume ratios of particles an inverse relationship may be expected, however, this premise would be 128 500 EPILITHON DETRITUS S MI T H 100 0 B AVENUE 100 0 C R *1102 9"l A P D w 1000 UPPER 100 43rd NAGEL 0 500 KELLOGG F O R E ST 100 W SP W SU SP SU • 4 mm o 1m m 250jim □ 75 Aim A 0 -45Pm SEASON Figure 28. Estimates of CR from particle-sized detritus and epilithon communities from selected riffle sites of Augusta Creek, Kalamazoo County, Michigan (1974-1975). 129 based on equal colonization of surfaces by a similar flora (i.e. bacteria). In heterogeneous layered substrates differences would be expected in types of colonizers (i.e. components) and particle location (i.e. autotrophic-heterotrophlc light availability). Epilithon CR was also lowest during spring periods; Increased during early summer; and remained stable at Smith and B Avenue Sites during the summer, but decreased at the Upper 43rd, Nagel and Kellogg Forest (Figure 28). The decrease at the autotrophic sites resulted during the period of algal senescence; detrital CR increased at the same time indicative of algal inputs to the detrital pool (Edberg and Hofsten 1973). Rates of epilithon CR ranged from nondetectable for 250 V>m sands at Stnith, Nagel, and Upper 43rd Sites in early spring to 214.4 Ul02 g “ lh“ l H). for 75 ym sands in June at the Kellogg Forest (Appendix Averaged by site, the highest CR was at the Kellogg Forest and Upper 43rd Sites with B Avenue and Nagel Sites approximately equal and Smith lowest (Table 21). The similarity of B Avenue and Nagel epilithon CR would Indicate similar levels of colonization, but the percent composition of autotrophs must have varied significantly based on the NCP values (Tables 9, 21). Ranked by increasing activity, the epilithon CR patterns followed the inverse relationship with particle-size proposed for detritus by Fenchel (1972). Comparing particle-sizes, (1970) and Hargrave a similar trend existed; averages for the 75 yim sands were the most active while the 1 m m were lowest at all sites (Table 21). Overall on a weight basis, detritus had higher CR and lower NCP than epilithon. 130 Particle-sized estimates of community metabolism Areal basis) On an areal basis, as on a weight basis, the autotrophic colonization was in the epilithon community, but on an areal basis, especially for epilithon, general trends were similar to in situ. This agreement further substantiates the magnitude of epilithon biomass in riffle sections. Average detrital NCP m - 2 , as on a weight basis, was generally negative with the highest and only positive value at Upper 43rd for 0.45 pm detritus (Table 23). The lowest average values were also for this segment at B Avenue and Smith Sites (Table 23). Averaged for particle-size, the lowest values of NCP were for the UPOM and highest for the CPOM fractions, which differed from a weight basis where UPOM had highest average values (Tables 21, 23). When particles were ranked by site for NCP (Table 24) the averages had a similar pattern to a weight basis, but generally there was direct relationship between rate and particle size not evident on a weight basis (Tables 22, 24). (AFDW) Areal ranked sites were identical to biomass (Tables 22, 24). The highest average NCP rates were from the epilithon of Upper 43rd 4 mm inorganic sediments, while the lowest rates were from 75 pm sands from the Smith site (Table 23). On an average particle-sized basis contributions of the 4 m m stones were highly separated from the other rates of NCP; the 75 p m sands had the lowest activity. On the weight basis the 75 p m fractions had high levels of activity (Tables 21, 23). Ranking sites by NCP m -2 (Table 23), Smith was lowest followed by B Avenue, Kellogg Forest, Nagel and Upper 43rd, while on the weight ranking the autotrophic sites shifted to Nagel, Upper 43rd, and Kellogg Forest. This shift illustrates the effects of epilithon 131 Table 23. Average estimates of community metabolism of various particle-sizes expressed on an areal basis for five riffle sites of Augusta Creek, Kalamazoo County, Michigan. Site X NCP 4mm X X NCP EPILITHON Smith B Avenue Upper 43rd Nagel Kellogg Forest X X CR DETRITUS Smith B Avenue Upper 43rd Nagel Kellogg Forest X X CR EPILITHON Smith B Avenue Upper 43rd Nagel Kellogg Forest X 250pm 75pm 0.45pm Total i -501.2 -784.0 136.7 -198.3 -265.7 mg 02«n'-2d -l DETRITUS Smith B Avenue Upper 43rd Nagel Kellogg Forest 1mm 32.0 35.6 28.2 11.9 4.1 - 42.3 -131.5 - 31.3 - 14.5 - 24.9 - 48.2 -143.1 - 31.8 - 26.6 - 43.6 - 50.1 -132.4 - 2.2 - 34.1 - 51.9 -328.7 -341.3 +225.7 -111.3 -141.3 - 22.4 - 48.9 - 58.7 - 54.1 -132.2 - mg 02in‘-2d-l .1 -380.6 5046.6 2446.9 1586.0 - 96.9 -108.3 137.4 155.3 70.3 -173.6 -138.2 117.7 344.0 87.4 - 22.3 - 15.4 23.0 22.0 95.6 1739.7 31.6 46.9 20.6 ------ -292.3 -194.9 5324.8 2968.3 1654.4 mg 02m -■2d-l 28.3 50.6 54.7 22.0 14.8 48.7 173.2 63.3 24.9 70.4 45.9 189.5 73.0 35.0 74.7 48.2 152.5 97.7 44.4 99.0 339.6 462.2 446.1 338.2 562.3 34.1 76.1 83.6 88.4 429.6 510.8 1028.0 748.6 464.4 821.1 mg 0?m"■2d -l 438.5 1339.3 1382.4 2070.6 1714.9 105.8 188.2 70.0 200.3 117.3 155.7 284.5 59.0 169.0 101.1 26.1 29.9 12.4 26.1 83.0 1389.1 136.3 153.9 35.5 726.2 1841.8 1523.8 2466.0 2016.4 132 Table 24. Ranked particle-size contributions on an areal basis to estimates of community metabolism from five riffle communities of Augusta Creek, Kalamazoo County, Michigan. Rank Site X NCP Particle size rank - Highest to lowest 1 2 3 4 5 DETRITUS Smith B Avenue Upper 43rd Nagel Kellogg Forest 4mm 4mm 0.45ym 4mm 4mm 1m m 1mm 75ym 1mm 1mm 250ym 75ym 4mm 250ym 250 ym 75ym 250ym 1m m 75ym 75ym 0.45ym 0.45ym 250ym 0.45ym 0.45ym Sites Average 0.45ym 1m m 250ym 4mm 75 pm Smith B Avenue Upper 43rd Nagel Kellogg Forest 4mm 75yra 4mm 4mm 4mm 75ym 1mm 1m m 250ym 75ym 1m m 250ym 250ym 1m m 250ym 250ym 4mm 75ym 75ym 1mm Sites Average 4mm 250ym 1mm 75ym Smith B Avenue Upper 43rd Nagel Kellogg Forest 0.45ym 0.45ym 0.45ym 0.45ym 0.45ym 1mm 250ym 75ym 75ym 75ym 75ym 1m m 250ym 250ym 250ym 250ym 75ym 1mm 1mm 1mm 4mm 4mm 4mm 4mm 4mm Sites Average 0.45ym 75ym 250ym 1mm 4mm Smith B Avenue Upper 43rd Nagel Kellogg Forest 4mm 4mm 4mm 4mm 4mm 250ym 250ym 1mm 1mm 1mm 1m m 1m m 250ym 250ym 250ym 75ym 75ym 75ym 75ym 75ym Sites Average 4mm 250ym 1mm 75ym X NCP X CR X CR EPILITHON DETRITUS EPILITHON 133 development and detrital content of the sediments, ^ n situ estimates of average NCP ranked more closely with epilithon than detrital values (Tables 9, 21, 23). Even though CR estimates g~l AFDW detritus were consistently higher than for epilithon (Table 21), when expressed on an areal format the reverse was true and the impact of the epilithic community established (Table 23). UPOM for which there is no inorganic counterpart had higher CR than all of the Inorganic fractions except 4 m m stones, noting the importance of the two extremes in particle-size and sediment types in the autotrophic-heterotrophic balance of riffle sections. Detrital CR was highest (average m~2) at the Kellogg Forest for UPOM and lowest at Nagel Site for 4 m m pieces. Averaged by particle-size the highest and lowest components remained the same and were similar to a weight basis. Areal particle-size ranking indicated the generally accepted trend of CR inversely related to particle-size (Fenchel 1970, Hargrave 1972, Naiman and Sedell 1979b). On a weight basis this pattern was characteristic only of the Nagel site (Table 22 ). Epilithon CR was higher than detrital on an area basis, and even the lowest Smith epilithon had higher CR m “ 2 than all except B Avenue and Kellogg Forest detrital particles (Table 23). On an average the 4 m m particles dominated CR at all of the sites and 75 pm sands contributed the least to epilithon CR. These average trends were characteristic of all sites with a direct relationship noted between particle-size and CR (Table 23). the reverse was true. On the weight basis (Tables 21, 23) This reversal indicated a more active, but lesser developed microflora on the sands. 134 When extrapolated to an areal basis, negative values of NCP, GCP, and NDM were consistently measured on detritus at the Smith Site with the 0.45 pm fraction predictably lowest; the others were significantly different and closely grouped (Figure 29). The 4 m m detritus, indicating some algal colonization, had the highest rates of NCP and the lowest rates of CR. CR was highest in the 0.45 ym fractions and increased with seasonal temperatures as did in situ CR (Figure 16). The consistently low NDM and P /R ratios of both detrital and epilithon fractions indicated heterotrophic colnization of all substrates and the heterotrophic character of this first order site (Figures 17, 18, 29, 30). Within the epilithic community, as well, little autotrophic potential existed (Figure 30). The only positive value for NCP was on 4 m m substrates during the February diatom bloom, which indicated a more diverse flora was present. The 75 y m sands had the least negative and most consistent levels of NCP, but as evidenced by GCP values, only in June rose above baseline in primary productivity. The 1 mm and 75 y m inorganic sediments had maxima in GCP just after the major diatom bloom which may be the result of diatom community changes within the sediments from the Meridion - Diatoma bloom forms to smaller motile migratory forms which may colonize smaller sediments. However, only taxonomic analyses could substantiate this hypothesis. The comparison of particle-size specific epilithon NCP trends to in situ measurements indicated similar patterns as did the detrital component, which reinforced the dominance of heterotrophic processes with the riffle community. B Avenue detritus, like Smith, had negative values of NCP, GCP, 135 SMITH - NCP DETRITUS GCP N D M 100 ■100 •1000 I W SP 1 SU W SP SU W SP SU ■o CM I E CM P/ R O 0> 1 E CR 0 1000 -A 1 100 • W SP • 4mm o 1m m ■ 250jum a 75um a o ■45jum 2 W SU SP SEASON Figure 29. Estimates of particle-sized community m etabolism of the detrital component of Smith riffle sediments, Augusta Creek, Kalamazoo County, Mic h i g a n (1974-1975). 136 S M I T H — EPI L I T H O N NDM GCP NCP 1000 100 -100 -1000 W SP SU W SP SU I W SP su •o CM » E CM O 0> E C R 1000 • 4 m m o 1 mm ■ 250Jum 100 ° W Figure 30. SP 7 5 Aim SU SEASON Estimates of particle-sized community metabolism of the epilithon of Smith riffle sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975). 137 and NDM which mimiced In situ trends (Figues 31, 14-18). On a comparative scale the UPOM particles were lowest and 4 m m detrital particles highest In autotrophic potential. The similarity In rates of NCP and CR of the other particles Indicated that after breakdown to 1 m m and finer, detrital particles had a consistent heterotrophic (probably largely bacterial) flora; the UPOM, where increased surface area to particle size was greatest, had maximized colonization and increased rates of CR. P/R ratios, indicating the heterotrophic nature of detritus in the B Avenue riffle sections, were similar in magnitude to those of Smith detritus and all were significantly less than one (Figure 31). The primary productivity capacity of the inorganic substrates at the B Avenue riffles also depicted a dependence on allochthonous substrates (Figure 32). the sampling period. NDM was below zero for all particles over NCP was positive only for 4 mm and 250 ym particles in the spring during the diatom bloom; levels of NCP were highest for most particles during that period. As in s i t u , the P/R ratios were consistently less than one with the highest levels in spring; patterns of epilithon CR were also similar to in situ (Figures 32, 14-18). When comparing detrital and epilithon autochthonous contributions to riffle community metabolism at this site it was obvious that the little algal colonization which occurred did so in the epilithon community. As on a weight basis, the small autotrophic potential demonstrated at the Upper 43rd Site in the detrital community, was due to the 0.45 ym particulate fraction in June (Figure 33). This June maximum was evident in in situ measurements of NCP, GCP, and NDM as 138 B AVEN UE — D E T R I T U S NCP GCP NDM ioo n -100 -1000 W SP SU W SP SU W SP SU ■o Cl Cl o 0) E P/ R CR 1 1000 100 • o ■ □ 4mm 1mm 250jum 75 Aim a 0’45j*m 0 ■1 W SP SU W SP SU SEASON Figure 31. Estimates of particle-sized community metabolism of the detrital component of B Avenue sediments, Augusta Creek, Kalamazoo County, M ichigan (1974-1975). 139 B AVENUE 5000 NCP m — EPILITHON GCP NDM 1000 100 •100 -1000 -5000 W SP ■o CM I W SU P/ R SP SU E CM O o> E C R 5000 •4mm o 1mm ■ 250jum 1000 a 75pm 100 SEASON Figure 32. W SP SU Estimates of particle-sized community metabolism of the epilithon of B Avenue riffle sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975). 140 UPPER 43RD —DETRITUS NCP 5 0 0 0 GCP N D M 1000 100 -100 -500 W SP W SP SU W SP SU ■o P/ R C R 1000 • ° ■ 100 W S P SU W SP d 4m m 1 mm 250JJm 75ju m A 0 .4 5 ju m SU S E A S O N Figure 33. Estimates of particle-sized community metabolism of the detrital component of Upper 43rd riffle sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975). 141 well (Figures 33, 14, 15, 18). Although NCP was positive only during June, GCP in the UPOM fraction was positive from February to August. During February, diatom colonization was evident on all available surfaces and could well have been a part of the FPOM pool. The June increases in the UPOM fraction as well as the 75 ym particles were related to periods of algal input. diatoms of the spring bloom: Length measurement of two common Diatoma vulgare Bory (30 to 60 ym length) and Merldon circulare (12 to 80 y m length) support this hypothesis. (Hustedt 1930) NDM in the 75 y m fraction indicated carbon was produced in excess during June and August, while the other particle sizes were generally below zero in NCP and NDM (Figure 33). GCP was positive for most particles; the 75 ym and 0.45 ym particles had the highest values. CR was highest for the UPOM with other sizes lower and similar (Figure 33). P/R ratios were generally below one, but much higher than the Smith and B Avenue values. These data suggest algal colonization and autochthonous contributions to the energy base of the Upper 43rd riffle by the detrital community. Based on levels of particle-sized epilithon primary productivity, the Upper 43rd epilithon community had a definite algal component (Figure 34). The only similarity with detrital contributions was levels of CR (Figure 33); even maxima of productivity occurred at different times with epilithon proceeding detrital, which added support to the algal input hypothesis. NDM and P/R ratios were above one Indicating an autotroph dominated system in most cases. The 4 mm inorganic particles had the greatest contributions to the autochthonous organic pool of the riffle section as indicated by values of NCP, GCP, NDM, and P/R (Figure 34). High rates of CR for the larger inorganic 142 U P P E R 4 3 R D - E P I N C P L l T H O N G C P N D M 10,0001 1000 100 0 W SP W SP su W SP •o CM cf P/R o> 50 E 10 C R 5000 1000 B • o ■ d 100 4 m m 1 m m 250jim 7 5 Aim w S E A S O N Figure 34. Estimates of particle-sized community metabolism of the epilithon of Upper 43rd riffle sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975). 143 substrates Indicated higher colonization both heterotroph and autotroph (Figure 34). all standpoints. The smallest sands had the least impact from While detrital patterns In NCP (Figure 33) did not follow in situ trends, the epillthic contributions did, which indicated a dependence on epilithon development for increased NCP. At the Nagel Site patterns of detrital community metabolism were similar to those of Upper 43rd (Figures 33, 35). negative and very similar for all NCP values were segments, GCP was positive during February diatom bloom for 4 and 1 m m pieces and 0.45 y m detritus in June and August (Figure 35). algal sloughing into the detrital pool. the The summer increases were related to CR was also highest for this site in the UPOM with other sizes lower and similar in value. P/R ratios were all less than one and intermediate between the extreme lows of B Avenue and Smith Sites and the higher Upper 43rd values. The only trend similar to in situ measurements was for CR (Figure 35). Particle-sized estimates of community metabolism at the Nagel site further expanded the importance of the larger stones as substrates for colonization (Figure 36). The patterns and magnitudes of parameters were similar to those of the Upper 43rd epilithon, and reflected in situ trends. This further substantiates the productivity potential of inorganic sediments within autotrophic riffles (Figures 34, 36). Consistently below zero levels for NCP, NDM and P/R characterized the heterotrophic nature of the Kellogg Forest detritus. The 4 mm fragments had the highest percentage algae/surface area as it yielded the highest estimates of NCP and NDM with the lowest rates of CR (Figures 37). The highest CR and corresponding lowest NCP and NDM 144 N A G E L - D E T R IT U S NCP N D M G C P 500 100 ■100 -500' W SP SU W SP SU W SP SU XJ • CM E cf o> P/R E 1 • 4mm ° 1mm CR 0 500 ■ 25Q iim ° 75iim A 0.45 Jim 100 -1 W SP su S E Figure 35. Estimates of particle-sized community metabolism of the detrital component of Nagel riffle sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975). 145 NAGEL — EPILITHON NCP 10,0 0 0 , 1000 • nr W SP GCP NDM 100 -100 I TJ su W SP W SP CM a 100 E P/ R o' o* E CR 10,000 10 1000 \ 100 \ • 4mm o 1mm ■ 250jum a 75jum SEASON Figure 36. Estimates of particle-sized community metabolism of the epilithon of Nagel riffle sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975). 146 were from the UPOM particles. GCP was positive for the UPOM segment which indicated potential algal inputs into this pool, yet NDM remained below aero for all segments. Nagel P/R ratios were similar to (below one) and intermediary to the extremes. The UPOM ratios were again the highest and most consistent (Figure 37), CR patterns were similar for all groups of detritus except 75 ym, which may have been increased by algal inputs, as at Upper 43rd. Illustrating the dependence of NDM on CR and in turn amounts of detritus in the sediments, estimates of detrital NDM and in situ NDM indicated similar p a t terns. General correspondence with in situ trends was seen only for the 0.45 Pm fraction, which notes the importance of this fraction in the autotrophic-heterotrophic community balance (Figures 37, 14-18). Autotrophic productivity of the epilithon at the Kellogg Forest was lowest during the early spring, increased in summer and decreased in late August (Figure 38). NCP and GCP values were positive for most particles over the experimental period and were similar to in situ (Figures 14, 15). CR rates followed the same pattern as NCP, GCP and in situ except 4 m m epilithon which remained high and stable (Figures 16, 38). NDM was characteristically above zero and was similar to in situ patterns (Figures 18, 38). In all cases 4 m m particles had the highest Impact on overall NCP, GCP, CR, and NDM others were similar in levels of activity. The P/R ratios of 1 m m sands were consistently above one, while each of the others at some period fell below the compensation point (Figure 38). The only inorganic particles which followed in situ P/R trends were the 4 m m stones (Figures 17, 38). This site indicated a photosynthetically active community with all 147 KELLOGG FORES T-DETRITUS NCP 1000 GCP NDM I 100 -100 -500* W SP SU W SP SU W SP SU ’I *0 CM E 6 1 O) E P/ R CR 1000 100 W Figure 37. • 4m m o imm ■ 250pm a 75jum a 0-45A>m ---- SP su SEASON SP SU Estimates of particle-sized community metabolism of the detrital component of Kellogg Forest riffle sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975). 148 KELLOGG F O R E S T — EPI L I T H O N GCP NCP 5000 N D M 1000 100 •100 •1000 -50001 W SP w SU SP W SU SP su •o eg P/R 5 2 .eg o O) E CR 5000 1 1000 • 4mm o 1mm ■ 250 Aim □ 75Aim 100 0 1 W SP SU W SP su SEASON Figure 38. Estimates of particle-sized community metabolism of the epilithon of Kellogg Forest riffle sediments, Augusta Creek, Kalamazoo County, Michigan (1974-1975). 149 inorganic sediments contributing to the autotrophic potential of the riffle reach. H o w e v e r , the majority of colonization occurred on 4 m m and larger particles. SYNTHESIS Babbling brooks won't tell... Sunlight dappling jubilantly won't tell... Neither will the stately adjacent tree or jewelweed glistening in a patch of sunlight bankslde... Of the complexity within. February 1982 Using a chamber m e t h o d , in situ measurements of community metabolism, light, temperature, and substrate composition were monitored over a two year period in five riffle sections of Augusta Creek, Michigan. These included the first order shaded Smith and B Avenue Sites, m eadow second order Upper 43rd Site, cleared third order Nagel Site, and the shaded third order section in the Kellogg Forest. During 1973-1974 daily estimates were made each month at B Avenue and Nagel Sites and daily seasonal estimates were made at the five sites during 1974-1975. Physical and biological differences between sites were illustrated and reinforced the complexity of patterns of stream metabo l i s m and productivity potential of woodland stream riffle sections. Temperature Typical temperate region seasonal patterns of temperature occurred at each site with maximum temperatures during the summer and minima during the winter. M a xima at the sites ranged from 19.9 C at 150 Smith and B Avenue Sites to 24.8 at Upper 43rd Site. Average daily temperatures ranged from 0.4 C at Upper 43rd in winter to 22.6 C at Nagel in August. First order sites were lowest, open sites highest, and shaded Kellogg Forest intermediary. Average daily temperatures plateaued at maximum temperatures in June and remained high until September at both shaded and open sections. Greatest daily temperature fluctuations ranged from 5.8 to 10.5 C with the highest fluctuations occurring at first order sites during spring and fall. Indicating the importance of canopy in maintaining cooler conditions, the largest annual averages were for the open sites. Light Average daily light measured at the riffle sections was highest (90,213 lux) at Upper 43rd in May and lowest (2,391 lux) at B Avenue in January, while averages over the study period ranged from 12,054 lux at Smith to 43,622 lux at Upper 43rd. First order sites were lowest, open sites highest and Kellogg Forest Intermediary. Smith and B Avenue sites received only 45% of the illumination of the shaded third order Kellogg Forest and only approximately 30% of the open sites. Open sites received approximately 140% of the shaded third order sites and Kellogg Forest approximately 66% of the light at open sites. Open sites displayed light patterns typical of temperate regions with spring-summer maxima and winter minima. June through September (leaf-out) light reductions were evident at Smith and B Avenue sites as were Increases in light after canopy abscission. Even though complex dappled light patterns were evident at the Kellogg 151 Forest site, significant light Inputs were evident. On a comparative scale lower light values were evident due to shading at these three sites even during winter. Ranking by average light input Smith was lowest followed by B Avenue, Kellogg Forest, Nagel, and Upper 43rd. Thus, light trends indicated the diversity of light patterns natural and man made along a continuum. Community Metabolism GCP values, ranging from 0.09 to 5.35 g O2 nT ^ d ” ! at Smith Site in winter and Nagel Site in August, respectively, varied seasonally and between sites over the annual period. Highest values were during the spring at naturally shaded sites and summer at open sites. On an average the lowest levels were at Smith followed by B Avenue, which had a wider channel and approximately two times the GCP of Smith; Kellogg Forest; Upper 43rd; and Nagel. Levels of GCP from the first order sites were lower than reported literature values. Open sites had approximately 13% higher GCP than the first order reaches. Clearing shifted the autotrophic capacity to values typical or larger rivers. NCP values followed the seasonal trends displayed by GCP and ranged from -0.25 g O 2 m ”2d“ l at Smith during the fall to 4.03 g O2 m “ 2d"l at Nagel Site in August. The low value, during the fall, may have been the effect of leaf inputs into the shaded Smith reach and the maximum value correlated with luxuriant algal development at Nagel Site. Indicating heterotrophic conditions and a dependence on allochthonous or stored organic matter, negative NCP values were 152 consistent at the first order sites. Forest Open sites and the Kellogg Site indicated positive NCP values, thus, autotrophic bases. Sites ranked by average NCP were Identical to GCP trends with the effects of clearing obvious from comparison of Kellogg Forest and Nagel Sites. Light saturated NCP occurred within the range of 15,000 to 24,000 lux at sites of Augusta Creek and combined sites indicated a saturation level of approximately 20,000 lux. Smith exhibited no saturation plateau, but light values rarely exceeded 20,000 lux at this site. Temperature did not affect light saturation thresholds. No photoinhibition was indicated at either autotrophic or heterotrophic sites. Rates of CR varied significantly between shaded and open sites. The range of values was 0.24 g O 2 m “ 2d“ l during January at B Avenue to 3.67 g O 2 m " 2 d ” l during July at Nagel Site. On an average basis B Avenue and Smith were similar and low, Kellogg Forest was intermediate, and Nagel and Upper 43rd were similar and higher in values. Kellogg Forest rates approximated rates of the heterotrophic sites with the only significant deviation occurring during periods of leaf-out when CR levels increased at Kellogg Forest corresponding with increased epilithon development. Seasonal patterns were similar at most sites with lowest rates during the winter and highest values during spring or summer. An invars®~EeLatioasMp--w&3..»ofced~ between CR and NCP at heterotrophic sites. CR as a percent of GCP varied from 445% at Smith to 55% at Kellogg Forest. Autotrophic sites averaged 65% while heterotrophic sites averaged 322%, which indicated a considerable deficit of organic carbon from autochthonous organisms at B Avenue and Smith Sites. Effects of clearing at Nagel compared to 153 the shaded Kellogg Forest Indicated Increased CR with greater epilithon development at the Nagel Site* P/R ratios at the five sites indicated consistent consuming modes at B Avenue and Smith and producing modes at the open and Kellogg Forest Sites over the year. Upper 43rd, closest to P/R » 1, indicated little excess autochthonous production for export or storage. Lowest P/R ratios were obtained on a snowy overcast day at B Avenue (-0.017) and highest (2.5) at the Nagel Site in August. ranked sites in increasing order: Kellogg Forest. was noted. Average P/R ratios Smith, B Avenue, Upper 43rd, Nagel, A trend of Increasing P /R with increased stream order The high levels at Kellogg Forest were attributed to lesser development of epilithon with increased light availability maximizing efficiency and, thus, indicated the autotrophic potential of shaded third order reaches. Highest ratios corresponded with trends of GCP and NCP with spring peaks at all sites and maximum in summer at the cleared Nagel site. at all sites. Low values were during the winter The low ratios obtained at Smith and B Avenue were similar to those reported from turbid or polluted systems and the open sites approximated values of larger rivers. Average P/R ratios from monthly estimates at B Avenue and Nagel Sites were not significantly different from averages obtained on a seasonal basis, which indicated seasonal monitoring was sufficient to characterize the autotrophicheterotrophic capacity of a riffle. NDM values were similar to NCP and P /R trends, but were lower in magnitude. Values ranged from -0.72 g O 2 m ^ d - * in fall at Smith to 2.68 g O 2 m -2d” l in August for Nagel. winter, Lowest NDM values were in the and at B Avenue and Smith consistent negative values indicated 154 dependence on allochthonous POM since no pool of autochthonous carbon was present. Lowest values, during fall at Smith, indicated the effects of increased CR concominant with CPOM inputs. Open sites and the Kellogg Forest Site were autotrophic in nature as indicated by positive NDM values with Nagel and Kellogg Forest capable of storage or export of autochthonous carbon and Upper 43rd Site was basically self-supporting with periods of potential dependence on the slight storage capacity or POM or allochthonous carbon corresponding to periods of low NCP and NDM. Average NDM was lowest at Smith followed by B Avenue, Upper 43rd, Kellogg Forest and Nagel, which Indicated that even though the highest P/R ratios were at Kellogg Forest, actual carbon production was greatest at Nagel and that the shaded Kellogg Forest produced more excess carbon than the open Upper 43rd Site. Detrital standing crop analyses indicated stable levels of detritus at each site with all but the extremes (Smith high and Upper 43rd low) similar in amounts. m~2. Standing crops ranged from 592 to 145 g No consistent seasonal trends were found between sites. First order sites had greater variance in standing crop levels while open and Kellogg Forest Sites remained stable. Average detrital standing crops for sites ranked in decreasing order were Ssith, Nagel, B Avenue, Kellogg Forest and Upper 43rd. The high levels of detritus at Nagel were attributed to inputs of filamentous algae and aquatic macrophytes as were the high average CPOM/FPOM ratios. This ratio varied from 0.06 during summer at Kellogg Forest to 0.68 at Nagel Site in winter with an inverse relationship between CPOM/FPOM and P/R. On an average basis sites ranked in CPOM/FPOM ratio in increasing order were Kellogg Forest, Smith, Upper 43rd, B Avenue, Nagel. 155 Patterns of community metabolism expressed on a g~^AFDW detritus basis did not reflect levels of standing crop, as values varied in activity differently from detrital ranking. CR maxima were highest during the fall in contrast to spring and summer on an area basis, which was attributed to a more labile (algal) input to the riffles at that time. First order sites were more stable in rates of community metabolism per gram detritus over the year which indicated a bacterial metabolic base. NDM values, indicating a dynamic equilibrium with each community of either an autotrophic or heterotrophic energy base, were consistent at each site. Since standing crop levels were not significantly different at most sites, little change in community metabolism trends were expected expressed on a weight basis. The concept of use of epilithon development (AFDW) as a mechanism to monitor autotrophic-heterotrophic capacities of riffle reaches was introduced. Values, obtained by ashing inorganic substrates including detrital and algal biomass, ranged from 1480 g m~2 at Smith during spring to 5030 g m~2 at Nagel during the same period. Obvious visual differences as well as biomass values varied with site and followed patterns of NCP and GCP in the riffles. Sites ranked Smith, B Avenue, Kellogg Forest, Nagel and Upper 43rd in average epilithon development. Seasonal variation was greatest at the open sites (C.V.%), while first order heterotrophic sites remained low and stable indicative of a bacterial base. Kellogg Forest was also stable, but had higher levels of epilithon development than the first order riffles. Even at Smith Site epilithon was a significant component of the total organic standing crop. Based on the low rates of NCP and GCP at this site it was detrital based epilithon, while open and shaded third order sites 156 had algal based epilithon. Peaks of epilithon occurred in fall at Smith during CPOM inputs and at open sites during spring and fall, w hich correlated with the diatom blooms. the larger substrates. Major changes occurred on Detritus averaged 12% of the TPOM; the range was 18% at Smith Site to 7.5% at Upper 43rd Site. between autotrophic and heterotrophic sites. Differences existed Use of the E/D ratio for characterization of riffle reaches was proposed. Average E/D ranged from 4.5 at Smith to 14.8 at Upper 43rd with increased importance of the epilithon (Algal and detrital) at the open sites. E/D had a direct relationship with NCP, GCP, and P/R for sites of Augusta Creek. However, higher values of E/D were obtained on an extremely heterotrophic riffle reach of a branch of the Yellowstone River, Montana, which indicates E/D indicative of the epilithon to non-attached detritus contribution to riffle energy bases, but NCP, P/R, and NDM estimates indicate the detrital-algal heterotrophic) contributions of the components. (autotrophic- E/D ratios used with P/R and NDM were of value in characterization of riffle sections. Regression Analyses Regression analyses involving the Independent variables stream order light, temperature, detritus AFDW, epilithon AFDW, and total particulate organic matter and the dependent variables NCP, GCP, CR, and NDM indicated stream order to be the most highly correlated parameter with all dependent variables (R = 0.52 to 0.74; F = 37 to 122). Therefore, the importance of geomorphology was evident from a functional standpoint. Light was also highly significant (P < 0.01) in cases of NCP, 157 GCP, and NDM based on average values (R ■ 0.52 to 0.57; F « 36 to 46) and on an individual basis (R *» 0.6; F ■ 1090). In multiple regression with temperature R - 0.597 for light while the temperature R value was only 0.005. Compared with other variable independent variables monitored, light was the most Important factor influencing NCP, GCP and NDM. On a site basis positive relationships were evident with light, but not as highly correlated. Significance increased with increasing order which indicated the increasing Importance of light on open sites. Temperature had a positive significant relationship with all average parameters with CR highest (R = 0.65; F = 70) and NCP lowest (R * 0.33; F =» 12). As CR varied little over the annual period in headwater first order streams, the relationship was less clear which indicated a stable community in dynamic equilibrium. Temperature was most strikingly related to NCP at open sites; on a site basis the most significant relationships were with temperature. The detrital content of the sediments had little effect on averaged or site basis community metabolism. Average NCP, and NDM were most highly correlated (R » 0.12; F - 1.3 and R ■ 0.12; F «* 1.4), but not highly significant (P ■ 0.26, 0.24). Epilithon development was highly significant with all averaged dependent variables with NCP correlating most closely (R = 0.42; F = 17.6). Although on a site basis significance levels decreased, a much stronger relationship existed for epilithon and rates of community metabolism than detritus. Total AFDW correlations were slightly reduced by addition of detritus in most cases, but were close to the values for epilithon. 158 Particle Sized Community Metabolism The Impact of various particle sizes (4 mm, 1 mm, 250 pm, 75 pm, 0.45 ym) for detritus and epilithon (Excepting 0.45 ym) was determined. Activity g” ^AFDW and areal estimates based on standing crop data were compared to examine the size acitvity and ecological impact respectively. All estimates were higher than in situ which indicated the inadequacy of short term estimates at light saturation levels for estimation of community metabolism, the possibility of disturbing particles in separation, or the potential of maximizing surface in Gilson vessels versus the packed condition typical of in situ conditions. However, relative comparisons m ay be made between particle sizes and sites. Absolute rates of detrital NCP ranged from -318 to 1688 ylOj g"2h“ l for 0.45 ym detritus at B Avenue and Upper 43rd respectively. At all sites most particle sizes were similar and negative in NCP except the 0.45 ym fraction, triiich was generally most negative in value. Average rates at Smith and B Avenue were considerably lower than the autotrophic sites and average particle size rates were highest for 0.45 ym followed by 1 mm, 250 ym, 4 mm, and 75 ym. CR of detritus particles ranged from 15 to 739 yl02 g"^h“ ^ for 4 m m particles at Nagel in Mar c h and 0.45 ym detritus at Upper 43rd in June. On an average 4 and 1 m m particles had lowest CR and 0.45 ym particles highest CR, but particles were mainly similar except for high rates in the 0.45 ym fraction. On a site basis 0.45 ym was highest at all sites, but no inverse trend was indicated in CR and particle size. Sizes ranked from lowest to highest CR were Upper 159 43rd, Kellogg Forest, B Avenue, Smith and Nagel. Detritus CR was higher for all fractions than epilithon on a weight basis. Inorganic sediment CR average 70% the rates of detritus with 4 and 1 mm particles about 35% their detrital counterparts and 250 and 75 ym were nearly equal; a similar, probably bacterial dominated flora, developed on both inorganic and organic substrates. O n epilithon, CR ranged from undetectable for 250 ym sizes at Smith, Nagel,and Upper 43rd to 215 yl02 g _1h“ l for 75 ym sands in June at the Kellogg Forest Site. with particle size. Rates exhibited an inverse relationship Sites ranked Kellogg Forest, Upper 43rd, B Avenue ■ Nagel, and Smith in increasing levels of CR. Epilithon CR values were similar at B Avenue and Nagel Sites, but a different floral composition was evident based on levels of primary productivity; B Avenue was bacterial dominated and Nagel algal dominated. Epilithon NCP had a greater range than detritus (-321 to +2174 yl02 g “ lh“ l) for 75 y m Smith sands to 4 m m Kellogg Forest gravel. Averages for particle-sizes indicated 1 m m to be lowest in NCP and 4 m m highest. All inorganic sediment sizes were colonized by autotrophs at Upper 43rd and the majority at Nagel and Kellogg Forest on an annual basis. Heterotrophic sites had little algal colonization indicated, but 4 and 1 m m fractions exhibited weak autotrophic potentials. Sites ranked Smith, B Avenue, Nagel, Upper 43rd, and Kellogg Forest in average increasing NCP. Since particle-size distribution affects total sediment community metabolism, estimates were calculated on an areal basis. Detrital NCP was generally highest on 4 m m particles with a direct inverse relationship evident between NCP and particle size. On an average 160 site basis 0.45 y m detritus was highest in NCP with no trend following for other particle-sizes. CR averages had an inverse relationship between particle size and CR, which was not evident on a weight basis. Epilithon NCP and CR on an areal basis were dominated by 4 m m stones followed by 250 ym, 1 mm, and 75 y m size. GCP, calculated only on an areal basis, indicated the detrital segement to be in a comsumlng mode with positive values only for 4 m m detritus at B Avenue in the fall, 0.45 ym seasonally, and all segements periodically at Upper 43rd, Nagel and Kellogg Forest. NDM remained below zero for all particle sizes except 0.45 ym in spring and summer at the Kellogg Forest, which indicated algal inputs to the FP0M pool. P/R ratios were all below one except for 0.45 ym particles in spring at Upper 43rd. Epilithon patterns were more complex with GCP from 4 mm, 1 mm, and 75 y m inorganic sediments contributing periodically at Smith; 4 mm annually and all other periodically at B Avenue; 4 and 1 m m and 250 ym sediments generally contributing annually at Upper 43rd; all sizes positive annually at Nagel; and Kellogg Forst had all except 75 ym sizes in a contributing mode annually. NDM was annually below zero for all particles at Smith and B Avenue and above for the majority of particles at the autotrophic sites. P/R ratios were below one for all sizes at heterotrophic sites and above one for most particles at autotrophic sites with exceptions in the smaller particle sizes. Overall, 0.45 ym detritus had the greatest CR and all sizes illustrated little productive capacity on a weight basis, while inorganic sediments had greatest NCP on larger substrates and CR in the smaller fractions. On an areal basis productivity was highest on 161 larger fractions of both detritus and Inorganic sediments; the majority of NCP occurred on the inorganics. CR was highest on smaller detritus and larger inorganics. Sites Smith, a first order site, was heterotrophic in nature, detrital based, dependent mainly on epilithic detritus for organic support with little autochthonous production. the larger substrates. What little GCP took place did so on P/R ratios were less than one, NDM below zero and E / D ratios constant and low (3.7 to 4.8), which Indicated the impact of allochthonous inputs to the riffle community. B Avenue, a slightly larger first order reach, was also heterotrophic dependent on allochthonous detritus trapped in the epilithon, but greater autochthonous activity was evident, which was at times contributing to the community support. Based on NDM and P/R ratios not enough NCP occurred for the riffle to be self-supporting. E /D ratios, averaging 7.9, and ranging from 3.5 to 16.5 indicated greater variability and changes in the epilithon flora compared to Smith Site and illustrated the periodic productive potential of the site. Upper 43rd, an open second order mead o w reach, was annually self-supporting, thus, considered an autotrophic site. Indicating the autotrophic dominance in the epilithon community, P/R ratios were near one and NDM was consistently positive. Epilithon development was high at this site as was CR which continually mediated P /R and NDM. E/D ratios were highest at this site averaging 14.8, which emphasized the importance of the epilithic community to riffle sections. Storage was 162 indicated at this site in the FPOM pool as high rates of NCP occurred as well as increased CR during summer and fall. High rates of NCP at Upper 43rd indicated the effects of lack of canopy at a second order site. The open third order Nagel Site illustrated the effects of clearing on rates of third order riffle NCP as rates approximated those of larger rivers and were significantly higher than the shaded Kellogg Forest Site. P/R and NDM indicated a producing mode to be characteristic of both sites, but at Nagel higher epilithon development occurred and subsequently increased CR, while autotrophic colonization was greater as indicated by increased rates of NCP. Indicating continued autochthonous support P/R ratios were well above one over annual periods and E / D ratios were similar to Upper 43rd at Nagel (x = 12) and intermediary at Kellogg Forest (x ■ 9.5). The shaded Kellogg Forest third order riffle was also autotrophic in nature, lesser in productivity than the open site, but also lesser in CR and epilithon development. This riffle section was self-supporting on an annual basis as indicated by the P /R ratios greater than one and positive NDM. E/D ratios ranging from 4.6 to 15.8 indicated the diverse base of this riffle. T he autotrophic-heterotrophic balance in riffle sections was dependent on the epilithon component both in development, which was m ediated by light availability, and composition which would also be affected by light, and on a species level temperature dictated. LITERATURE CITED LITERATURE CITED Adams, Michael S. and Walter Stone, 1973. 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Estimates of Community Metabolism (g 02/m2), Augusta Creek, Michigan. 24 October 1974 (light), 25 October 1974 (dark), SMITH SITE, STANDARD RON g 0 2 / m 2 .hr NET COMMUNITY PRODUCTIVITY COMMUNITY RES P I R A T I O N -.018 -.034 -.018 -. 195 -.360 -. 193 - 6.059 -11. 16 -5.993 mean se c v (JO -. 023 .005 3 8.300 - . 250 :055 38.300 - 7.736 1.711 3 8.300 g 0 2 / m 2 . hr g 02/rn2.da g 0 2 / m 2 .mo .031 .038 .036 .751 .910 .876 23.287 28.198 27.156 mean se c v (JO .035 .002 9 .870 .846 .048 9 .870 26.214 1.494 9.870 Rep 1 Rep 2 Rep 3 mean se c v (JO g 0 2 / m 2 .rco .013 .004 .018 . 139 .045 . 197 4.304 1.391 6.092 .012 .004 60.400 . 127 .044 60.400 3.929 1.370 60.400 g 02/m 2 . d a Rep 1 Rep 2 Rep 3 g 02/'m2.da -.612 — .865 - .679 -.719 .075 18.178 1 1 1 1 GROSS COMMUNITY 1 PRODUCTION/ 1 24 H R RESPIRATION 1 1 1 1 1 PG/R24 Rep 1 R ep 2 R ep 3 . 185 .049 .224 me a n se c v (JO . 153 .053 60.063 174 Rep 1 R e p .2 Rep 3 mean se c v (JO NET DAILY METABOLISM g 0 2 / m 2 .mo Rep 1 Rep 2 Re p 3 g 02/m 2 . h r GROSS C O M M U N I T Y PRODUCTIVITY g 0 2 / m 2 .da Table A-l (con’t ) . Estimates of Community Metabolism (g 02/ra2), Augusta Creek, Michigan. 15 January 1975 (light). 16 January 1975 (dark), SMITH SITE, STANDARD RUN g 0 2 ' m 2 . hr NET COMMUNITY PRODUCTIVITY GROSS COMMUNITY PRODUCTIVITY - .007 -.008 -.006 - . 064 - . 073 - . 060 -1.988 -2.251 -1.871 me a n se c v (JS) -. 007 .000 9. 557 -.066 .004 9 . 557 -2.037 .112 9.557 02''m2. hr g 0 2 / ib2 .da g 0 2 / m 2 .mo Rep 1 Rep 2 Rep 3 .011 .019 .020 .271 .461 .473 8.407 14.285 14.657 mea n se c v (/•) .017 .003 28.160 .402 .065 28.160 12.450 2.024 28.160 B 02/m2.hr g 02/m 2 . d a g 02/m2.mo Rep 1 Rep 2 Rep 3 .004 .OH .013 .042 . 108 . 125 1.315 3.362 3.888 mean se c v ( ") .010 .003 47.601 .092 .025 47.601 2.855 .785 47.601 g 02/m2.da NET DAILY ME T A B O L I S M g 0 2 / m 2 .mo Rep 1 Rep 2 Rep 3 g COMMUNITY RE S P I R A T I O N g 02.'m2. da Rep 1 Rep 2 Rep 3 — .229 - . 352 - . 347 mean se c v (” ) - . 310 .040 22.602 1 1 1 GROSS COMMUNITY 1 PROD U C T I O N / 1 24 H R R E S P I R A T I O N 1 1 I 1 1 PG/R24 Rep 1 Rep 2 Rep 3 . 157 09 rr .265 mean se c v (75) .219 .032 25.656 Table A-l (con't) . Estimates of Community Metabolism (g 02/m2) , Augusta Creek., Michigan. 20 May 1973 (light), 21 May 1973 (dark), SMITH SITE, STANDARD RDM g 02/m2.hr MET COMMUNITY PRODUCTIVITY COMMUNITY RESPIRATION Rep 1 Rep 2 Rep 3 -.007 - .004 - .010 - . 100 - . 062 -. 143 -3.109 -1. 9 2 0 -4.435 mean se c v (JI) -.007 .002 3 9.875 -. 102 .023 39.875 -3.155 .726 39.875 g 0 2 / m 2 .hr g 02/m2.da g 02/m2.mo Rep 1 Rep 2 Rep 3 .021 .028 .009 .492 .677 .211 15.252 20.981 6.547 me a n se c v (55) .019 .006 50.966 .460 . 135 50.966 14.260 4. 196 50.966 g 02/m2.hr g 02/m 2 . d a g 02/m2.mo Rep 1 Rep 2 Rep 3 .014 .024 -.001 .202 .354 - .013 6 . 264 10.974 - . 412 mean se c v (55) .012 .007 102.00 . 181 . 107 102.00 5.609 3.303 102.00 g 0 2 / m 2 . da NET DAILY METABOLISM Rep 1 Rep 2 R ep 3 - . 290 -.323 -.225 mea n se c v (55) -.279 .029 17.930 1 1 1 GROSS COMMUNITY 1 PROD U C T I O N / 1 24 H R R E S P I R A T I O N 1 1 1 1 1 PG/R24 Rep 1 Rep 2 Rep 3 .411 .523 - . 063 mean se c v (5?) .290 . 180 107.13 176 G R OS S C O M M U N I T Y PRODUCTIVITY g 02/m2.tno g 02/m2.da Table A-l (cos’t). Estimates of Community Metabolism (g 0 2 / m 2 ) , Augusta Creek. Michigan. 17 July 1975 (light), 18 July 1975 (dark), SMITH SITE, STANDARD RUN g 0 2 / 1112.hr NET COMMUNITY PRODUCTIVITY COMMUNITY RESPIRATION G R OS S C O M M U N I T Y PRODUCTIVITY g 02/m2.mo Rep 1 Rep 2 Rep 3 -.019 -.012 -.015 -.287 -. 175 -.225 -8.886 -5.415 -6.989 mea n se c v (J5) -.015 .002 24.492 -.229 .032 24 . 4 9 2 - 7.097 1.004 2 4.492 g 02/m2.hr g 02/m 2 . d a g 02/m2. mo Rep 1 Rep 2 Rep 3 .028 .016 .023 .662 .379 .564 20.534 11.755 17.484 me a n se c v ( 3) .022 .003 2 6 .865 .535 .083 26.865 16.591 2.573 26.865 g 0 2 / m 2 .hr g 0 2 / m 2 .da g 0 2 / m 2 .mo Rep 1 Rep 2 Rep 3 .008 .004 .008 . 125 .661 . 125 3.888 1.898 3.888 mean se c v (75) .007 .001 35.635 . 104 .021 35.636 3.224 .663 35.635 g 02/012.43 NET DAILY METABOLISM g 0 2 / m 2 . da Rep 1 Rep 2 Rep 3 - . 537 -.318 -.439 me a n se c v (R) -.431 .063 25 . 4 3 8 1 1 1 GROSS COMMUNITY 1 P R O DUCTION/ 1 24 H R R E S P I R A T I O N I 1 1 1 1 PG/R24 Rep 1 Rep 2 Re p 3 . 189 . 161 .222 me a n se c v C55) . 191 .018 15.985 Table A-l. Estimates of Community Metabolism (g 02/ro2) , Augusta Creek., Michigan. 26 January 1973 (light), 25 January 1973 (dark), B AVENUE. STANDARD RUN RET COMMUNITY PRODUCTIVITY 1 -. 002 -.023 -.728 mean se c v (75) -. 002 - . 023 -.728 Rep g 02/m2. hr COMMUNITY RESPIRATION .011 .274 8.482 mean se c v (75) .011 .274 8. 482 178 Rep 1 g 02/m 2 . d a g 0 2 / m 2 .mo 1 .009 .088 2.729 me a n se c v ( 75) .009 .088 2.729 Rep g 02/tn2.da NET DAILY ME T A B O L I S M g 0 2 / m 2 .mo g 02/m2.da g 02/m2. hr G ROSS C O M M U N I T Y PRODUCTIVITY g 0 2 / m 2 .mo g 02/m 2 . d a g 02/m2. hr Rep 1 -. 186 mean se c v ( 75) -.186 PG/R24 1 1 GROSS COMMUNITY 1 PRODUCTION/ ! 24 H R R E S P I R A T I O N 1 1 Rep 1 .322 mean se c v ( 75) .322 Table A-l (con’t). Estimates of Community Metabolism (g 0 2 /m2). Augusta Creek, Michigan. 0? June 1973 (light), 96 June 1973 (dark), B AVENUE, STANDARD RUN g 0 2 / m 2 .hr NET COMMUNITY PRODUCTIVITY COMMUNITY RESPIRATION GROSS COMMUNITY PRODUCTIVITY g 02/m 2 . d a g 02/m2.mo Rep 1 R ep 2 .016 .003 .247 .046 7 .42 3 1.366 m ea n se c v (J5) .010 .007 9 7.436 . 146 . 101 9 7.456 4.395 3.028 97.456 g 02/tn2.hr g 02/m 2 . d a g 0 2 / m 2 .mo Rep 1 Rep 2 .037 .021 .890 .511 26.712 15.336 mean se c v (!?) .029 .008 38.261 .701 . 190 38.261 21.024 5.688 38.261 g 02/m2.hr g 0 2 / m 2 .da g 0 2 / m 2 . mo Rep 1 Rep 2 .053 .024 .811 .369 24. 3 1 8 11 .066 mean se c v (T5) .039 .015 52.965 .590 17.692 6.626 52.965 on , 52.965 g 9 2 / m 2 .da 1 P G/R24 mea n se c v (J5) © Rep 1 Rep 2 1 NET DAILY METABOLISM b 0) 1 -.142 -.111 .031 39.797 1 GROSS COMMUNITY I PROD U C T I O N / 1 24 H R R E S P I R A T I O N 1 1 1 Rep 1 Rep 2 .910 .722 mean se c v (” ) .816 .094 16.361 Table A-l (con't). Estimates of Community Meta b o 1 Ism (g 02/m2), Augusta Creek, Michigan. 05 July 1973 (light),.04 July 1973 (dark), B AVENUE, STANDARD RUN g 02/m 2 . h r NET COMMUNITY PRODUCTIVITY COMMUNITY RE S P I R A T I O N Rep 1 Rep 2 -.007 -.016 -. 106 -.244 -3.292 -7.571 mean se c v ( 75) -.012 .005 55.711 -.175 .069 55.711 -5.432 2. 140 55.711 g 0 2 / m 2 .hr g 02/m2.da g 02/m2.mo Rep 1 Rep 2 .039 .021 .924 .492 28.644 15.252 mea n se c v (75) .030 .009 43.145 .708 .216 43.146 21 . 9 4 8 6.696 43.146 g 0 2 / m 2 .hr g 02/m2.da g 02/m2. mo Rep 1 Rep 2 .031 .004 .478 .067 14.814 2.069 mean se c v (75) .018 .014 106.76 .272 .206 106.76 8.441 6.372 106.76 g 0 2 / m 2 .da NET DAILY METABOLISM g 02/m2.mo Rep 1 Rep 2 — .446 -.425 mean se c v (75) - . 436 .01© 3.376 1 1 1 GROSS COMMUNITY 1 PRODU C T I O N / 1 24 H R R E S P I R A T I O N 1 1 1 PG/R24 Rep 1 Rep 2 .517 . 136 mean se c v (71) .326 . 191 82.635 180 GR OS S C O M M U N I T Y PRODUCTIVITY g 02/m2.da Table A— 1 (con’t). Estimates of Community Metabolism (g 02/ra2), Augusta Creek, Michigan. 02 August 1973 (light), 91 August 1973 (dark). B AVENUE, STANDARD RUN g 0 2 / m 2 .hr NET COMMUNITY PRODUCTIVITY .006 .006 .032 .032 2.548 2.548 mean se .006 .082 2.548 (n) g 02/tti2.hr g 0 2 / m 2 .mo Rep 1 Re p 2 .025 .037 .600 .893 18.600 27.677 mean se c v (3) .031 .006 27.739 .746 . 146 27.739 23.138 4.538 27.739 g 02/m2.hr g 0 2 / m 2 . da g 0 2 / m 2 .mo Rep 1 Rep 2 .031 .043 .443 .619 13.724 19.177 mean se c v ( 3) .037 .006 23.442 .531 .068 23.442 16.450 2.727 23.442 g 0 2 / m 2 .<1a NET DAILY METABOLISM g 02/mC.da Rep 1 Rep 2 -. 157 -.274 mean se c v (3) -.216 .058 38.313 1 I 1 1 GROSS COMMUNITY 1 P R O DUCTION/ 1 24 H R R E S P I R A T I O N 1 1 1 PG/R24 Rep 1 Rep 2 mean se c v (3) .733 .693 .715 .022 4 . 433 181 GROSS COMMUNITY PR O D U C T I V I T Y g 02/m2.mo Rep 1 Rep 2 CV COMMUNITY RESPIRATION g 0 2 / m 2 .da Table A-l (con’t). Estimates of Community Metabolism (g 02/m2), Augusta Creek, Michigan. 29 August 1973 (light), 39 August 1973 (dark), B AVENUE, STANDARD RUN g 02/tn2.lir N E T COMMUNITY PRODUCTIVITY COMMUNITY RESPIRATION CROSS C O M M U N I T Y PRODUCTIVITY Rep 1 Rep 2 - . 004 - . 012 -.053 -. 166 -1.649 -5.154 mean se c v (75) -.008 .004 72.853 110 .057 7 2 .853 -3.401 1.752 72.853 g 0 2 / m 2 .hr g 02/m2.da g 0 2 / m 2 .mo Rep 1 Rep 2 .018 .031 .439 .742 13.615 22.990 me a n se c v ( 75) .025 .006 36.218 .590 . 151 36.218 18.302 4.687 36.218 g 0 2 / m 2 . hr g 0 2 / m 2 . da g 02/m2.mo Rep 1 Rep 2 me a n se c v (3) .014 .018 . 190 .245 5.896 7.586 .016 .002 17.732 .217 .027 17.732 6.741 .845 17.732 g 02/m2.da NET DAILY METABOLISM g 0 2 / m 2 .mo g 02/m2.da Rep 1 Rep 2 -.249 - .497 mean se c v (75) -.373 . 124 47.001 1 1 1 GROSS COMMUNITY 1 PROD U C T I O N / 1 24 H R R E S P I R A T I O N 1 1 1 PG/R24 Rep 1 Rep 2 .433 .330 mean se c v (75) .382 .052 19.091 Table A-l (con’t). Estimates of Comnmnlty Metabolism (g 02/tn2) , Augusta Creek., M ichigan. 27 September 1973 (light), 26 September 1973 (dark), B AVENUE, STANDARD RUN g 0 2 / m 2 . hr NET COMMUNITY PRODUCTIVITY COMMUNITY RES P I R A T I O N Rep 1 Rep 2 -.011 -.014 -. 131 -. 167 -3 . 9 4 4 -5.019 mean se c v ( %) -.013 .001 16.971 -. 149 .018 16.970 -4.481 .538 16.971 g 0 2 / m 2 . hr g 0 2 / m 2 .da g 0 2 / m 2 .mo Rep 1 Rep 2 .032 .039 .775 .941 23.256 28.224 mean se c v ( 55> .036 .003 13.643 .858 .083 13.648 25.740 2.484 13.648 g 02/m2. hr g 02/tn2.da g 0 2 / m 2 .mo Rep 1 Rep 2 mea n se c v (a) .021 .023 .255 .301 7.636 9.034 .023 .002 11.361 .278 .023 11.861 8.335 .699 11.861 g 0 2 / m 2 .da NET DAILY METABOLISM g 0 2 / m 2 •mo Rep I Rep 2 me a n se c v <3) -.521 -.640 -.580 .059 14.503 1 1 1 GROSS COMMUNITY 1 PROD U C T I O N / 1 24 H R R E S P I R A T I O N 1 1 1 PG/R24 Rep 1 Rep 2 .328 .320 mean se c v (T.) .324 .084 1.788 183 C ROSS C O M M U N I T Y PRODUCTIVITY g 0 2 / m 2 .da Table A-l (con’t). Estimates of Community Metabolism (g 0 2 / m 2 ) , Augusta Creek. Michigan. 05 November 1973 (light), 06 November 1973 (dark), B AVENUE, STANDARD RUN g 02/m2- hr NET COMMUNITY PRODUCTIVITY COMMUNITY RE S P I R A T I O N -.005 - .004 -.054 - .039 -1.619 -1. 161 mean se c v (55) -.005 .001 23.311 -.046 .008 23.311 - 1.390 .229 23.311 g 02/m2.hr g 02/m2.da g 02/ffl2.mo Rep 1 Rep 2 .015 .014 .358 .326 10.728 9 . 792 mea n se c v (55) .014 .001 6.451 .342 .016 6.451 10.260 .468 6.451 g 0 2 / 012 .da g 02/m2.mo Rep 1 R ep 2 .010 .010 .098 . 100 2 .932 2 .993 mea n se c v ( 55) .010 .000 1.459 .099 .001 1.457 2.962 .031 1.456 g 0 2 / m 2 .da NET DAILY METABOLISM g 0 2 / m 2 .mo Rep 1 Rep 2 g 0 2 / m 2 .hr G ROSS C O M M U N I T Y PRODUCTIVITY g 02/m2.da R ep 1 Rep 2 - . 260 — .2°7 mea n se c v (55) -.243 .017 9.680 1 I 1 1 GROSS COMMUNITY 1 PROD U C T I O N / 1 24 H R R E S P I R A T I O N 1 1 1 PG/R24 Rep 1 Rep 2 .273 .306 mean se c v (55) .289 .016 7.891 Table A-l (con't). Estimates of Community Metabolism (g 02/m2), Augusta Creek, Michigan. 11 December 1973 (light), 12 December 1973 (dark), B AVENUE, STANDARD RUN g 0 2 / m 2 .hr NET COMMUNITY PRODUCTIVITY COMMUNITY RESPIRATION Rep 1 Rep 2 -.019 - . 004 -. 172 - . 032 -5.333 - . 9 93 mea n se c v (7?) -.011 .008 9 7 .029 -. 102 .070 9 7 .029 -3.163 2. 170 97.029 g Q2/tn2.hr g 02/m 2 . d a g 02/ra2.mo Rep 1 Rep 2 .011 .009 .266 .214 8.258 6.622 mean se c v (7?) .010 .001 15.556 .240 .026 15.556 7.440 .818 15.556 g 02-'m2. hr g 02/m2.da B 02/m2.mo Rep 1 Rep 2 -.008 .005 - .070 .049 -2.184 1.532 mean se c v ( 71) -.001 .007 805.49 -.011 .060 805.49 -.326 1.858 805.49 g 02/rr.2.da NET DAILY METABOLISM g 02/m2.mo Rep 1 Rep 2 -.337 -. 164 mean se c v (71) -.251 .086 4 8.740 1 1 1 GROSS COMMUNITY 1 PROD U C T I O N / 1 24 E R R E S P I R A T I O N 1 1 1 PG/R24 Rep 1 Rep 2 -.265 .231 mean se c v (75) -.017 .248 2111.9 185 GROS S C O M M U N I T Y PRODUCTIVITY g 02/m 2 . d a Table A-l (con’t). Estimates of Community Metabolism (g 02/m 2 > . Augusta Creek, Michigan. 23 January 1974 (light). 22 January 1974 (dark), B AVENUE, STANDARD RUN g 02/m2 . h r NET COMMUNITY PRODUCTIVITY COMMUNITY RESPIRATION g 02/m2.da Rep 1 Rep 2 .000 .009 .002 .000 .060 .000 mean se c v (n> .000 .000 141.42 .0 0 1 .001 141.42 .030 .030 141.42 g 0 2 / m 2 . hr g 02/m2.da g 02/m2.mo Rep 1 Rep 2 .011 .012 .259 .298 8.035 9.226 mea n se c v (J5) .012 .001 9 . 753 .278 .019 9 . 753 8.630 .595 9.753 g 02/ra2.hr CROSS COMMUNITY PRODUCTIVITY Rep 1 Rep 2 me a n se c v (") Rep I Rep 2 mean se cv ( g 02/m2.da g 02/ra2.mo .012 . 106 . 120 3.301 3.721 .012 .001 8.461 . 113 .007 8.461 3.511 .210 8.461 .011 B 02/rn2. .008 .007 119.80 . 115 .097 119.80 3 .553 3.010 119.80 g 02/m2. hr B 02/m2.da B 0 2 / m 2 .mo Rep 1 Rep 2 .025 .017 .598 .410 18.526 12.722 .021 .004 26 . 2 6 4 .504 .094 2 6.264 15.624 2.902 26.264 g 0 2 / m 2 .hr B 02/nt2.da B 0 2 / m 2 .mo Rep 1 Rep 2 mean se c v IN) 3 NET DAILY METABOLISM B 0 2 / m 2 .mo Rep 1 Rep 2 mean se c v (%> GROSS COMMUNITY PRODUCTIVITY g 02/m2.da Rep 1 Rep 2 me a n se c v (%> .026 .032 .381 .461 11.313 14.302 .029 .003 13.480 .421 .040 13.480 13.058 1.245 13.480 G 2 / m 2 .da - . 217 .051 - . 083 . 134 22 8 . 5 8 1 1 i GROSS COMMUNITY 1 PROD U C T I O N / 1 24 H R R E S P I R A T I O N 1 1 1 PC/R24 Rep 1 Rep 2 .638 1. 124 me a n se c v , Augnsta Creek, Michigan. 18 December 1973 (light), 19 December 1973 (dark), NAGEL SITE, STANDARD RUN g 0 2 / m 2 .hr NET COMMUNITY PRODUCTIVITY COMMUNITY RESPIRATION Rep 1 Rep 2 . 139 .076 1.262 .690 39.127 21.383 mean se c v (J5) . 107 .031 41.470 .976 .286 41.470 30.255 8.872 41.470 g 0 2 / m 2 .hr g 0 2 / m 2 . da g 0 2 / m 2 .mo Rep 1 Rep 2 .036 .025 .876 .588 27.156 18.228 mean se c v (3) .031 .006 27.821 .732 . 144 2 7.820 22.692 4.464 27.821 g 0 2 / m 2 . hr g 0 2 / m 2 .da g 02/tn2.mo Rep 1 Rep 2 . 175 . 100 1.594 .913 49.424 28.295 mean se c v (53) . 138 .037 38.448 1.254 .341 38.448 38.859 10.565 3 8.448 S 02/m2.da NET DAILY METABOLISM Rep 1 Rep 2 mean se c v (3) .718 .325 .521 . 197 53.369 1 1 1 1 CROSS COMMUNITY 1 PRODUCTION/ 1 24 H R R E S P I R A T I O N 1 1 1 PG/R24 Rep 1 Rep 2 mean se c v (3) 1.820 1.552 1.686 . 134 11.226 202 GROSS COMMUNITY PRODUCTIVITY g 0 2 / m 2 .mo g 0 2 / m 2 .da Table A— 1 (con’t). Estimates of Community Metabolism (g 02/m2), Augusta Creek, Michigan. 20 March 1974 (light)., 21 March 1974 (dark), NAGEL SITE, STANDARD RUN g 02/m2 . h r NET COMMUNITY PRODUCTIVITY COMMUNITY RES P I R A T I O N Rep 1 Rep 2 . 177 . 141 2. 142 1.705 66.407 52.870 m ean se c v (55) . 159 .010 16.050 1.924 .218 16.050 59.638 6.769 16.050 g 0 2 / m 2 .hr g 02/m2.da g 0 2 / m 2 .mo Rep 1 Rep 2 .073 .054 1.762 1.289 54.610 39.953 mean se c v (55) .064 .010 21.920 1.525 .236 21.920 47.281 7.328 21.920 g 02/re2.hr g 0 2 / m 2 .da g 02/m2.rao Rep 1 R ep 2 .250 . 194 3 .032 2 .357 94.007 73 . 0 6 3 mean se c v (55) *222 .028 17.729 2.695 .338 17.729 83.535 10.472 17.729 S 02/m2.da NET DAILY METABOLISM g 0 2 / m 2 . mo Rep 1 Rep 2 1.271 1.063 mea n se c v (55) 1. 170 . 101 12.262 1 1 f GPvOSS C O M M U N I T Y t PRODUCTION/ I 24 H R R E S P I R A T I O N t 1 1 PG/R24 Rep 1 R ep 2 1.721 1.829 mean se c v (55) 1.775 .054 4.274 203 G R OSS C O M M U N I T Y PRODUCTIVITY g 0 2 / m 2 . da Table A- 1 (con’t). Estimates of Community Metabolism (g 0 2 /m2), Augusta Creek, Michigan. 23 April 1974 (light), 22 April 1974 (dark), BAGEL SITE, STANDARD RUN g 02-'m2. hr NET COMMUNITY PRODUCTIVITY COMMUNITY RESPIRATION g 0 2 / m 2 .no g 0 2 / te2 .da Re p 1 Rep 2 . 155 .082 2. 122 1. 122 63.675 33.669 mean se c v (B) . 118 .036 43.592 1.622 .500 43.592 48.672 15.003 43.592 g 0 2 / m 2 .hr g 0 2 / m 2 . da g 02/ra2.mo Rep I Rep 2 . 127 . 151 3.038 3.634 91.152 109.01 mea n se c v (75) . 139 .012 12.616 3.336 .298 12.616 100.08 8.928 12.616 -p' g 02/m2.hr GR OS S C O M M U N I T Y PRODUCTIVITY g 02/ra2.mo Rep 1 Rep 2 .281 .233 3.859 3.200 115.78 95.985 me a n se c v (15) .257 .024 13.221 3.529 .330 13.221 105.88 9.899 13.221 g 02/m2.da NET DAILY METABOLISM g 0 2 / m 2 .(la Rep 1 Rep 2 .821 - . 434 mean se c v (75) . 193 .628 45 8 . 7 7 1 1 1 GROSS COMMUNITY 1 PROD U C T I O N / 1 24 H R R E S P I R A T I O N 1 1 1 PC/R24 Re p 1 Re p 2 1.270 .880 mean se c v (75) 1.075 . 195 25.625 T able A — 1 ( c o n ’t ) . Est imates of C o m m u n i t y M e t a b o l i s m (g 0 2 Z m 2 ) , A ugusta Creek, Michigan, 2 4 April 1974 ( light)., 2 2 April 1974 (dark), N A G E L SITE, S T A N D A R D RUN g 02Zm 2 . h r WET COMMONITY PRODUCTIVITY COMMUNITY RE S P I R A T I O N Re p 1 Rep 2 .251 . 124 3 . 454 1.713 103.61 51.390 mean se c v (JS) . 188 .063 47.643 2.583 .870 47.643 77. 4 9 8 26. 1 0 8 47.643 g 02Zm2.hr g 02Zm2.da g 0 2 Z m 2 .mo Rep 1 Rep 2 . 127 . 151 3.038 3.634 91.152 109.01 mean se c v (55) . 139 .012 12.616 3 .336 .298 12.616 100.08 8.928 12.616 g 02Zin2.hr g 02Zm2.da g 02Zm2.mo Rep 1 Rep 2 m ean se c v (3) .377 .276 5. 197 3.798 155.90 113.93 .327 .05 1 2 1.997 4 . 497 .700 2 1.997 134.92 20.985 21.997 g 9 2 Z n 2 .da NET DAILY METABOLISM Rep 1 Rep 2 2. 158 . 164 mea n se c v (3) 1. 161 .997 121.43 1 I 1 GROSS COMMUNITY 1 PROD U C T I O N Z 1 24 H R R E S P I R A T I O N 1 1 1 P GZR24 Rep 1 R ep 2 mean se c v (3) 1.710 1.045 1.378 .333 34.139 205 G ROS S C O M M U N I T Y PRODUCTIVITY g 0 2 Z m 2 .mo g 0 2 Z m 2 .da Table A-l (can’t). Estimates of Community Metabolism (g 0 2 /m2), Augusta Creek, Michigan. 29 May 1974 (light), 29 May 1974 (dark), NAGEL SITE. DIEL RON g 0 2 / m 2 . hr NET COMMUNITY PRODUCTIVITY COMMUNITY RESPIRATION . 128 . 173 1.915 2 . 600 59.381 80.585 m ea n se c v ( 75) . 150 .023 2 1.425 2.258 .342 2 1.425 69.982 10.602 21.425 g 02/m2.hr g 0 2 / 1112.da g 0 2 / m 2 .mo Rep 1 Rep 2 . 113 . 119 2.719 2.868 84.295 88.908 mea n se c v (75) . 116 .003 3 .766 2.794 .074 3.767 86.602 2.306 3.766 g 0 2 / m 2 . da g 0 2 / m 2 .mo Rep 1 Rep 2 .241 .293 3 .615 4.392 112.06 136.15 mean se c v (!5) .267 .026 13.724 4.003 .389 13.724 124.11 12.043 13.723 S 02/m2.da NET DAILY METABOLISM g 02/m2.mo Rep 1 Rep 2 g 02/tn2.hr GROSS COMMUNITY PRODUCTIVITY g 0 2 / m 2 . da Rep 1 Rep 2 .896 1.524 mean se c v (75) 1.210 .314 36.714 1 1 1 GROSS COMMUNITY 1 P R O D U C T 1011/ 1 24 H R R E S P I R A T I O N 1 1 1 P G/R24 Rep 1 Rep 2 1.329 1.531 mean se c v ( 75) 1.430 . 101 9.986 Table A - 1 (con’t). Estimates of Community Metabolism (g 02/ic2) ■ Augusta Creek, Michigan. 01 July 1974 (light), 01 July 1974 (dark), NAGEL SITE, DIEL RON g 02/tn2.hr NET COMMUNITY PRODUCTIVITY Rep . 133 2.032 62.982 me a n se c v (71) . 133 2.032 62. 9 8 2 Rep 3.672 113.83 mean se c v (J5) . 153 3.672 113.83 207 . 153 g 0 2 / 102 .da g 02/m2.mo 1 .286 4. 362 135.22 mean se c v (J5> .286 4.362 135.22 Rep 3 NET DAILY METABOLISM g 0 2 / m 2 .mo g 0 2 / 102 .da 1 S 0 2 / m 2 .hr GROSS COMMUNITY PRODUCTIVITY g 02/m2.rno 1 g 0 2 / m 2 .hr COMMUNITY RE S P I R A T I O N g 02/m2. da Rep 02/ra2.da 1 .690 mean se c v (/•) .690 1 1 1 GROSS COMMUNITY 1 PRODU C T I O N / 1 24 H R R E S P I R A T I O N PG/F24 Rep 1 1.188 me a n se c v <%) 1. 188 Table A - 1 ( c o n ’t). E s t i m a t e s of C o m m u n i t y M e t a b o l i s m (g 0 2 / m 2 ) , Augusta Creek, Michigan. 05 August 1974 (light), 05 August 1974 (dark), N A G E L SITE, S T A N D A R D R O N g 0 2 / m 2 .hr NET COMMUNITY PRODUCTIVITY COMMUNITY RES P I R A T I O N g 02/m 2 . d a Rep 1 Rep 2 Rep 3 .251 . 163 3.564 2.332 111.09 72.302 m ea n se CV (55) .207 .044 29.911 2.958 :626 29.911 91.697 19.394 29.911 g 0 2 / m 2 .hr g 0 2 / m 2 .da B 0 2 / m 2 .mo .094 2.249 69.713 .091 2. 179 67.555 mean se c v (5J) .092 .001 2. 224 2 . 214 .035 2. 223 68.634 1.079 2.223 8 0 2 / m 2 .mo Rep 1 Rep 2 Rep 3 .344 4 .923 152.63 mea n se c v (55) .344 4 .923 152.63 g 0 2 / m 2 .(1a NET DAILY METABOLISM g 02/m 2 . d a Rep 1 Rep 2 Rep 3 2.675 mean se c v (55) 2.673 1 1 GROSS COMMUNITY 1 PROD U C T I O N / 1 24 H R RESPIRATION 1 1 PG/H24 Rep 1 Rep 2 Re p 3 2. 189 mean 2. 189 1 se 1 c v ( 55) 208 Rep 1 Rep 2 Rep 3 g 02/m 2 . h r GROSS COMMUNITY PRODUCTIVITY B 02/m2.mo Table A - 1 (con1t)• Estimates of Community Metabolism (g 02/m2>, Augusta Creek, Michigan. 06 November 1974 (light), 07 November 1974 (dark), NAGEL SITE, STANDARD RUN g 02/m 2 . h r NET COMMUNITY PRODUCTIVITY COMMUNITY RESPIRATION Rep 1 Rep 2 Rep 3 . 121 . 197 .083 1.225 1.991 .844 36.742 59.716 25.315 mean se c v (71) . 134 .033 43.164 1.353 .337 43.164 40.591 10.116 43.164 g 0 2 / m 2 .hr g 0 2 / m 2 . da g 02/m2. mo Rep 1 Rep 2 Rep 3 .031 .066 .027 .744 1.572 .660 2 2.320 47.160 19.800 mean se c v (75) .041 .012 50.811 .992 .291 50.811 29.760 8.730 50.811 g 0 2 / m 2 .hr g 02/m2.da g 02/m2.rno Rep 1 Rep 2 Rep 3 . 152 .262 .111 1.539 2.654 1. 122 46.162 79.622 33.672 mean se c v (75) . 175 .045 44 . 7 0 0 1.772 .457 4 4 .700 53.152 13.717 44.700 g 02/rn2.da NET DAILY METAEOLISM g 02/m2.mo Rep 1 Rep 2 Re p 3 .795 1.082 .462 nMs«*n se c v (75) .780 . 179 39.773 1 1 1 GROSS COMMUNITY 1 PRODUCTION/ 1 24 H R RESPIRATION 1 1 1 1 t PG/R24 Rep 1 Rep 2 Rep 3 2.068 1.688 1.701 mean se c v (75) 1.819 . 125 11.868 209 GR O S S C O M M U N I T Y PRODUCTIVITY g 02/m 2 . d a Table A-l (con’t). Estimates of Community Metabolism (g 02/tn2) , A u g u s t a Creek. Michigan. 30 January 1975 (light). 31 January 1975 (dark), NAGEL SITE, STANDARD RON g 0 2 / m 2 .h r ' NET COMMUNITY PRODUCTIVITY COMMUNITY RESPIRATION Rep 1 Rep 2 Rep 3 .021 .032 .028 .212 .315 .278 6 . 557 9.758 8.619 mean se c v ( 75) .027 .003 19.525 .268 .030 19.524 8.311 .937 19.525 g 0 2 / m 2 . hr g 0 2 /1 1 1 2 . da g 0 2 / m 2 .mo Rep 1 Rep 2 Rep 3 .031 .023 .024 .746 .554 .586 23.138 17. 186 18.154 mean se c v (75) .026 .002 16.385 .629 .059 16.385 19.493 1.844 16.385 g 0 2 / m 2 . hr g 02/m2.da g 02/m2.mo Rep 1 Rep 2 R ep 3 .052 .055 .052 .520 .544 .520 16.130 16.869 16.130 mean se c v ( 75) .053 .001 2.605 .528 .008 2 . 604 16.377 .246 2 .604 g 0 2 / m 2 . da NET DAILY METABOLISM Rep 1 Rep 2 Rep 3 -.226 - . 010 -.065 mean se c v (75) -. lOl .065 111.58 1 1 i 1 GROSS COMMUNITY 1 PRODUCTION/ 1 24 H R R E S P I R A T I O N i 1 1 1 1 PG/ R 2 4 Rep 1 Rep 2 R ep 3 .697 .982 .838 mean se c v (75) .856 .084 16.946 210 GROSS COMMUNITY PRODUCTIVITY g 02/m2.mo g 02/m2.da Table A— 1 (con’t) . Estimates of C o m m u n i t y Metabolism (g 02/m2) , Augusta Creek., Michigan. 31 January 1974 (light), 01 February 1974 (dark), NAGEL SITE. STANDARD RUN g 02/m2.hr NET COMMUNITY PRODUCTIVITY COMMUNITY RESPIRATION Rep 1 Rep 2 .057 .027 .573 .265 17.772 8.221 mean se c v (JJ) .042 .015 51.961 .419 . 154 51.961 12.996 4.775 51.961 g 0 2 / m 2 . hr g 02/m2.da g 0 2 / m 2 .mo Rep 1 Rep 2 .021 .027 .504 .660 15.624 20.460 mean se c v (JJ) .024 .003 18.953 .582 .078 18.953 18.042 2 .418 18.954 g 0 2 / m 2 .hr g 02/n2.da g 0 2 / m 2 .mo Rep 1 Rep 2 .079 .054 .783 .539 24 .262 16.721 me a n se c v (Jl) .066 .012 26.023 .661 . 122 2 6.023 20.491 3.771 2 6.023 g 02/m2.da NET DAILY METABOLISM Rep 1 Rep 2 .279 -. 121 m ea n se c v ( 75) .079 .200 357.31 1 I 1 GROSS COMMUNITY 1 PROD U C T I O N / 1 24 E R RESP I R A T I O N 1 1 1 PG/R24 Rep 1 Rep 2 mean se c v (3) 1.553 .817 1. 185 .368 43.898 211 GROSS COMMUNITY PRODUCTIVITY g 0 2 / m 2 .mo g 02/m2.da PLEASE NOTE: Page 212 missing in numbering only. Text follows. Filmed as received. UNIVERSITY MICROFILMS Table A— 1 (con’t) . Estimates of Community Metabolism (g 02/m2) . Augusta Creek, Michigan. 24 July 1975 (light), 25 July 1975 (dark), NAGEL SITE, STANDARD RUN g 02/m2.da g 0 2 / m 2 .hr NET COMMUNITY PRODUCTIVITY COMMUNITY RESPIRATION Re p 1 Rep 2 Rep 3 . 109 .052 . 106 1.606 .759 1.557 49.773 23.516 48.266 mean se c v (55) .089 .019 36.387 1.307 .275 36.357 40.518 8.512 3 6.387 g 0 2 / m 2 .hr g 0 2 / m 2 .da g 02/m2.mo Rep 1 Rep 2 Rep 3 . 130 . 113 .098 3. 115 2.722 2.342 96.571 84.370 72.614 mean se c v (55) . 114 .009 14.173 2 . 726 .223 14. 173 84.518 6.916 14.173 g 02/m2.hr g 02/m 2 . d a g 0 2 / m 2 .mo Re p 1 Rep 2 Rep 3 .239 . 165 .203 3.518 2.429 2.995 109.04 75.298 92.833 mean se c v ( 55) .292 .021 18.267 2 .980 .314 18.266 92.391 9 . 7 44 18.266 g 0 2 / m 2 . da NET DAILY METABOLISM Rep 1 Rep 2 Rep 3 .402 - .293 .652 I I 1 GROSS COMMUNITY 1 PRODUCTION/ 1 24 H R R E S P I R A T I O N PG/R24 Rep 1 Rep 2 Rep 3 I. 129 .893 1.278 me a n se 1.100 . 112 /a a 1 1 mean se ___ €W7\ .254 .283 «r\r% "9/t 1 1 1 ___ t *7\ « 213 GROSS COMMUNITY PRODUCTIVITY g 0 2 / m 2 .mo Table A - 1 (con’t). Estimates of Community Metabolism (g 0 2 /m2), Augusta Creek., Michigan. 26 November 1974 (light), 27 November 1974 (dark), KELLOGG FOREST, STANDARD RUN g 02/m 2 . h r NET COMMUNITY PRODUCTIVITY COMMUNITY RESPIRATION .061 . 133 .046 .576 1.257 .430 17.293 37.705 12.899 mea n se c v (55) .080 .027 58.486 .754 .255 58.486 22.633 7 .642 58.486 g 02-'m2. hr g 0 2 / m 2 .da g 0 2 / m 2 .mo Rep 1 Rep 2 Rep 3 .470 .610 .252 14.112 18.238 7 .560 .019 .004 40.598 .444 . 104 40.598 13.320 3. 122 40 . 5 9 8 g 02/w 2 . h r g 02/m2.da g C 2 / m 2 .mo Rep 1 Rep 2 Rep 3 .081 . 158 .056 .762 1.497 .529 22.850 44.906 15.876 me a n se c v (3) .098 .031 54.360 .929 .292 5 4 .360 27 . 8 7 8 8.749 54.360 Rep 1 Rep 2 Rep 3 .291 .887 .277 mean se c v ( 3) .485 .201 71.763 1 1 1 GROSS COMMUNITY 1 PROD U C T I O N / 1 24 H R R E S P I R A T I O N 1 1 1 1 P G/R24 Rep 1 Rep 2 Rep 3 1.619 2.456 2. 100 mean se c v (3) 2.058 .242 2 0.392 214 .020 .025 .011 g 02/ra2.da RET DAILY METABOLISM g 0 2 / 1112.1110 Rep 1 Rep 2 Rep 3 mea n se c v (3) GROSS COMMUNITY PRODUCTIVITY g 02/m2.da Table A— 1 (con't). Estimates of Community Metabolism (g 0 2 /m2), Angusta Creels., Michigan. 06 Febrnary 1973 (light!, 05 February 1975 (dark), KELLOGG FOREST, STANDARD RON g 0 2 / m 2 .hr RET COMMUNITY PRODUCTIVITY COMMUNITY RESPIRATION g 02/m2.da “ 777777” Rep 1 Rep 2 Rep 3 !©34 .042 !346 .426 9! 682 11.938 mean se c v i%y .038 .004 14.759 .386 .040 14.759 10.810 1. 128 14.758 g 02/m2.hr g 0 2 / 0 2 .da g 02/m2.m0 Rep 1 Re p 2 Rep 3 .013 .014 .013 .302 .338 .317 8.467 9.475 8 . 8 70 mean se c v (J5) .013 .000 5.676 .319 .010 5.677 8.938 .293 5.677 g 0 2 / m 2 .hr GROSS COMMUNITY PRODUCTIVITY Re p 1 Rep 2 Rep 3 mean se cv < rs> g 0 2 / m 2 .mo g 0 2 / m 2 . da 777777“ 777777 .048 .055 ’ !490 .561 13.709 15.708 .052 .003 9.611 .525 .036 9.611 14.708 1.000 9.611 g 0 2 / m 2 .da RET DAILY METABOLISM g 02/m2.mo Rep 1 Rep 2 Rep 3 . 151 .244 mea n se c v (") . 198 .047 33.263 1I 1 GROSS COMMUNITY 1 PRODU C T I O N / 1 24 H R R E S P I R A T I O N | 1 1 1 1 PG/R24 Rep 1 Rep 2 Rep 3 1.447 1.771 me a n se c v (?I) 1.609 . 162 14.241 Table A - 1 (con11) . Estimates of Community Metabolism (g 02/m2), Augusta Creek, Michigan, 13 May 1975 (light), 14 May 1975 (dark), KELLOGG FOREST, STANDARD RUN g 0 2 / m 2 .hr NET COMMUNITY PRODUCTIVITY COMMUNITY RESPIRATION Rep 1 Rep 2 Rep 3 ,087 , 107 .077 1.265 1.555 1. 114 39.205 48 . 2 0 8 34.524 mean se c v (55) .090 .009 17. 110 1.311 : 130 17.110 40.646 4.015 17.110 g 0 2 / m 2 . hr g 02/m2.da g 02/m2.mo Rep 1 Rep 2 Rep 3 .047 .037 .020 1. 135 .881 .487 35.191 27.305 15.103 mean se c v ( 55) .035 .008 39 . 1 2 8 .834 . 188 39.128 2 5.866 5.843 39.128 g 02/m 2 . h r g 02/n2.da g 0 2/m2.mo Rep 1 Rep 2 Rep 3 . 134 . 144 .097 1.951 2.088 1.408 6 0.496 6 4.727 43.662 mean se c v (55) . 125 .014 19.795 1.816 .208 19.795 56.295 6. 434 19.795 g 0 2 / m 2 . da NET DAILY METABOLISM Rep 1 Rep 2 Rep 3 .816 1.207 .921 mean se c v (55) .982 . 117 2 0.612 1 1 1 GROSS COMMUNITY 1 PRODUCTION/ 1 24 H R R E S P I R A T I O N I 1 1 1 I PG/R24 Rep 1 Rep 2 Rep 3 1.719 2.371 2.891 me a n se c v C55) 2.327 .339 25.232 216 GROSS COMMUNITY PRODUCTIVITY g 02/m2.mo g 02/m2.da Table A-l (con’t) . Estimates of Coimnnnity Metabolism (g 02/m2) , Angnsta Creek, Michigan. 28 July 1975 (light), 29 July 1975 (dark), KELLOGG FOREST, STANDARD RON g 02/ni2.da g 02/m2.hr NET COMMUNITY PRODUCTIVITY COMMUNITY RESPIRATION Rep 1 Rep 2 Rep 3 .079 . 116 .073 1.150 1.694 1.063 35.665 52 . 5 0 2 32.949 menu se c v ( /?) .089 .014 2 6 .236 1.302 . 197 26.236 40 . 3 7 2 6. 115 26.236 g 0 2 / m 2 .hr g 0 2 / m 2 .da g 0 2 / m 2 .mo Rep 1 Rep 2 Rep 3 .077 .027 .073 1.843 .660 1.740 57.139 20.460 53.940 mean se c v (S5) .059 .016 46.335 1.414 .378 46.335 43.846 11.730 46.335 g 02/m2.hr g 02/ra2.da g 0 2 / m 2 .mo Rep 1 Rep 2 Rep 3 . 156 . 144 . 145 2 . 272 2.095 2. 121 70.425 64.948 65.763 mea n se c v (J5) . 148 .004 4.407 2. 163 .055 4 . 407 67.045 1.706 4 .407 g 02/m2.da NET DAILY METABOLISM Rep 1 Rep 2 Rep 3 .429 1.435 .381 mean se c v (%) .748 .344 79.533 1 PG/R24 I 1 GROSS COMMUNITY 1 PRODUCTION/ 1 24 H R RESPIRATION t I 1 1 1 Rep 1 Rep 2 Rep 3 mean se cv ( 1.233 3. 174 1.219 1.875 .650 59.989 217 G R OS S C O M M U N I T Y PR O D U C T I V I T Y g 02/m2.mo Table A—2# Estimates of Community Metabo 1 ism/g AFDW Detritus, Angaata Creek., Michigan. 15 January 1975 (light), 16 January 1975 (dark), SMITH SITE, STANDARD HUN g A F D W DETRITUS/tn2 NET COMMUNITY PRODUCTIVITY - . 000 -.000 -. 000 -.000 -.000 -.000 - . 004 -.005 -.005 mean se c v (55) 43 9 . 8 7 40.682 16.019 -.000 .000 16.294 -.000 .000 16.294 -.005 .000 16.294 B 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W 51 7 . 2 0 4 2 3 . lO 37 9 . 3 0 .000 .000 .000 .001 ,O01 .001 .016 .034 .039 me a n ae c v ( J5) 439.87 40.682 16.019 .000 .000 39.833 .001 .000 39 . 8 3 3 .030 .007 39.833 8 02/hr.g AFD W g 02/da.g AFDW 8 02/mo.b AFDW Rep 1 Rep 2 Rep 3 517.20 423. 10 379.30 .000 .000 .000 .000 .000 .000 .003 .008 .010 mean se c v ( J5) 4 39.87 4 0.682 16.019 .000 .000 57 . 2 2 2 .OOO .OOO 57. 2 2 2 .007 .002 57.222 Rep 1 Rep 2 Rep 3 -.000 -.001 -.001 mean se c v (/•) -.001 .000 34.619 1 1 1 1 1 1 1 1 1 1 PG/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 Rep 3 . 157 .235 .265 mean se c v t!5) .219 .032 25.656 218 Rep 1 Rep 2 Rep 3 g 02/da.g AFDW NET DAILY METABOLISM 8 02/mo.g AFDW 5 17.20 42 3 . 1 0 3 79.30 B A F D W D E T R I T U S / m2 GROSS COMMUNITY PRODUCTIVITY g 02/da.g AFDW Rep 1 Rep 2 Rep 3 B A F D W DETR I T U S / m 2 COMMUNITY RE S P I R A T I O N B 02/hr.g AFDW Table A-2 (con’t). Estimates of Community M e t a b o 1Ism/g AFDW Detritus, Augusta Creek, Michigan. 20 Ma y 1975 (light), 21 May 1975 (dark), SMITH SITE, STANDARD RUM b WET COMMUNITY PRODUCTIVITY AFD W DETRITUS/m2 -.000 -.000 -.000 -.001 -.000 -.001 -.036 -.005 -.018 mea n se c v (55) 253.34 98.104 67.071 -.000 .000 8 0.967 -.001 .000 80. 9 6 7 -.019 .009 80.967 g 0 2 / h r .g A F D W g 02/da.g AFDW B 0 2 / m o .g A F D W 8 6.630 426.30 247. 10 .000 .000 .000 .006 .002 .001 . 176 .049 .026 mean se c v (55) 253.34 98.104 67.071 .000 .000 96.035 .003 .002 96.035 .084 .047 96.035 g 02/hr.g AFDW g 02/da.g AFDW 8 0 2 / m o .g A F D W Rep 1 Rep 2 Rep 3 86.630 426.30 247.10 .000 .000 - . 000 .002 .001 - . 000 .072 .026 - . 002 me a n se c v (55) 2 53.34 9 8.104 67.071 .000 .000 116.40 .001 .001 116.40 .032 .022 116.40 Rep 1 Rep 2 Re p 3 -.003 -.001 -.001 mea n se c v ( 55) -.002 .001 86 . 9 7 0 1 1 1 1 1 1 1 1 1 1 PG/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 Rep 3 .411 .523 -.063 mea n se c v (5?) .290 . 180 107.13 219 Rep 1 Rep 2 Rep 3 g 0 2 / d a .g A F D W NET DAILY METABOLISM g 0 2 / m o .g A F D W 8 6.630 4 26.30 247.10 B AFDW DETRITUS/m2 GR OS S C O M M U N I T Y PRODUCTIVITY g 02/da.g AFDW Rep 1 Re p 2 Rep 3 B A F D W D E T R I T U S / m2 COMMUNITY RES P I R A T I O N g 0 2 / h r .g A F D W Table A-2 (con't). Estimates of Comnranlty M e tabo1Isra/g AFDW Detritus, Augusta Creek, Michigan. 17 July 1975.(light), 18 July 1975 (dark), SMITH SITE, STANDABD RUN g A F D W DETRITUS/m2 NET COMMUNITY PRODUCTIVITY - . 000 -.000 -.00© — .001 -.001 -.001 -.018 -.019 - .020 mean se c v (3) 379.80 6 2.195 28.363 -.000 .000 4.762 -.001 .OOO 4 . 763 -.019 .001 4.763 g 02/hr.g AFDW g 02/da.g AFDW g 02/mo.g AFDW 4 97.50 286. 10 355.80 .000 .000 .OOO .001 .001 .002 .041 .041 .049 mean se c v (55) 3 79.80 6 2.195 28 . 3 6 3 .000 .000 10.484 .001 .000 10.484 .044 .003 10.484 g 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W Re p 1 Rep 2 Rep 3 497.50 286.10 355.80 .000 .000 .000 .000 .000 .000 .008 .007 .011 mean se c v (JS) 379.80 62.195 28.363 .000 .000 2 6.226 .000 .000 26.226 .008 .001 26.226 Rep 1 Rep 2 Rep 3 -.001 -.001 -.001 mean se c v (R) -.001 .000 7.085 1 I t 1 1 1 1 1 1 1 PG/R24 GROSS COMMUNITY PROD U C T I O N / 24 H R RESPIRATION Rep 1 Rep 2 Re p 3 . 189 . 161 •222 mean se c v (55) . 191 .018 15.985 220 Re p 1 Rep 2 Rep 3 g 0 2 / d a .g A F D W NET DAILY METABOLISM g 02/mo.g AFDW 4 9 7.50 286.10 355.80 g AFD W DETRITUS/m2 GROSS COMMUNITY PRODUCTIVITY g 0 2 / d a .g A F D W Rep 1 Rep 2 Rep 3 g A F D W DETRlTUS/«ii2 COMMUNITY RESPIRATION g 02/hr.g AFDW Table A-2. Estimates of Conmnuilty Metabollsm/g AFDW Detritus, Augusta Creek., Michigan. 02 August 1973 (light), 01 August 1973 (dark). B AVENUE, STANDARD RUN e A F D W D E T R I T U S / m2 NET COMMUNITY PRODUCTIVITY .000 .000 .000 .000 .010 .010 mean se c v (N) 2 48.60 5.000 2.844 .000 .000 2.844 .000 .000 2.844 .01O .OOO 2.845 g 02/hr.g AFDW g 0 2 / d a .g A F D W g 02/mo.g AFDW 253.60 243.60 .000 .000 .002 .004 .073 .114 me a n se c v ( JJ) 248.60 5 . 000 2.844 .000 .OOO 30.463 .003 .OOl 30 . 4 6 3 .093 .020 30 .463 g 0 2 / h r .g A F D W g 02/da.g AFDW g 02/mo.g AFDW Rep 1 Rep 2 253.60 243.60 .000 .000 .002 .003 .054 .079 mean se c v ( N) 24 8 . 6 0 5.000 2 . 844 .000 .000 2 6.199 .002 .OOO 2 6.199 .066 .012 26.199 Rep 1 Rep 2 -.001 -.001 mean se c v (!?) -.001 .OOO 40.935 1 1 1 1 1 1 1 1 PG/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 .738 .693 mean se c v (/•) .715 .022 4.438 221 Rep 1 Re p 2 g 02/da.g AFDW NET DAILY METABOLISM e 02/mo.g AFDW 253.60 2 4 3.60 g A F D W D E T R I T U S / m2 GROSS COMMUNITY PRODUCTIVITY g 02/da.g AFD W Rep 1 Rep 2 g A F D W DETRITUS/ra2 COMMUNITY RESPIRATION g 0 2 / h r .g A F D W Table A-2 (con’t). Estimates of Community Metabo 1 ism/g AFDW Detritus, Augusta Greek., Michigan. 29 August 1973 (light), 30 August 1973 (dark), B AVENUE, STANDARD RUN g AFD W DETRITUS/m2 N E T COMMUNITY PRODUCTIVITY -.000 -.000 -.000 mean se c v ( 55) 265.40 4. 600 2.451 -.000 .000 -.000 .000 71.037 71.037 - .013 .006 7 1.036 g 0 2 / h r .g A F D W g 0 2 / d a .g A F D W er 0 2 / m o . g A F D W -.0 0 1 - .006 -.019 260.80 270.00 .000 .000 .002 .003 .052 .085 mean se c v ( 55) 2 65.40 4.600 2.451 .000 .OOO 3 3 .9 1 7 .002 .001 33.917 .069 .016 33. 9 1 7 g 02/hr.g AFDW g 02/da.g AFDW e 02/mo.g AFDW Rep 1 Rep 2 2 60.80 2 7 0.00 .000 .000 .001 .001 .023 .028 mean se c v (55) 2 6 5.40 4 .600 2.451 .000 .000 15.314 .001 .000 15.314 .025 .003 15.314 PG/R24 R ep 1 Rep 2 mea n se c v (55) -.0 0 1 -.002 -.001 .000 44.808 GROSS COMMUNITY PRODUCTION/ 2 4 H R RESP I R A T I ON Rep 1 Rep 2 .433 .330 mean se c v (55) .382 .052 19.091 222 Rep 1 Rep 2 g 02/da.g AFDW NET DAILY METABOLISM g 02/mo.g AFDW 2 60.80 270.00 fir A F D W DETRITUS/iii2 GROSS COMMUNITY PRODUCTIVITY g 0 2 / d a .g A F D W Rep 1 Rep 2 S AFDW DETRITUS/m2 COMMUNITY RE S P I R A T I O N g 0 2 / h r .g A F D W Table A-2 (con't). Estimates of Common!ty Meta b o 1ism/g AFDW Detritus, Augusta Creek, Michigan. 27 September 1973 (light), 26 September 1973 (dark), B AVENUE, STANDARD RUN b NET COMMUNITY PRODUCTIVITY A FDW DETRITUS/m2 - . 000 -.000 - .000 - .002 -.012 -.051 mean se c v (51) 2 15.37 116. 12 76.251 - .000 .000 87.557 -.001 .001 87 . 5 5 7 -.031 .019 87. 5 5 7 B 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W 3 3 1.50 99.250 .000 .000 .002 .009 .070 .284 mean se . c v (51) 2 1 5.37 116.12 76.251 .0 0 0 .0 O O 85.452 .006 .004 85.452 . 177 . 107 85.452 g 02/hr.g AFDW g 0 2 / d a .g A F D W g 0 2 / m o .g A F D W Rep 1 Rep 2 331.50 9 9.250 .000 .000 .0 0 1 .003 .023 .091 mean se c v (JS) 2 15.37 116. 12 76.251 .0 0 0 .0 0 0 84.300 .002 .001 84.300 .057 .034 84.300 Rep 1 Rep 2 mean se c v (B) - .002 -.006 -.004 .002 85.999 1 1 1 1 1 1 1 1 P C/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 .328 .320 mean se c v ( 55) .324 .004 1.788 223 Rep 1 Rep 2 g 02/da.g AFDW NET DAILY METABOLISM g 02/mo.g AFDW 3 31.50 9 9.250 B A F D W DETRITUS/m2 G R OSS C O M M U N I T Y PRODUCTIVITY g 0 2 / d a .g A F D W Rep 1 Rep 2 g AFD W DETRITUS/m2 COMMUNITY RESPIRATION B 02/hr.g AFDW Table A-2 (con't), Estimates of Community H e t a b o l U n / g AF D W Detritus, Augusta Creek, Michigan. 05 November 1973 (light), 06 November 1973 (dark), B AVENUE, STANDARD RUN g AFD W DETRITUS/m2 NET COMMUNITY PRODUCTIVITY -.000 -.000 -.000 -.000 - .012 -.005 mean se c v (55) 192.10 5 2.100 38.355 -.000 .000 59.028 -.000 .000 59.028 -.008 .003 59.028 B 02/hr.g AFDW g 0 2 / d a .g A F D W B 0 2 / m o .g A F D W 140.00 2 44.20 .000 .000 .003 .001 .077 .040 mean se c v (55) 19 2 . lO 52.100 3 8.355 .000 .000 4 4 .259 .002 .001 44.259 .058 .018 44.259 8 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W Rep 1 Rep 2 140.00 2 4 4.20 .000 .000 .001 .000 .021 .012 mean se c v (55) 192.10 5 2.100 38.355 .000 .000 37.001 .001 .000 37.001 .017 .004 37.001 Rep 1 Rep 2 -.002 -.001 mean se c v (55) -.001 .000 47.160 PG/R24 1 I 1 1 1 1 1 1 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep I Rep 2 .273 .306 mean se c v (55) .289 .016 7.891 224 Rep 1 Rep 2 g 02/da.g AFDW NET DAILY METABOLISM B 02/mo.g AFDW 140.00 244.20 8 A F D W DETRITUS/m2 GROSS COMMUNITY PRODUCTIVITY g 02/da.g AFDW Re p 1 Rep 2 B A F D W DETRITUS/m2 COMMUNITY RESPIRATION g 02/hr.g AFD W Table A-2 (con’t). Estimates of Community MetabolIsm/g AF D W Detritus, Augusta Creek. Michigan. 11 December 1973 (light), 12 December 1973 (dark.), B AVENUE, STANDARD RDM g A F D W D E T R I T U S / m2 RET COMMUNITY PRODUCTIVITY -.000 - .000 -.001 -.000 -.028 -.003 mean se c v (J5) 248.80 55.600 31 . 6 0 4 - .000 .000 111.53 -.000 .000 111.53 -.015 .012 111.53 g 02/hr.g AFDW g 02/da.g AFDW 8 02/mo.g AFDW 193.20 304.40 .000 .000 .901 .001 .043 .022 mean se c v (5?) 248.80 55.600 31.604 .000 .000 4 6 .029 .001 .000 46 . 0 2 9 .032 .010 46.029 g 02/hr.g AFDW g 0 2 / d a .g A F D W g 0 2 / m o .g A F D W Re p 1 Rep 2 193.20 30 4 . 4 0 -.000 .000 -.000 .000 -.011 .005 mean se c v (5?) 24 8 . 8 0 55.600 31.604 - . 000 .000 3 6 8.30 -.000 .000 36 8 . 3 0 - . 0 03 .008 36 8 . 3 0 Rep 1 Rep 2 -.002 -.001 mean se c v (M) -.001 .001 74.598 1 1 1 1 1 1 1 1 PG/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 -.265 .231 mean se c v (JS) - . 0 17 .248 2111.9 225 Rep 1 Rep 2 g 02/da.g AFDW NET DAILY METABOLISM g 0 2 / m o .g A F D W 193.20 3 04.40 8 A F D W D E T R I T U S / m2 GROSS COMMUNITY PRODUCTIVITY g 02/da.g AFDW Rep 1 Rep 2 8 A F D W D E T R I T U S / m2 COMMUNITY RESPIRATION g 02/hr.g AFDW Table A-2 (con’t). Estimates of Community Metabo1Ism/g AFDW Detritus, Angusta Creek, Michigan. 23 January 1974 (light), 22 January 1974 (dark), B AVENUE, STANDARD RUN g A F D W D E T R I T U S / m2 NET COMMUNITY PRODUCTIVITY .000 .000 .000 .000 .000 .000 mean se c v (ID 242.60 25.100 14.632 .000 .000 141.42 . ooo .000 141.42 .000 .000 141.42 g 02/hr.g AFDW g 02/da.g AFDW g 02/mo.g AFDW 217.50 267.70 .OOO .000 .QOl .001 .037 .034 me a n se c v (JS) 24 2 . 6 0 25.100 14.632 .000 .000 4.914 .001 .000 4.913 .036 .001 4. 913 g 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W Rep 1 Rep 2 217.50 267.70 .000 .000 .OOO .000 .015 .014 mea n se c v (%) 2 42.60 25.100 14.632 .000 .000 6.209 .000 .000 6 . 209 .015 .001 6.209 Rep 1 Rep 2 me a n se c v (") -.001 -.001 -.001 .000 4 . 002 1 1 1 1 1 1 1 1 PG/R24 CROSS COMMUNITY PROD U C T I O N / 24 H R R E S P I R A T I O N Rep 1 Rep 2 .411 .403 me a n se c v (5J) .407 .004 1.303 226 R ep 1 Rep 2 g 02/da.g AFDW NET DAILY METABOLISM g 0 2 / m o .g A F D W 217.50 267.70 g A F D W D E T R I T U S / m2 G ROSS C O M M U N I T Y PRODUCTIVITY g 0 2 / d a .g A F D W Rep 1 Rep 2 g AFDW DETRITUS/m2 COMMUNITY RESPIRATION g 02/hr.g AFDW Table A-2 (con't). Estimates of Community Jletabo 1Ism/g AF D W Detritus, Augusta Creek, Michigan. 05 March 1974 (light), @4 March 1974 (dark), B AVENUE, STANDARD RUN 8 A FDW DETRITUS/m2 NET COMMUNITY PRODUCTIVITY -.000 .000 -.000 .ooo - . 014 .003 mean se c v (55) 21 5 . 7 0 100.70 66.023 -.000 .000 209.43 -.000 .000 209.43 -.006 .008 209 . 4 3 g 02/hr.g AFDW g 02/da.g AFDW 8 0 2 / m o .g A F D W 115.00 316.40 .000 .002 ...... .072 mean se c v (J5) 215.70 100.70 66 . 0 2 3 .000 .002 .072 227 Re p 1 Re p 2 g 02/hr.g AFDW g 02/da.g AFDW 8 0 2 / m o .g A F D W Rep 1 Rep 2 115.00 316.40 .000 ...... .001 .020 mean se c v ( f?) 21 5 . 7 0 100.70 66.023 .000 .001 .020 g 02/da.g AFDW NET DAILY METABOLISM 8 02/mo.g AFDW 115.00 316.40 B A F D W DETRI T U S / m 2 GROSS COMMUNITY PRODUCTIVITY g 02/da.g AFDW Rep 1 Rep 2 8 A F D W D E T R I T U S / m2 COMMUNITY RESPIRATION g 02/hr.g AFDW Rep 1 Rep 2 mean se c v (7.) - . 002 1 1 1 1 1 1 1 1 PG/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 mean se c v (JS) .283 ..... .283 Table A-2 (con* t) . Estimates of Community Me tabo 1 Isra/g AFDW D e t r 1t us, Augusta Creek, Michigan.. 09 April 1974 (light), 10 April 1974 (dark), B AVENUE, STANDARD RUN g A F D W D E T R I T U S / m2 NET COMMUNITY PRODUCTIVITY -.000 -.000 -.000 -.000 - . 010 -.011 mean se c v (3) 320.30 92.000 40.621 -.000 .000 3.589 -.000 .000 3.588 -.011 .000 3.589 g 02/hr.g AFDW g 02/da.g AFDW g 02/mo.g AFDW 228.30 412.30 .000 .000 .003 .002 .078 .046 me a n se c v (3) 3 20.30 9 2.000 40.621 .000 .OOO 35.763 .002 .001 35.763 .062 .016 3 5.763 g 0 2 / h r .g A F D W g 0 2 / d a . g AF D W g 0 2 / m o .g A F D W R ep 1 Rep 2 2 28.30 4 12.30 .000 .000 .001 .000 .032 .014 mean se c v (3) 3 20.30 9 2.000 40.621 .000 .OOO 53.631 .001 .000 53.631 .023 .009 53.631 Rep 1 R ep 2 -.002 -.001 mean se c v < 3) -.001 .000 25.048 1 I 1 1 1 1 1 1 PG/R24 GROSS COMMUNITY PROD U C T I O N / 24 H R R E S P I R A T I O N Rep 1 Rep 2 mean se c v (3) .413 .311 . .362 .051 19.768 228 Rep 1 Rep 2 g 02/da.g AFDW NET DAILY METABOLISM g 0 2 / m o .g A F D W 228.30 412.30 g A F D W DETRI T U S/ m 2 G R OS S C O M M U N I T Y PRODUCTIVITY g 0 2 / d a .g A F D W Rep 1 Rep 2 g A F D W D E T R I T U S / m2 COMMUNITY RESPIRATION g 02/hr.g AFDW Table A-2 (con’t). Estimates of Community M e t a b o H s m / g AFDW Detritus, Angusta Creek, Michigan. 15 Ma y 1974 (light), 16 May 1974 (dark), B AVENUE, STANDARD RUN ts A F D W DETRITUS/'m2 NET COMMUNITY PRODUCTIVITY .000 .OOO .000 .001 .002 .028 mean se c v ( 55) 2 76.80 4 6.000 2 3 .502 .000 .000 125.62 .000 .000 125.62 .015 .013 125.62 g 02/hr.g AFDW g 02/da.g AFDW e 0 2 / m o .g A F D W 322.80 230.80 .000 .000 .002 .002 .057 .055 me a n se c v (55) 276.80 46.000 2 3.502 .000 .OOO 2.850 .002 .OOO 2.849 .056 .OOl 2 . 850 g 0 2 / d a .g A F D W e 0 2 / m o .g A F D W g 02/hr.g AFDW Rep 1 Rep 2 322.80 2 30.80 .000 .000 .001 .002 .037 .062 mean se c v (55) 2 76.80 46.000 2 3 .502 .000 •O O O 36.406 .002 .OOO 3 6.406 .049 .013 36.406 Rep 1 Rep 2 -.001 .000 mean se c v (55) -.000 .000 2 80.40 1 PG/R24 t 1 1 1 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 1 mea n 1 1 se c v ( 55) .638 1. 124 .881 .243 39.050 229 Rep 1 Rep 2 g 02/da.g AFDW NET DAILY METABOLISM e 0 2 / m o .g A F D W 322.80 230.80 e A F D W D E T R I T U S / m2 GROSS C O M M U N I T Y PRODUCTIVITY g 0 2 / d a .g A F D W Rep 1 Rep 2 e A F D W D E T R I T U S / m2 COMMUNITY RE S P I R A T I O N g 02/hr.g AFDW Table A-2 (con’t). Estimates of Community Metabo1ism/g A F D W D e t r 1tu b , Augnsta Greek, Michigan. 17 October 1974 (light), 18 October 1974 (dark), B AVENUE, STANDARD RUN g A F D W DETRITUS/m2 RET COMMUNITY PRODUCTIVITY -. 000 .000 -.000 -.000 .000 -.000 - . 004 .003 -.002 mean se c v (55) 592.00 39.902 11.674 -.000 .000 2 97.57 -.000 .000 29 7 . 5 7 -.001 .002 297.57 g 02/hr.g AFD W g 02/da.g AFDW g 0 2 / m o .g A F D W 6 23.90 6 3 9. 4 0 5 1 2.70 .000 .000 .000 .001 .001 .001 .024 .025 .035 mean se c v (55) 592.00 3 9 .902 11.674 .000 .000 2 0 .275 .001 .000 20.275 .028 .003 20.275 g 02/hr.g AFDW g 02/da.g AFDW g 02/mo.g AFDW Rep 1 Rep 2 Rep 3 6 2 3.90 639.40 5 12.70 .OOO .000 .000 .000 .000 .000 .007 .014 .014 me a n se c v (55) 5 92.00 3 9.902 11.674 .000 .000 3 2.406 .000 .000 3 2.407 .012 .002 32 . 4 0 7 Rep 1 Rep 2 Rep 3 -.001 -.000 -.001 mea n se c v (55) -.001 .000 30.339 1 1 1 1 1 I 1 1 1 1 PC/R24 GROSS COMMUNITY PRODUCTION/ 24 H R RESPI R A T I O N R ep 1 Rep 2 Rep 3 .304 .562 .396 mean se c v (55) .421 .076 31.147 230 Re p 1 Rep 2 Rep 3 g 02/da.g AFDW NET DAILY METABOLISM g 0 2 / m o .g A F D W 623.90 639.40 512.70 g A F D W DETRITUS/-m2 C R OS S C O M M U N I T Y PRODUCTIVITY g 0 2 / d a .g A F D W Rep 1 Rep 2 Re p 3 g AFD W DETRITUS/m2 COMMUNITY RESPIRATION g 02/hr.g AFDW Table A—2 (con’t). Estimates of Community Metabollsm/g AFDW Detritus, Augusta Creek, Michigan. 09 January 1973 (light), 08 January 1975 (dark), B AVENUE, STANDARD RUN g A F D W DETRITUS/m2 NET COMMUNITY PRODUCTIVITY - . 00© -.000 -.000 -.000 -.000 - . 0 00 -.002 -.012 -.001 mean se c v (JJ) 176.98 54.151 52.997 - .000 .000 120.52 -.000 .000 120.52 -.005 .004 120.52 g 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W 166.90 88.630 275.40 .000 .000 .000 .001 .004 .001 .019 . 128 .027 mean se c v (55) 176.98 54.151 52.997 .000 .000 104.38 .002 .001 104.38 .058 .035 104.38 g 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W Rep I Rep 2 Rep 3 166.90 88.630 275.40 .000 .000 .000 .000 .001 .OOO .006 .037 .009 mean se c v (55) 176.98 54.151 52.997 .000 .000 9 9.879 .001 .000 99.879 .017 .010 99.879 Rep 1 Rep 2 Rep 3 -.00© - .003 -.001 me a n se c v (55) -.001 .001 106.34 1 1 1 1 1 i 1 1 1 1 PG/R24 CROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 Rep 3 .288 .290 .340 me a n se c v (55) .306 .017 9.704 231 Rep 1 Rep 2 Rep 3 g 02/da.g AFDW NET DAILY METABOLISM g 0 2 / m o .g A F D W 166.90 88.630 275.40 g AFD W DETRITUS/m2 G R OS S C O M M U N I T Y PRODUCTIVITY g 02/da.g AFDW Rep 1 Rep 2 Rep 3 g A F D W D E T R I T U S / m2 COMMUNITY RESPIRATION g 0 2 / h r .g A F D W Table A-2 (con’t). Estimates of Community Meta b o 1Ism/g AFDW Detritus, AnguBta Creek, Hlchlgan. 16 April 1975 (light), 17 April 1975 (dark), B AVENUE, STANDARD RUN 8 A F D W DETRITUS/m2 RET COMMUNITY PRODUCTIVITY .000 -.000 .000 .001 -.000 .OOl .045 -.009 .025 mea n se c v (5S) 144.97 19.961 23.849 .000 .000 133.42 .001 .OOl 133.42 .020 .016 133.42 g 02/hr.g A F D W g 02/da.g AFDW g 0 2 / m o .g A F D W Rep 1 Rep 2 Rep 3 .000 .000 .000 .005 .005 .003 . 160 . 135 .087 144.97 19.961 2 3 .849 .000 .000 29.101 .004 .001 29.101 . 127 .021 29.101 g 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W Rep 1 Rep 2 Rep 3 135.80 115.90 183.20 .000 .000 .000 .004 .002 .002 mean se c v (5!) 144.97 19.961 23.849 .000 .000 40.494 .003 .OOl 40.494 6 02/da.g AFDW Rep 1 Rep 2 Rep 3 -.001 - . 002 -.000 mean se c v (JO -.001 .001 79.824 1 1 1 1 1 1 1 1 1 1 . 134. .067 .073 .091 .021 40.494 PG/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 Rep 3 .838 .494 .841 mean se c v (JO .725 .115 27.531 232 135.80 115.90 183.20 8 A F D W D E T R I T U S / m2 NET DAILY METABOLISM B 02/mo.g AFDW 135.80 115.90 183.20 mean se c v (JO G R OS S C O M M U N I T Y PRODUCTIVITY g 02/da.g AFDW Rep 1 Rep 2 Rep 3 8 A F D W D E T R I T U S / m2 COMMUNITY RE S P I R A T I O N g 02/hr.g AFDW Table A-2 (con’t). Estimates of Community Metabollsra/g AF D W Detritus, Augusta Creek, Michigan. 14 July 1975 (light). 15 July 1975 (dark), B AVENUE, STANDARD RUN e A F D W D E T R I T U S / m2 NET COMMUNITY PRODUCTIVITY -.000 .000 -.000 -.000 .OOO -.000 -.001 .002 -.001 mean se c v (3) 330.20 41. 116 21.567 .000 .000 2796.1 .000 .000 2796.1 .000 .001 2796. 1 g 0 2 / h r .g A F D W g 02/da.g AFDW e 02/mo.g AFDW 367.20 248.10 375.30 .000 .000 .000 .003 .004 .002 .082 .111 .076 mean se c v (51) 33 0 . 2 0 4 1.116 21.567 .000 .000 20.858 .003 .OOO 20.858 .089 .011 20.858 g 02/hr.g AFDW g 02/da.g AFDW g 02/mo.g AFDW Rep 1 Rep 2 Rep 3 367.20 248.10 375.30 .000 .000 .000 .002 .002 .001 .050 .071 .046 mea n se c v (3) 330.20 41. 116 21.567 .000 .000 23.941 .002 .000 23.941 .056 .008 23.941 Rep 1 Rep 2 Rep 3 mea n se c v (3) -.001 -.001 -.001 -.001 .000 15.709 1 1 1 1 1 1 I 1 1 1 PG / R 2 4 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 Rep 3 mean se c v (3) .616 .644 .610 .623 .010 2.868 233 Rep 1 Rep 2 Rep 3 g 02/da.g AFDW NET DAILY METABOLISM g 02/mo.g AFDW 367.20 248.10 375.30 g AFD W DETRITUS/m2 GROSS COMMUNITY PRODUCTIVITY g 02/da.g AFDW Rep 1 R ep 2 Rep 3 e A F D W DETRITUS/m2 COMMUNITY RESPIRATION g 02/hr.g AFDW Table A-2 (con't). Estimates of Community Metabolism/g AFDW Detrltns, Augusta Creek, Michigan. 30 October 1974 (light), 31 October 1974 (dark), U P P E R 43, STANDARD RUN e A F D W DETRITUS/'m2 NET COMMUNITY PRODUCTIVITY .000 .001 .005 .OH . 141 .348 mean se c v (55) 218.35 44.750 2 8.984 .001 .000 59.990 .008 .003 5 9.990 .244 . 104 59. 9 9 0 B 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W 263. 10 173.60 .000 .001 .009 .014 .280 .428 mean se c v (55) 218.35 4 4.750 28.984 .OOO .000 29.597 .011 .002 29.597 .354 .074 2 9.597 8 02/hr.g AFDW g 02/da.g AFDW g 02/mo.g AFDW Rep 1 Rep 2 2 63.10 173.60 .001 .002 .008 .017 .262 .534 mean se c v (55) 218.35 4 4.750 2 8.984 .001 .000 48.248 .013 .004 48.248 .398 . 136 48 . 2 4 8 Rep 1 Rep 2 -.001 .003 mean se c v (55) .001 .002 197.59 1 1 1 1 I 1 1 P G/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 .937 1.248 mean se c v (55) 1.092 . 155 20.085 234 Rep 1 Rep 2 g 02/d a .g A F D W NET DAILY METABOLISM g 02/mo.g AFDW 2 63.10 173.60 B A F D W D E T R I T U S / m2 G R OSS C O M M U N I T Y PRODUCTIVITY g 02/da.g AFDW Rep 1 Rep 2 8 A F D W D E T R I T U S / m2 COMMUNITY RESPIRATION 8 02/hr.g AFDW Table A-2 (con't). Estimates of Community Metabollsra/g AFDW Detritus, Augusta Greek, Michigan. 21 January 1975 (light), 22 January 1975 (dark), TIPPER 43, STANDARD RON S AFDW DETRITUS/m2 NET COMMUNITY PRODUCTIVITY .000 .000 .000 .001 .001 .001 .029 .021 .023 mean se c v (J5) 235.03 35.480 26.146 .000 .000 16.970 .001 .OOO 16.971 .024 .002 16.971 g 02/hr.g AFDW g 02/da.g AFDW g 02/mo.g AFDW 301.40 180.10 223.60 .000 .000 .000 .002 .001 .002 .053 .034 .062 mea n se c v (JO 23 5 . 0 3 35 . 4 8 0 26.146 .000 .000 29.088 .002 .000 29.088 .050 .008 29 . 0 8 8 g 02/hr.g AFDW g 02/da.g AFDW g 0 2 / mo.g A F D W Rep 1 Rep 2 Re p 3 301.40 180.10 223.60 .000 .000 .000 .002 .OOl .002 .050 .035 .048 mean se c v (JS) 23 5 . 0 3 35 . 4 8 0 2 6.146 .000 .000 19.046 .001 .OOO 19.046 .044 .005 19.046 Rep 1 Rep 2 Rep 3 -. OOO .000 - .000 mea n se c v (J5) - . 000 .OOO 143.23 1 1 1 1 1 1 1 1 1 1 PG/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 Rep 3 .948 1.020 .770 mean se c v (%) .912 .074 14.071 235 Rep 1 Rep 2 Rep 3 g 02/da.g AFDW NET DAILY METABOLISM g 02/mo.g AFDW 3 0 1.40 180.10 223.60 g AFD W DETRITUS/m2 G R OS S C O M M U N I T Y PRODUCTIVITY g 02/da.g AFDW Rep 1 Rep 2 Re p 3 g A F D W DETRITUS/'m 2 COMMUNITY RE S P I R A T I O N g 02/hr.g AFDW Table A-2 (con't). Estimates of Community Metabolisn/g AFDW Detritus, Augusta Creek, Hlchigan. 27 Hay 1975 (light), 28 May 1975 (dark), UPPER 43, STANDARD RUN e A F D W D E T R I T U S / m2 RET COMMUNITY PRODUCTIVITY .001 .001 .000 .019 .009 .007 .579 .284 .202 mean se c v (M) 179. 10 2.532 2 . 449 .001 .OOO 55.904 .011 .004 55 . 9 0 4 .355 .114 55.904 g 02/hr.g AFDW g 02/da.g AFDW g 02/mo.g AFDW 182.30 180.90 174.10 .000 • .000 .000 .012 .009 .011 .364 .284 .341 m ea n se c v (5?) 179. 10 2.532 2.449 .000 .000 12.538 .011 .001 12.538 .330 .024 12.538 g 0 2 / h r .g A F D W g 02/da.g AFDW g 0 2 / m o .g A F D W Rep 1 Rep 2 Rep 3 182.30 180.90 174.10 .002 .001 .001 .026 .015 .013 .805 .461 .414 mean se c v (JI) 179.10 2 . 532 2.449 .001 .000 3 8.180 .018 .004 3 8.179 .560 . 123 38.180 Rep 1 Rep 2 Rep 3 mea n se c v (8) .014 .006 .002 .007 .004 82.457 1 1 1 1 1 1 r i i i P G/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 Rep 3 2.212 1.623 1.214 mean se c v (J5) 1.683 .290 29.822 236 Rep 1 Re p 2 Rep 3 g 02/da.g AFDW NET DAILY METABOLISM g 0 2 / m o .g A F D W 182.30 180.90 174.10 e A F D W D E T R I T U S / m2 GROSS COMMUNITY PRODUCTIVITY g 02/da.g AFDW Rep 1 Rep 2 Rep 3 e A F D W DETRITUS/ra2 COMMUNITY RESPIRATION g 02/hr.g AFDW Table A-2 (con’t). Estimates of Community Metabollsm/g AFDW Detritus. Augusta Creek, Michigan. 21 July 1975 (light), 22 July 1975 (dark), UPPER 43, STANDARD RUN g A F D W DETRITUS/m2 NET COMMUNITY PRODUCTIVITY .000 .000 .002 .004 .059 . 134 me a n se c v (55) 328.03 34.914 18.435 .000 .000 55.539 .003 .001 55.539 .097 .038 55.539 g 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W 34 7 . 2 0 37 6 . 6 0 260.30 .000 .000 .000 .006 .007 .012 . 181 .202 .360 mean se c v (55) 328.03 34 . 9 1 4 18.435 .000 .OOO 39.573 .008 .002 39 . 5 7 3 .248 .057 39.573 g 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W Rep 1 Rep 2 Rep 3 347.20 376.60 260.30 .000 .001 .006 .008 . 171 .259 mean se c v ( K) 328.03 34.914 18.435 .000 .000 29.112 .007 .OOl 29.113 .215 .044 29. 112 Rep 1 Rep 2 Rep 3 mea n se c v (55) -.000 002 • • • • • • .001 1 1 1 1 1 1 1 1 1 1 PG/R24 GROSS COMMUNITY PRODUCTION/ 24 H R RESPIRATION Rep 1 Rep 2 Rep 3 .941 1.284 mean se c v ( 55) 1.112 . 172 21.816 237 Rep 1 Rep 2 R ep 3 g 02/da.g AFDW NET DAILY METABOLISM g 02/mo.g AFDW 34 7 . 2 0 3 7 6. 6 0 260.30 B AFD W DETRITUS/m2 GROSS C O M M U N I T Y PRODUCTIVITY g 0 2 / d a .g A F D W Rep 1 Rep 2 Rep 3 S A F D W DETRITUS/m2 COMMUNITY RE S P I R A T I O N g 0 2 / h r .g A F D W Table A-2 (con’t). Estimates of Community Metabo 1 Ism/g AF D W Detritus, Augusta Creek., Michigan. 04 September 1973 (light), 05 September 1973 (dark), NAGEL SITE, STANDARD RUN B A F D W D E T R I T U S / m2 N ET COMMUNITY PRODUCTIVITY .002 .001 .020 .016 .600 .492 mean se c v (J5) 164.80 3 4.100 2 9.262 .OOl .000 13.959 .018 .002 13.959 .546 .054 13.959 g 02/hr.g AFDW g 02/da.g AFDW g 02/mo.g AFDW 130.70 198.90 .001 .001 .021 .013 .622 .386 mean se c v (%) 164.80 34.100 2 9.262 .001 .000 33.121 .017 .004 33.121 .504 .118 33.121 g 02/hr.g AFDW g 02/da.g AFDW e 0 2 / m o .g A F D W Rep 1 Rep 2 130.70 198.90 .002 .002 .031 .023 .938 .702 mean se c v ( %y 164.80 3 4.100 29.262 .002 .000 20.361 .027 .004 20.361 .820 .118 20.361 Rep 1 Rep 2 .011 .011 mea n se c v ( 3) .011 .000 .000 1 1 1 1 1 1 1 ) PG/R24 G R OSS C O M M U N I T Y PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 1.506 1.816 me a n se c v (3) 1.661 . 155 13.207 238 Rep 1 Rep 2 g 02/da.g AFDW NET DAILY METABOLISM g 02/mo.g AFDW 130.70 198.90 B A F D W DETRITUS/m2 GROSS COMMUNITY PRODUCTIVITY g 02/da.g AFDW Rep I Rep 2 8 A F D W DETRITUS/m2 COMMUNITY RESPIRATION g 0 2 / h r .g A F D W Table A-2 (con't). Estimates of Community Metabo1ism/g AFDW Detritus, Augusta Creek, Michigan. 04 October 1973 (light). 03 October 1973 (dark), NAGEL SITE, STANDARD RUN e A F D W DETRITUS/ra2 NET COMMUNITY PRODUCTIVITY .001 .001 .006 .007 . 182 -.202 mean se c v (55) 2 2 3.75 3.851 2.434 .001 .000 7.420 .006 .OOO 7.420 . 192 .010 7.420 g 02/hr.g A FDW g 0 2 / d a .g A F D W g 0 2 / m o .g A F D W 227.60 219.90 .000 .000 .005 .006 . 167 .200 mean se c v (55) 2 23.75 3.851 2.434 .000 .000 12.665 .006 .001 12.665 . 184 .016 12.666 g 02/hr.g A F D W g 02/da.g AFDW g 0 2 / m o .g A F D W Rep 1 Rep 2 227.60 219.90 .001 .001 .008 .0 1 0 .263 .299 mean se c v (55) 223.75 3.851 2.434 .001 .000 9 . 083 .009 .001 9.083 .281 .018 9 . 083 Rep 1 Rep 2 .003 .003 mea n se c v (55) .003 .OOO 2 .292 1 1 1 1 1 1 1 1 PG/R24 G ROSS C O M M U N I T Y PRODUCTION/ 24 H R RESPI R A T I O N Rep 1 Hep 2 1.570 1.492 me a n se c v ( 55) 1.531 .039 3.603 239 Rep 1 Rep 2 g 02/da.g AFDW NET DAILY METABOLISM 0 02/mo.g A F D W 227.60 219.90 fS A F D W D E T R I T U S / m2 GROSS COMMUNITY PRODUCTIVITY g 02/da.g AFDW R ep 1 Rep 2 e A FDW DETRITUS/m2 COMMUNITY RE S P I R A T I O N g 02/hr.g AFDW Table A-2 (con’t). Estimates of Commnnlty Metabo1ism/g AFDW Detritus, Augusta Creek, Michigan. 12 November 1973 (light), 13 November 1973 (dark), NAGEL SITE, STANDARD RUN 8 A F D W DETRITUS/'m2 NET COMMUNITY PRODUCTIVITY .001 .001 .007 .010 .224 .297 mean se c v (J?) 2 86.45 47.950 23 . 6 7 3 .001 .000 19.729 .009 .001 19.728 .261 .036 19.728 g 0 2 / h r .g A F D W g 0 2 / d a .g A F D W g 02/mo.g AFDW 334.40 238.50 .OOO .000 .006 .006 . 169 . 185 mean se c v (JJ) 2 8 6.45 47.950 23 . 6 7 3 .000 .000 6 . 290 .006 .000 6.290 . 177 .008 6.290 g 02/hr.g AFDW g 02/da.g AFDW 8 02/mo.g AFDW Rep 1 Rep 2 3 34.40 2 38.50 .001 .001 .010 .012 .294 .373 mean se c v ( ») 286.45 4 7 .950 23.673 .001 .000 16.786 .011 .001 16.785 .334 .040 16.785 Rep 1 Rep 2 .004 .006 me a n se c v (J5) .005 .001 28.613 1 I 1 1 1 1 1 1 1 PG/R24 GROSS COMMUNITY PRODUCTION/ 24 H R RESP I R A T I O N Rep 1 Rep 2 1.741 2.021 me a n se c v (%) 1.881 . 140 10.552 240 Rep 1 R ep 2 g 02/da.g AFDW NET DAILY METABOLISM g 0 2 / m o .8 A F D W 334.40 238.50 8 AFD W DETRITUS/m2 GROSS COMMUNITY PRODUCTIVITY g 02/da.g AFDW Rep 1 Rep 2 8 A F D W DETRlfUS/m2 COMMUNITY RESPIRATION g 02/hr.g AFDW Table A—2 (con’t). Estimates of Community Me tabo1ism/g AFDW Detrltns, Angusta Creek, Michigan. 18 December 1973 (light), 19 December 1973 (dark), NAGEL SITE, STANDARD RUN 8 A F D W D E T R I T U S / m2 NET COMMUNITY PRODUCTIVITY .000 .000 .004 .003 .114 .084 mean se c v (55) 297.70 44.200 20.997 .000 .000 2 1 .405 .003 .000 21.405 .099 .015 21.405 g 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W 341.90 253.50 .000 .000 .003 .002 .079 .072 mean se c v (55) 297.70 44.200 20.997 .000 .000 7.028 .002 .OOO 7 .029 .076 .004 7.029 g 02/hr-g A FDW g 02/da.g AFDW g 0 2 / mo.g A F D W Re p 1 Rep 2 341.90 253.50 .001 .000 .005 .004 . 145 .112 mea n se c v (55) 297.70 44.200 20.997 .000 .000 18. 185 .004 .001 18. 185 . 128 .016 18.185 Rep 1 Rep 2 .002 .001 mea n se c v ( 55) .002 .000 34.293 1 I 1 1 1 1 1 1 PG/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 1.820 1.552 me a n se c v ( 55) 1.686 . 134 11.226 241 Rep 1 Rep 2 g 02/da.g AFDW NET DAILY METABOLISM g 0 2 / m o .g A F D W 34 1 . 9 0 233.50 g AFDW DETRITUS/m2 GROSS COMMUNITY PRODUCTIVITY g 02/da.g AFDW Rep 1 Rep 2 8 A FDW DETRITUS/m2 COMMUNITY RESPIRATION g 02/hr.g AFDW Table A-2 (con’t). Estimates of Community Metabo 1 ism/g AFDW Detritus, Augusta Creek, Michigan. 31 January 1974 (light), 01 February 1974 (dark), NAGEL SITE, STANDARD RUN e AFDW DETRITUS/m2 NET COMMUNITY PRODUCTIVITY g 0 2 / m o .g A F D W 219. 10 379.90 .000 .000 .003 .OOl .081 .022 mean se c v (55) 299.50 80.400 37.964 .000 .000 8 1.852 .002 .001 81.852 .051 .030 81 . 8 5 2 g 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W 219.10 379.90 .000 .000 .002 .002 .071 .054 mean se c v (55) 299.50 80.400 37.964 .000 .000 19.720 .002 .OOO 19.720 .063 .009 19.72© g 0 2 / h r .g A F D W g 02/da.g AFDW g 0 2 / m o .g A F D W Rep 1 Rep 2 219. 10 379.90 .000 .000 .004 .001 .111 .044 m ean se c v (55) 299.50 80.400 37.964 .000 .000 60.975 .002 .001 60.975 .077 .033 60.975 g 0 2 / d a .g A F D W RET DAILY METABOLISM Rep 1 Rep 2 -.ooo mean se c v (55) .000 .001 235.53 .001 1 1 1 1 1 1 1 1 PC/R24 G R OSS C O M M U N I T Y PROD U C TI O N / 24 H R RESP I R A T I O N Rep 1 Rep 2 1.553 .817 mean se c v ( 55) 1.185 .368 43.898 242 Rep 1 Rep 2 g A FDW DETRITUS/m2 GROSS COMMUNITY PRODUCTIVITY g 02/da.g AFDW Rep 1 Rep 2 g A F D W DETRITUS/m2 COMMUNITY RESPIRATION g 02/hr.g AFDW Table A - 2 ( c o n ’t). Estimates of C o m m u n i t y Metabo 1 ism/g A F D W Detritus, Augnsta Creek, Michigan. 2 0 M a r c h 1974 (light), 21 M a r c h 1974 (dark), N A G E L SITE, S T A N D A R D R O N e AF D W DETRITUS/m2 NET COMMUNITY PRODUCTIVITY g 02/mo.g AFDW 2 2 3.10 4 9 7.70 .001 .000 .010 .003 .298 . 106 mean se c v (5S) 3 60.40 137.30 5 3.877 .001 .000 6 7.029 .007 .003 67 . 0 2 9 .202 .096 67.029 g 02/hr.g AFDW g 02/da.g AFDW B 02/mo.g AFDW 2 23.10 497.70 .000 .000 .008 .003 .245 .080 mean se c v (J5> 3 60.40 137.30 53.877 .000 .000 71.570 .005 .003 71 . 5 7 0 . 163 .082 71 .570 g 02/hr.g AFDW g 02/da.g AFDW 8 02/mo.g AFDW Rep 1 Rep 2 223.10 497.70 .001 .000 .014 .005 .421 . 147 mean se c v (") 360.40 137.30 5 3.877 .001 .000 6 8.342 .009 .004 68. 3 4 2 .284 . 137 68.342 8 02/da.g AFDW \ PG/R24 1 NET DAILY METABOLISM Rep 1 Rep 2 .006 .002 m ean se c v (J5) .004 .002 64.024 1 1 1 1 1 1 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 1.721 1.829 me a n se c v (JJ) 1.775 .054 4.274 243 Rep 1 Rep 2 8 AFDW DETRITUS/m2 GROSS COMMUNITY PRODUCTIVITY g 02/da.g AFDW Rep 1 Rep 2 e A F D W DETR I T U S / m 2 COMMUNITY RESPIRATION g 02/hr.g AFDW Table A-2 (con’t). Estimates of Community MetabolIsm/g AFDW Detritus, Augusta Creek, Nlchlgan. 23 April 1974 (light), 22 April 1974 (dark), NAGEL SITE, STANDARD RUN e A F D W D E T R I T U S / m2 NET COMMUNITY PRODUCTIVITY .000 .000 .006 .004 . 169 . 127 me a n se c v (J5) 321.35 5 5.950 2 4.623 .000 .000 20.046 .005 .OOl 2 0.046 .148 .021 20.045 g 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W Rep 1 Rep 2 3 77.30 265.40 .000 .001 .008 .014 .242 .411 mea n se c v (55) 321.35 55.950 2 4.623 .000 .000 3 6.669 .Oil .003 36.669 .326 .085 36.669 g 02/hr.g AFDW g 02/da.g AFDW g 02/rao.g A F D W Rep 1 Rep 2 3 7 7.30 2 6 5.40 .001 .001 .010 .012 .307 .362 mean se c v (55) 321.35 5 5.950 2 4.623 .001 .000 11.590 .011 .001 11.590 .334 .027 11.590 g 02/da.g AFDW NET DAILY METABOLISM g 02/mo.g A F D W 3 7 7.30 2 65.40 e A F D W D E T R I T U S / m2 GROSS COMMUNITY PRODUCTIVITY g 02/da.g AFDW Rep 1 Rep 2 e A F D W DETRITUS/m2 COMMUNITY RESPIRATION g 02/hr.g AFDW Rep 1 Rep 2 .002 -.002 mea n se c v (5?) .000 .002 997.60 1 1 1 1 1 1 1 1 PG/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 1.270 .880 mean se c v (55) 1.075 . 195 25.625 Table A**2 (con’t). Estimates of Community MetabolIsm/g AFDW Detritus, Augusta Greek, Michigan. 24 April 1974 (light)., 22 April 1974 (dark), NAGEL SITE, STANDARD RUN g A F D W DETRITUS/fn2 NET COMMUNITY PRODUCTIVITY .001 .000 .009 .006 .275 . 194 mean se c v ( J5) 321.35 53.950 24.623 .001 .0 0 2 4 .455 .008 .001 2 4.455 .234 .040 24.455 g 02/hr.g AFDW g 02/da.g AFDW B 02/mo.g AFDW 377.30 2 65.40 .000 .001 .008 .014 .242 .411 mean se c v ( J5) 321.35 5 5.950 24.623 .000 .000 3 6.669 .011 .003 36.669 .326 .085 3 6.669 g 0 2 / h r .g A F D W g 02/da.g AFDW B 02/mo.g AFDW Rep 1 Rep 2 3 77.30 2 65.40 .001 .001 .014 .014 .413 .429 mean se c v ( J5) 321.35 5 5 .950 2 4 .623 .001 .000 2.699 .014 .000 2.699 .421 .008 2.699 Rep 1 Re p 2 .006 .001 mea n se c v ( JS) .003 .003 113.82 1 1 I 1 1 1 1 1 PG/R24 GROSS COMMUNITY PROD U C T I O N / 24 H R RESP I R A T I ON Rep 1 Rep 2 1.710 1.045 me a n se c v (J5) 1.378 .333 34.139 245 Rep 1 Rep 2 g 02/da.g AFDW NET DAILY METABOLISM g 02/mo.g AFDW 377.30 263.40 B A F D W DETRITUS/m2 GROSS C O M M U N I T Y PR O D U C T I V I T Y g 02/da.g AFDW Rep 1 Rep 2 e A F D W DETRITUS/m2 COMMUNITY RESPIRATION g 02/hr.g AFDW Table A ~ 2 (con't). Estimates of C o m m u n i t y M e t a b o l i s m / g A F D W Detritus, Augusta Creek, 29 M a y 1974 (light), 29 M a y 1974 (dark). N A G E L SITE, D I E L R U N g A F D W D E T R ITUS/m2 NET COMMUNITY PRODUCTIVITY .001 .001 .012 .008 .383 .263 mean se c v (55) 2 30.75 7 5.650 46.364 .OOl .000 2 6 .243 .010 .002 26.243 .323 .060 26.243 g 0 2 / h r .g A F D W g 0 2 / d a .g A F D W g 0 2 / m o .g A F D W 155. lO 306.40 .OOl .000 .018 .009 .543 .290 me a n se c v ( 55) 230.75 75.650 4 6.364 .001 .000 4 2.973 .013 .004 42.973 .417 . 127 42.973 g 02/hr.g AFDW g 02/da.g AFDW g 02/mo.g AFDW Rep 1 Rep 2 155. 10 3 06.40 .002 .001 .023 .014 .723 .444 me a n se c v ( 55) 230.75 7 5 .650 46.364 .001 .000 33.713 .019 .004 33.713 .583 . 139 33.713 Rep 1 Rep 2 .006 .005 mea n se c v (55) .005 .000 10.548 1 1 1 1 1 1 1 PG/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 1.329 1.531 mean se c v (55) 1.430 . 101 9.986 246 Rep 1 Rep 2 g 02/da.g A F D W NET DAILY METABOLISM g 0 2 / m o .g A F D W 155.10 306.40 B AFD W DETRITUS/m2 GROSS COMMUNITY PRODUCTIVITY g 0 2 / d a .g A F D W Rep 1 Rep 2 g A FDW DETRITUS/m2 COMMUNITY RESPIRATION g 02/hr.g AFDW Michigan. Table A-2 (con't). Estimates of Community Ketabollsm/g AFDW Detrltns, Augusta Creek, Hlchlgan. 01 July 1974 (light), 01 July 1974 (dark), NAGEL SITE, DIEL RUN g AFD W DETRITUS/m2 NET COMMUNITY PRODUCTIVITY .000 .007 .223 mean se c v (55) 2 83.00 .000 .007 .223 g 02/hr.g AFDW g 02/da.g AFDW g 02/mo.g AFDW 2 83.00 .001 .013 .402 mean se c v (55) 2 83.00 .001 .013 .402 247 Rep 1 g 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W 1 2 83.00 .001 .015 .478 mean se c v ( 55) 2 83.00 .001 .015 .478 Rep PG/R24 g 02/da.g AFDW RET DAILY METABOLISM g 02/mo.g AFD W 2 83.00 g A FDW DETRITUS/m2 GROSS COMMUNITY PRODUCTIVITY g 0 2 / d a .g A F D W Rep 1 g A F D W D E T R I T U S / m2 COMMUNITY RESPIRATION g 02/hr.g AFDW 1 .002 m ea n se c v ( 55) .002 R ep GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N 1 1. 188 me a n se c v ( 55) 1. 188 Rep Table A-2 (con't). Estimates of Community MetaboIIsm/g AF D W Detritus, Augusta Creek, Michigan. 05 August 1974 (light), 05 August 1974 (dark), NAGEL SITE, STANDABD RON g A F D W DETRITUS/m2 N ET COMMUNITY PRODUCTIVITY Rep 1 Rep 2 Rep 3 mean se c v (R) .014 .014 .440 .444 235.40 37.882 27.873 .001 .000 .649 .014 .000 .649 .442 .002 .647 g 02/hr.g AFDW g 02/da.g AFDW ,276 25 2 . 6 0 162.90 290.70 .000 .000 .007 .232 mean se c v ( R) 235.40 37.882 27.873 .OOO .000 12.127 .008 .001 12.127 .254 .022 12. 127 g 02/hr.g AF DW g 02/da.g AFDW g 0 2 / m o .g A F D W •••••• 252.60 162.90 290.70 .001 me a n se c v (R) 235.40 37 . 8 8 2 27 . 8 7 3 .001 mean se c v (R) .011 .019 .604 #**••• .019 PG/R24 1 1 1 1 1 1 1 .604 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 Rep 3 me a n se c v <%) 2. 189 2. 189 248 Rep 1 Re p 2 Re p 3 Rep 1 Rep 2 Rep 3 .009 g 02/mo.g AFDW Rep 1 Rep 2 Rep 3 g 02/da.g AFDW NET DAILY METABOLISM g 02/mo.g AFDW .001 .001 e A F D W DETRITUS/m2 G R OSS C O M M U N I T Y PRODUCTIVITY g 0 2 / d a .g A F D W 252.60 162.90 2 90.70 g A F D W D E T R I T U S / m2 COMMUNITY RESPIRATION g 02/hr.g AFDW Table A-2 (con’t). Estimates of Community M e t a b o 1lsm/g AFDW Detritus, Augusta Creek, Michigan. 06 November 1974 (light), 07 November 1974 (dark), NAGEL SITE, STANDARD RUN e A F D W D E T R I T U S / m2 NET COMMUNITY PRODUCTIVITY .000 .000 .000 .004 .004 .002 .118 . 124 .068 mean se c v (55) 3 8 8.23 4 9 .905 2 2 .265 .000 .000 29.404 .003 .001 29.404 .103 .018 29.404 g 0 2 / h r .g A F D W g 02/da.g AFDW g 0 2 / m o .g A F D W 312.30 482.30 37 0 . 1 0 .000 .000 .000 .002 .003 .002 .071 .098 .053 mean se c v (55) 38 8 . 2 3 49.905 22.265 .000 .000 29.996 .002 .000 29.996 .074 .013 29.996 g 02/hr.g AFDW g 02/da.g AFDV g 02/mo.g AFDV Rep 1 Rep 2 Rep 3 3 12.30 482.30 37 0 . 1 0 .000 .001 .000 .005 .006 .003 . 148 . 165 .091 mean se c v (55) 3 88.23 49.905 2 2.265 .000 .000 2 8 . BOO .004 .001 28.800 . 135 .022 28.800 Rep 1 Rep 2 Rep 3 .003 .002 .001 mean se c v (55) .002 .000 33.681 PG/R24 1 1 1 1 1 t 1 1 1 1 G R OSS C O M M U N I T Y PRODUCTION/ 24 H R RESPIRATION Rep 1 Re p 2 Re p 3 2 .068 1.688 1.701 mean se c v (55) 1.819 . 125 11.868 249 Rep 1 Rep 2 Rep 3 g 0 2 / d a .g A F D W NET DAILY METABOLISM B 02/mo.g AFDW 312.30 482.30 370. 10 g A F D V DETRITUS/"m 2 GROSS COMMUNITY PRODUCTIVITY g 02/da.g AFDV Rep 1 Rep 2 R ep 3 B A F D W D E T R I T U S / m2 COMMUNITY RESPIRATION g 02/hr.g AFDW Table A-2 (coo’t). Estimates of Community Meta b o 1 lsm/g AFDW Detritus, Augusta Creek, Michigan. 30 January 1973 (light), 31 January 1975 (dark), NAGEL SITE, STANDARD RON e A F D W D E T R I T U S / m2 MET COMMUNITY PRODUCTIVITY .000 .000 .000 .001 .001 .001 .017 .024 .022 mean . se c v (5S) 393.33 7.789 3.430 .000 .000 16.855 .001 .000 16.855 .021 .002 16.855 6 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W 3 8 4.00 4 0 8.80 3 87.20 .000 .000 .000 .002 .001 .002 .060 .042 .047 mean se c v (%) 3 9 3.33 7.789 3 . 430 .000 .000 18.972 .002 .000 18.972 .050 .005 18.973 g 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W Rep 1 Rep 2 Hep 3 3 84.00 4 08.80 3 87.20 .000 .000 .000 .001 .001 .001 .042 .041 .042 mean se c v ( J5) 3 93.33 7.789 3 .430 .000 .000 .890 .001 .000 .890 .042 .OOO .889 Rep 1 Rep 2 Rep 3 -.001 -.000 - .000 me a n se c v ( J5) - .000 .000 112.35 1 1 I I 1 1 ■ 1 1 1 1 PC/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 Rep 3 .697 .982 .888 mean se c v (55) .856 .084 16.946 250 Re p 1 Hep 2 Hep 3 g 02/da.g AFDW NET DAILY METABOLISM g 0 2 / m o .g A F D W 384.00 408.80 38 7 . 2 0 B A FDW DETRITUS/m2 GROSS COMMUNITY PRODUCTIVITY g 02/da.g AFDW Hep 1 Rep 2 Rep 3 B AFD W DETRITUS/m2 COMMUNITY RESPIRATION S 02/hr.g AFDW ii I n Table A—2 (con’t). Estimates of Community Metabollsm/g AFDW Detritus, Augusta Creek, Michigan. 06 May 1975 (light), 07 Ifay 1975 (dark). NAGEL SITE, STANDARD RUN B A F D W D E T R I T U S / m2 NET COMMONITV PRODUCTIVITY .000 .000 .000 .005 .002 .004 . 150 .055 . 133 mean se c v (55) 325.60 63.118 3 3 .576 .000 .000 45.084 .004 .001 45.084 .113 .029 45.084 g 02/hr.g AFDW g 02/da.g AFDW B ’ 0 2 / m o .g A F D W 21 2 . 9 0 332.70 431.20 .000 .000 .000 .005 .002 .004 . 144 .056 .117 mean se c v ( 55) 325.60 63.118 33.576 .000 .000 42.626 .003 .001 42.626 . 106 .026 42.626 g 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W Rep 1 Rep 2 Rep 3 212.90 332.70 431.20 .001 .000 .000 .008 .003 .007 .236 .088 .203 mean se c v (55) 325.60 63 . 1 1 8 33.576 .000 .000 44.119 .006 .001 44.119 . 176 .045 44.119 Rep 1 Rep 2 R ep 3 .003 .001 .003 m ea n se c v ( 55) .002 .001 46.942 1 1 1 1 1 i i 1 1 1 PG/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 Rep 3 1.641 1.577 1.734 mean se c v (55) 1.651 .046 4.782 251 Rep 1 Rep 2 Rep 3 g 02/da.g AFDW NET DAILY METABOLISM g 0 2 / m o .g A F D W 212.90 332.70 431.20 B A F D W D E T R ITUS/m2 GROSS COMMUNITY PR O D U C T I V I T Y g 02/da.g AFDW Rep 1 Rep 2 Rep 3 B A F D W DETRITUS/m2 COMMUNITY RESPIRATION g 02/hr.g AFDW Table A—2 (con’t). Estimates of Community Me tabo1lsra/g AF D W Detritus, Augusta Creek, Michigan. 24 Jnly 1975 (light), 25 July 1975 (dark), NAGEL SITE, STANDARD RUN e A F D W D E T R ITUS/ m 2 NET COMMUNITY PRODUCTIVITY .001 .000 .000 .008 .003 .005 .257 .081 . 154 me a n se c v (55) 265.40 36.454 23.791 .000 .000 53.690 .005 .002 53.690 . 164 .051 5 3.690 AFD W DETRITUS/m2 g 02/hr.g AFDW g 02/da.g AFDW B 0 2 / mo.g A F D W 193.90 288.80 313.50 .001 .000 .000 .016 .009 .007 .498 .292 .232 mean se c v (51) 265.40 36.454 23.791 .000 .000 4 1.006 .011 .003 41 . 0 0 6 .341 .081 41.006 g 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W Rep 1 Rep 2 Rep 3 193.90 288.80 313.50 .001 .001 .001 .018 .008 .010 .562 .261 .296 me a n se c v ( 55) 265.40 36.454 23.791 .001 .000 44.197 .012 .003 44.197 .373 .095 44.198 Rep 1 Rep 2 Rep 3 .002 -.001 .002 me a n se c v (55) .001 .001 170.38 1 P G/R24 « 1 1 1 1 1 1 t GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 Rep 3 1. 129 .893 1.278 mean se c v ( 55) 1 . 100 . 112 17.690 252 Rep 1 Rep 2 Rep 3 g 02/da.g AFDW NET DAILY METABOLISM B 0 2 / m o .g A F D W 193.90 288.80 313.50 e A F D W DETRI T U S / m 2 GROSS COMMUNITY PRODUCTIVITY g 02/da.g AFDW Rep 1 Rep 2 Rep 3 b COMMUNITY RESPIRATION g 02/hr.g AFDW Table A-2 (con’t ) . Estimates of Community Me tabo1Ism/g AFDV Detritus, Augusta Creek, Michigan. 26 November 1974 (light), 27 November 1974 (dark), KELLOGG FOREST, STANDARD RUN e A F D V D E T R I T U S / m2 NET COMMUNITY PRODUCTIVITY .001 .000 .000 .005 .005 .002 . 143 . 140 .051 mean se c v (55) 214.80 47 . 2 4 4 38.095 .000 .000 4 7.006 .004 .001 47.006 .111 .030 47.006 g 02/hr.g AFDV g 02/da.g AFDV g 0 2 / m o .g A F D V Re p 1 Rep 2 Rep 3 .000 .000 .000 .004 .002 .001 .117 .068 .030 214.80 47.244 38.095 .000 .000 6 1.049 .002 .001 61. 0 4 9 .071 .025 61.049 g 02/hr.g AFDV g 02/da.g AFDV g 0 2 / m o .g A F D V Rep 1 Rep 2 Rep 3 120.80 270.10 253.50 .001 .001 .000 .006 .006 .002 . 189 . 166 .063 mean se c v (50 2 1 4. 8 0 47.244 38.095 .000 .000 48.383 .005 .001 48. 3 8 3 . 139 .039 48 . 3 8 3 g 02/da.g AFDV Rep 1 Rep 2 Rep 3 .002 .003 .001 mean se c v (55) .002 .001 48.746 1 1 1 1 1 1 1 1 1 1 PG/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Re p 1 Rep 2 Rep 3 1.619 2.456 2. 100 mean se c v ( 55) 2.058 .242 20.392 253 120.80 270.10 253.50 g A F D V D E T R I T U S / m2 NET DAILY ME T A B O L I S M g 02/mo.g AFDV 120.80 27 0 . 1 0 25 3 . 5 0 mean se c v (JO G ROSS C O M M U N I T Y PRODUCTIVITY g 02/da.g AFDV R ep 1 Rep 2 Rep 3 g AFDV DETRITUS/m2 COMMUNITY RESPIRATION g 02/hr.g AFDV Table A-2 (con't). Estimates of Commnnity Metabo1 Ism/g AFDW Detritus, Augusta Creek, Michigan. 06 February 1975 (light), 05 February 1975 (dark), KELLOGG FOREST, STANDARD RON g A F D W D E T R I T U S / m2 NET COMMUNITY PRODUCTIVITY Rep 1 Rep 2 R ep 3 me a n se c v (5?) 282.00 108.40 53.40 NET DAILY METABOLISM g 0 2 / m o .g A F D W ’!©oo .000 .002 .001 .054 .030 .000 .000 40.242 .002 .000 40 . 2 4 3 .042 .012 40.243 g 02/hr.g AFDW g 02/da.g AFDW g 0 2 / m o .g A F D W Rep 1 Rep 2 Rep 3 .000 178.60 39 5 . 4 0 TTT7TT .000 .000 .002 .001 .053 .022 mean se c v (/?) 191.33 114.32 103.49 .000 .000 5 7.363 .001 .001 57.363 .038 .015 57.363 g 0 2 / h r .g A F D W g 02/da.g AFDW g 02/rao.g A F D W g A F D W D E T R IT U S / m 2 G ROSS C O M M U N I T Y PRODUCTIVITY g 02/da.g AFDW •••••• *178.60 395.40 g A F D W DETRITUS/m2 COMMUNITY RESPIRATION g 0 2 / h r .g A F D W Rep 1 Rep 2 R ep 3 .000 178.60 395.40 .000 .000 .003 .001 • ••••• .077 .040 mean se c v (3) 191.33 114.32 103.49 .000 .OOO 44.958 .002 .001 44.958 .058 .019 44 . 9 5 8 PG/ R 2 4 g 02/da.g AFDW 1 | Rep 1 Rep 2 Rep 3 ...... 1 I 1 1 1 mea n se c v ( R) .001 ,000 1 1 me a n se .000 1 c v (%) .001 .001 G R OSS C O M M U N I T Y PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 Rep 3 1.447 1.771 1.609 . 162 14.241 Table A-2 (con’t). Estimates of Community Metabo1Ism/g AFDV Detritus, Augusta Creek, Michigan. 13 May 1975 (light), 14 May 1975 (dark), KELLOGG FOREST, STANDARD RON e AFDV DETRITUS/m2 NET COMMUNITY PRODUCTIVITY .000 .000 .00O .004 .004 .003 . 117 . 127 .098 mean se c v (JS) 3 55.47 12.839 6 .256 .000 .000 12.724 .004 .OOO 12.724 .114 .008 12.724 g 02/hr.g AFDV g 02/da.g AFDV B 02/mo.g AFDV 3 3 5. 8 0 3 7 9.60 351.00 .000 .000 .000 .003 .002 .001 . 105 .072 .043 mea n se c v (J5) 3 55.47 12.839 6.256 .000 .000 42.191 .002 .001 42.191 .073 .018 42.191 g 0 2 / h r .g A F D V g 0 2 / d a .g A F D V B 02 / m o . g A F D V Rep 1 Rep 2 Rep 3 3 35.80 379.60 351.00 .000 .900 .000 .006 .006 .004 . 180 . 171 . 124 mea n se c v ( ?5) 3 55.47 12.839 6.256 .000 .000 18.821 .005 .001 18.821 . 158 .017 18.821 Rep 1 Rep 2 Rep 3 .002 .003 .003 me a n se c v (55) .003 .000 14.168 1 I t 1 1 I 1 1 1 1 P G/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 Rep 3 1.719 2.371 2.891 me a n se c v C 55) 2.327 .339 25.232 255 Rep 1 Rep 2 Rep 3 g 02/da.g AFDV RET DAILY METABOLISM B 0 2 / m o .g A F D V 335.80 3 79.60 3 5 1. 0 0 e A F D V D E T R ITUS/tn2 GROSS C O M M U N I T Y PRODUCTIVITY g 02/da.g AFDV Rep 1 Rep 2 Rep 3 g AFD V DETRITUS/m2 COMMUNITY R ESP I R A T I O N g 02/hr.g AFDV Table A-2 (con’t). Estimates of Commandty Hetabollsn/g AFDV Detritus, Augnsta Creek, Michigan. 28 July 1975 (light), 29 July 1975 (dark), KELLOGG FOBEST, STANDARD RON g A F D V DETRITUS/in2 WET COMMUNITY PRODUCTIVITY Rep 1 Rep 2 Rep 3 246.30 293.10 278.40 m ea n se c v (%) 272.60 13.818 8.780 B AFDV DETRITUS/m2 COMMUNITY RESPIRATION e 02/mo.g AFDV .000 .000 .000 .005 .006 .004 . 145 . 179 .118 .000 .000 2 0 .669 .005 .001 20.669 . 147 .018 20.669 g 02/hr.g AFDV g 02/da.g AFDV e 0 2 / m o .g A F D V 1 .000 .000 .000 .007 .002 .006 .232 .070 . 194 mean se c v (5S) 272.60 13.818 8.780 .000 .000 51.327 .005 .002 51.327 . 165 .049 51.327 g 02/hr.g AFDV g 0 2 / d a .g A F D V g 0 2 / m o .g A F D V Rep 1 Rep 2 Rep 3 246.30 293.10 278.40 .001 .000 .001 .009 .007 .008 .286 .222 .236 mean se c v (55) 272.60 13.818 8.780 .001 .000 13.604 .008 .001 13.604 .248 .019 13.604 Rep 1 Rep 2 Rep 3 .002 .005 .001 mean se c v (%> .003 .001 72.614 1 1 1 1 1 1 1 1 1 1 PG/R24 GROSS COMMUNITY PRODUCTION/ 24 H R R E S P I R A T I O N Rep 1 Rep 2 Rep 3 1.233 3. 174 1.219 me a n se c v ( ?S) 1.875 .650 59.989 256 246.30 293.10 278.40 g 02/da.g AFDV NET DAILY METABOLISM g 02/da.g AFDV Rep 1 Rep 2 Rep 3 e AFDV DETRITUS/m2 G ROSS C O M M U N I T Y PRODUCTIVITY g 02/hr.g AFDV APPENDIX B Table B-l. S u m m a r y of c o m m u n i t y m e t a b o l i s m estimates DA TES NCP (g 02/'m2. hr) CR . GCP for S M I T H SITE, 1 NCP 24 0c t . 1974 ( 1 is h t ) 25 Oct. 1974 (dark) mean se c v (FI) -.023 .005 33.30© .035 .002 9.370 .012 .004 60.400 1 -.250 1 .055 1 3 8 .300 15 Jan. 16 Jan. mean se c v (r?) -. 007 .000 9.557 .017 .003 28 . 1 6 0 .010 .003 47.601 1 1 1 2 0 Flay 1975 ( 1 ight) 21 M a y 1975 (dark) mean se c v ( F3) -.007 .002 39.875 .019 .006 50.966 IT J u l y 1975 ( 1 ight) 18 J u l y 1975 (dark) mean se c v ( FI) -.015 .002 24.492 .022 .003 26.365 1975 (light) 1973 (dark) A u g u s t a Creek, (g 02^'m2. da ) CR GCP Michigan. RDM .846 .048 9 . 870 . 127 .044 60 . 4 0 0 -.719 .075 18.178 -.066 .004 9.557 .402 .065 28.160 .092 .025 47.601 - . 310 .040 22. 6 0 2 .012 .007 102.00 1 - . 102 1 .023 1 39.875 .460 . 135 5 0 .966 . 181 . 107 102.00 -.279 .029 17.930 .007 .001 3 d .635 1 -.229 ! .032 1 2 4.492 .535 .083 26.865 . 104 .021 35.636 -.431 .063 25.433 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 PG/R24 .153 .053 60.068 .219 .032 25.656 .290 . 180 107.13 .191 .018 15.933 257 Table B-2. Summary of community metabolism estimates for B AVENUE, Augusta Greek, Michigan. DA TES NCP Ig 0 2 / m 2 . h r ) CR GCP 26 Jan. 25 Jan. 1973 (light) 1973 (dark) mean se c v (” ) - .002 .011 .009 0 7 Jane 06 June 1973 (light) 1973 (dark) mean se c v (55) .010 .007 97.456 .029 .003 38.261 . .039 .015 5 2.965 0 7 June 07 June 1973 ( light) 1973 (dark) mean se c v ( S) .010 .007 9 7.456 .028 .003 16.503 .033 .010 37.335 05 J u l y 1973 (light) 0 4 J u l y 1973 (dark) mean se c v (3) -.012 .005 55.711 .030 .009 4 3 .145 0 2 Aug. 1973 (light) 01 A u g . 1973 (dark) me a n se c v (J5) .006 .000 .000 29 Aug. 1973 (light) 3 0 A u g . 1973 (dark) me a n se c v (%) 2 7 Sep. 26 Sep. 1973 (light) 1973 (dark) 05 Nov. 06 Nov. 1 1 1 1 1 1 | NCP (g 02/niC .da) GCP CR NDM 1 PG/R24 1 .322 -.023 .274 .088 -. 186 1 . 146 1 . 101 1 97.456 | 1 . 146 1 . 101 1 97.456 .701 . 190 38.261 .590 .221 52.965 -.111 .031 39.797 1 .816 t .094 1 16.361 .668 .078 16.503 .569 . 150 37.335 -.099 .072 103.05 1 .837 I . 127 1 21.497 .018 .014 106.76 1 -. 175 1 .069 ! 55.711 | .708 .216 4 3.146 .272 .206 106.76 -.436 .010 3.376 1 .326 I . 191 1 82.635 .031 .006 2 7 .739 .037 .006 23.442 1 1 1 .082 .000 .000 .746 . 146 2 7.739 .531 .088 23 . 4 4 2 -.216 .058 38.313 1 1 1 -.008 .004 7 2.853 .025 .096 36.213 .016 .002 17.732 -.110 .057 72.853 .590 . 151 36.218 .217 .027 17.732 - . 373 . 124 47.001 1 .382 1 .052 1 19.091 mean se c v ( J3) -.013 .001 16.971 .036 .093 13.648 .023 .002 11. £61 -. 149 .018 16.97© .858 .083 13.648 .278 .023 11.861 -.580 .059 14.503 1 1 1 .324 .004 1.788 1973 ( 1 ight) 1973 (dark) mea n se c v ( 55) -.005 .091 23.311 .014 .001 6.451 .010 .000 1.459 - . 046 .008 23.311 .342 .016 6.451 .099 .001 1.457 -.243 .017 9.680 1 1 1 .289 .016 7.891 11 Dec. 12 Dec. 1973 ( U c h t ) 1973 (dark) mean se c v (55) -.011 .008 97.029 .010 .001 15.556 -.001 .007 805.49 - . 102 .070 9 7.029 .240 .026 15.556 -.011 .060 805.49 -.251 .086 48.740 1 -.017 1 .248 I 2111.9 2 3 Jan. 2 2 Jan. 1974 (light) 1974 (dark) me a n se c v (55) .090 .090 141.42 .012 .001 9 . 753 .012 .601 8.461 1 1 1 | 1 1 1 | 1 I 1 | 1 1 1 1 1 1 1 .001 .001 141.42 .278 .019 9 . 753 . 113 .007 8.461 -. 165 .012 10.661 1 I 1 I .715 .022 4.438 .407 .004 1.303 Table B - 2 ( c o n ’t) S u m m a r y of c o m m u n i t y m e t a b o l i s m est imates for B AVENUE, DA TES NCP (g 0 2 / m 2 .hr) CR GCP 05 Mar. 1974 < light) 04 M a r . 1974 (dark) mean se c v ( yi) -.001 .003 464.67 .011 .007 09 A p r . 1974 ( 1 ight) 10 A p r . 1974 (dark) mean se c v (J5) -.099 .003 43.339 .026 .001 5.238 .017 .002 14.601 15 M a y 16 Ma y mean se c v (i?) .003 .007 119.30 .021 .004 26.264 .029 .003 13.430 1974 (light) 1974 (dark) 1974 ( 1 ight) 1974 (dark) mean se c v ( yj) -.002 .093 342.32 .022 .001 7.995 .020 .004 32.597 09 Jan. 0 8 Jan. 1975 (light) 1975 (dark) me a n se c v (yj) -. 002 .001 75.235 .010 .003 55.443 .008 .092 53.724 16 Apr. 1975 ( light) 17 A p r . 1975 (dark) mean se c v (ys> .003 .005 116.52 .025 .000 19.036 .033 .007 39.767 14 J u l y 1975 ( 1 ight) 15 J u l y 1975 (dark) me a n se c v (yo -. ooo .091 8 08.93 .039 .001 4.595 .038 .091 3.361 NCP 1 1 - .012 1 .039 1 464.67 1 1 1 | 1 1 1 | 1 I 1 | 1 1 1 1 1 1 1 | 1 1 1 I (g 02/102 .da) GCP CR NDM .266 .075 .002 -.114 .035 43.389 .616 .023 5 . 238 .222 .023 14.601 -.394 .046 16.407 . 115 .097 119.80 .504 .094 2 6 . ?64 .421 .040 13.430 - . 083 . 134 22 8 . 5 8 -.019 .033 342.32 .529 .024 7.995 .223 .042 32.597 -.305 .038 21 . 7 0 0 -.019 .008 75.235 .236 .076 5 5.443 .072 .022 53 . 7 2 4 -. 164 .054 56.763 . 106 .072 116.52 .592 .065 19.086 .437 .100 39 . 7 6 7 -. 155 .056 61.969 - . 002 .009 808.93 .924 .025 4.595 .576 -.349 .017 8.465 .0 1 1 3. 362 1 1 1 ! 1 1 1 1 1 1 | 1 1 1 | 1 1 1 | 1 1 1 | 1 1 1 | 1 1 1 I PG/R24 .283 .362 .051 19.768 .881 .243 39 . 0 5 0 .421 .076 31.147 .306 .017 9 .704 .725 .115 27.531 .623 .010 2.868 259 17 Oct. 18 Oct. 1 1 Angus ta Creek, Michigan. Table B-3. Sumnary of community metabolism estimates for UPPER 43. Augusta Creek. Michigan. DATES RCP (g 02/m2. hr) CR GCP 1 1 NCP (g 0 2 / m 2 . d a ) CR GCP RDM 30 O c t . 1974 ( 1 ight) 31 Oc t . 1974 (dark) mean se c v (75) . 151 .036 33.958 .099 .000 .641 .250 .037 2 0.712 1 1 1.570 1 .377 1 33.958 2 . 384 .011 .640 2.606 .382 20 . 7 1 2 .222 .371 236.25 21 Jan. 22 Jan. mean se c v (75) .020 .005 43.358 .016 .004 43 . 3 1 4 . .036 .009 4 1 .544 1 . 190 1 .048 1 43.358 .386 .097 43 . 314 .344 .083 41 . 5 4 4 -. 042 .032 130.87 27 Ma y 1975 ( light) 2 8 M a y 1975 (dark) me a n se c v (75) . 138 .©46 57.615 .079 .006 12.741 .217 .050 40 . 0 0 3 1 2.064 1 .686 1 57.615 1.904 . 140 12.741 3 . 250 .751 40.008 1.346 .650 83.693 21 J n l y 1975 ( light) 22 J u l y 1975 (dark) mean se c v (75) .077 .033 60.321 . 104 .012 19.982 . 171 .042 34 . 5 6 8 1 1.145 1 .488 1 60.321 1 2 . 502 .289 19.982 2.528 .618 34 . 5 6 8 .288 .408 200.42 1975 ( 1 ight) 1975 (dark) 1 1 PG / R 2 4 1 1 1.092 1 . 155 1 20.085 1 .912 1 .074 1 14.071 1 1 1.683 1 .290 1 29 . 8 2 2 1 1.112 1 .172 1 21.816 1 260 Table B-4, Summary of community metabolism estimates for NAGEL SITE. D A TES NCP (g 02/m2. hr) CR G CP 1 NCP A u g u s t a Creek, Michigan. (g 02/m2.da) CR GCP NDM 1 PG/R24 0 8 Aug. 09 Aug. 1973 (light) 1973 (dark) mea n se c v (R> .234 .067 33.361 .093 .021 31.948 .377 .046 1 7 . ISO 1 4 . 027 1 .950 1 33.361 2.242 .506 3 1 .949 5. 353 .650 17.181 3. 112 1. 157 52. 5 7 3 1 2.586 1 .874 1 47.819 04 Sep. 05 Sep. 1973 ( 1 ight) 1973 (dark) mea n se c v (55) .225 .025 15.623 .no .003 4.035 . .335 .022 9. 175 1 2 . 937 1 .324 1 15.623 2. 636 .076 4.055 4.368 .233 9. 175 1.732 .359 29.315 1 1.661 1 . 155 1 13.207 04 0c t . 1973 ( 1 ight) 03 Oc t . 1973 (dark) mea n se c v (55) . 119 .004 4.991 .055 .004 10.248 . 174 .008 6 . 657 1 i 1 1.383 .049 4.991 1.325 .096 10.248 2.024 .095 6.657 .699 .001 . 148 12 Nov. 13 Nov. me a n se c v (55) .245 .007 4.039 .070 .009 17.513 .315 .016 7.027 1 1 1 2.431 .069 4.039 1.676 .208 17.513 3. 124 . 155 7 . 027 1.448 .052 5. 113 f 1.881 1 . 140 1 10.552 18 D e c . 1973 ( 1 ight) 19 D e c . 1973 (dark) me a n se c v (S) . 107 .031 41.470 .031 .006 27.821 . 138 .037 38.448 ! .976 I .286 ! 4 1.470 .732 . 144 2 7 . 8 20 1.254 .341 38.448 .521 . 197 53.369 1 1.686 1 . 134 1 11.226 31 Jan. 1974 ( light) 01 F e b . 1974 (dark) me a n se c v (55) .042 .0 15 51.961 .024 .003 18.953 .066 .012 26 . 0 2 3 1 .419 1 . 154 i 51.961 .532 .© 7 S 18.953 .661 . 122 26 . 0 2 3 .079 .200 357.31 1 1.185 1 .368 1 43. 8 9 8 20 M a r . 1974 ( 1 ight) 21 M a r . 1974 (dark) mean se c v (55) . 159 .018 16.050 .064 .010 21 . 9 2 0 on o !0 2 8 17.729 I 1.924 I .218 1 16.050 1.525 .236 2 1 . 9 20 2.695 .338 17.729 1. 170 . 101 12.262 1 1 1 23 Apr. 1974 ( 1 ight) 22 A p r . 1974 (dark) mean se c v (55) . 118 .036 43.392 . 139 .012 12.616 .257 .024 13.221 i 1.622 ! .500 1 4 3.592 3 . 336 .298 12.616 3.529 .330 13.221 . 193 .628 4 58.77 1 1.075 1 . 195 1 25.625 24 A p r . 1974 ( 1 ight) 22 A p r . 1974 C dark) me a n se c v (55) . 183 .063 47.643 . 139 .012 12.616 .327 .051 21.997 ! 2.583 1 .87© ! 4 7.643 3 . 336 .298 12.616 4.497 .700 21 . 9 9 7 1. 161 .997 121.43 1 1.378 1 .333 1 34.139 29 M a y 1974 (light) 29 M a y 1974 (dark) mean se c v (55) . 150 .023 2 1 .425 . 116 .093 3 .766 .267 .026 13.724 1 2 . 258 1 .342 1 21.425 2 . 794 .074 3. 767 4.003 .339 13.724 1.210 .314 36.714 1 1 1 1973 (light) 1973 (dark) 1 1 1 1.531 .039 3.603 1.775 .054 4.274 1.430 . 101 9.986 Table B-4 (con’t). Summary of community metabolism estimates for RAGEL SITE. Augusta Greek, Michigan. DATES NCP (g 02/m2. hr) CR . GCP 1 NCP (g 0 2 /m2 .d a ) GCP CR NDM PG/'R24 mean se c v (%) . 133 . 153 .286 1 2.032 3 . 672 4 .362 .690 1. 188 05 Aug. 05 Aug. 1974 (light) 1974 (dark) mean se c v (JI) .207 .044 29.911 .092 .001 2 . 224 .344 1 2.958 1 .626 1 29.911 2 . 2 14 .035 2 . 22 3 4 .923 .011 2. 189 06 Nov. 07 Nov. 1974 (light) 1974 (dark) mean se c v (51) . 134 .033 4 3.164 .041 .012 50.311 . 175 .045 44.700 1 1.353 1 .337 1 4 3 .164 .992 .291 50.811 1.772 .457 4 4.700 .780 . 179 39.773 1.819 . 125 11.868 30 Jan. 31 Jan. 1975 (light) 1975 (dark) mean se c v (51) .027 .003 19.525 .026 .002 16.335 .053 .001 2.605 1 .268 1 .030 1 19.524 .629 .059 16.385 .528 .008 2 . 604 -. 101 .065 111.58 .856 .084 16.946 06 M a y 1975 (liS ht) 07 M a y 1975 (dark) mean se c v ( 51) .081 .026 55.378 .045 .012 48.434 . 126 .038 52.913 1 1.159 1 .371 1 55.378 1.071 .300 4 8 . 434 1.796 .549 52.913 .724 .249 59 . 5 6 8 1.651 .046 4.782 06 M a y 1975 (light) 07 M a y 1975 (dark) mean se c v (51) - . 017 .006 - . 012 -.249 . 134 -. 170 -.304 -1 . 2 6 2 24 J u l y 1975 (light) 25 J n l y 1975 (dark) me a n se c v ( 51) .089 .019 36.387 . 114 .009 14.173 .202 .021 18.267 1 1.307 1 .275 1 3 6.387 2 . 726 .223 14.173 2 .980 .314 18.266 .254 .283 192.76 1. 100 . 112 17.690 1 262 01 J u l y 1974 (light) 01 J u l y 1974 (dark) Table B-5. Summary of conmntnity metabolism es t I m a t e s ,for KELLOGG FOREST, Augusta Creek, Michigan. D ATES NCP (g 02/m2. hr) CR . GCP 1 NCP (g 02/'m2 .da) GCP CR NDM 1 PG / R 2 4 26 Nov. 27 Nov. 1974 ( 1 ight) 1974 (dark) me a n se c v (R> .080 .027 58.486 .019 .004 40.598 .098 .031 5 4.360 1 .754 1 .255 1 58.486 .444 . 104 40.598 .929 .292 54.360 .485 .201 71.763 1 2.058 1 .242 1 20.392 06 Feb. 05 Feb. 1975 (light) 1975 (dark) mean se c v ( 55) .038 .004 14.759 .013 .000 5.676 ■ .052 .003 9.611 1 .386 1 .040 1 14.759 .319 .010 5 . 677 .525 .036 9.611 .001 ! 1.609 1 . 162 1 14.241 13 M a y 1975 ( light) 14 M a y 1975 (dark) mea n se c v ( %) .090 .009 17.110 .035 .008 39.128 . 125 .014 19.795 1 1.311 1 . 130 I 17.110 .834 . 188 39.128 1.816 .208 19.795 .982 .117 20. 6 1 2 1 2 .327 1 .339 1 25 . 2 3 2 2 8 J n l y 1975 ( 1 ight) 29 J u l y 1975 (dark) mean se c v (JS) .089 .014 26.236 .059 .016 46.335 . 148 .004 4 . 407 1 1.302 1 . 197 I 2 6.236 1.414 .378 4 6.335 2 .163 .055 4.407 .748 .344 79.533 1 1.875 1 .650 1 59.989 263 Table B - 6. Summary of community m e t a b o l i s m estimates on a detrital A F D W b asis for Smith Site, Augusta Creek, M i c h i g a n . DateS 24 Oct. 25 Oct. 1974 (light) 1974 (dark) 1975 (light) 1975 (dark) 20 May 1975 (light) 21 May 1975 (dark) 17 July 1975 (light) 18 July 1975 (dark) AFDW G / M ~2 N CP . , G 0 2 /G- 1D" 1 mean S.E. C.V. % 479.83 94.38 34.07 - 0.001 mean S.E. C.V.% 439.87 40.68 16.01 0.000 mean S.E. C.V.% 253.34 98.10 67.07 - 0.001 mean S.E. C.V.% 379.80 62.20 28.36 0.000 81.13 CR G 0 2 /G"*D-1 GCP GO / G ^ D -1 N DM GO^Xrkrl 0.002 0.001 49.73 0.000 - 0.002 0.001 61.17 0.001 0.000 - 0.001 0.000 ••• t• 0.000 34.62 39.83 0.003 0.002 96.04 0.001 0.001 116.40 - 0.001 0.001 0.000 0.000 0.000 • •• * • 0.000 80.97 4.76 10.48 - 0.002 0.001 86.98 - 0.001 0.000 7.085 264 15 Jan. 16 Jan. . Table B - 7. Summary of community m e t a b o l i s m estimates on a detrital A F D W basis for B Avenue Site A u g u s t a Creek, Michigan. Dates AFDW_ G/M NCP -1 -1 G 0 2 /G d ' CR -1 -1 G O 2 /G D GCP -1 -1 G 0 2/G ad NDM g o 2 /g 0.003 0.001 30.46 0.002 0.000 26.20 - 0.001 0.000 40.94 I 1973 (light) mean 1973 (dark) S.E. C.V.% 248.60 5.00 2.84 0.000 29 Aug. 30 Aug. 1973 (light) mean 1973 (dark) S.E. C.V.% 265.40 4.60 2.45 0.000 • 1 • • • 0.002 0.001 33.92 0.001 0.000 15.31 - 0.001 0.000 44.81 1973(light) mean 1 9 7 3 (dark) S.E. C.V.% 215.37 116.12 76.25 - 0.001 - 0.001 87.56 0.006 0.004 85.45 0.002 0.001 84.30 - 0.004 0.002 86.00 05 Nov. 06 Nov. 1973 (light) mean 1973 (dark) S.E. C.V.% 192.10 52.10 38.36 0.000 0.002 0.001 44.26 0.001 0.000 37.00 - 0.001 0.000 47.16 11 Dec. 12 Dec. 1973 (light) mean 1973 (dark) S.E. C.V.% 248.80 55.60 31.60 0.000 0.001 0.000 46.03 0.000 - 0.001 0.001 74.60 23 Jan. 22 Jan. 1974 (light) me a n 1974 (dark) S.E. C.V.% 242.60 25.10 14.63 0.000 0.001 0.000 4.91 0.000 • • • • • - 0.001 0.000 4.00 05 Mar. 04 Mar. 1974 (light) mean 1974 (dark) S.E. C.V.% 215.70 100.70 66.02 0.000 0.002 0.001 - • t • • » • • • • • 27 Sept. 26 Sept. • • • • • • • • • « 0.002 • a * • • 265 02 Aug. 01 Aug. Table B - 7 (cont.). AFDW, G/ M 09 April 1974(light) m e a n 10 April 1 9 7 4 (dark) S.E. C.V.% 320.30 92.00 40.62 0.000 0.002 0.001 35.76 0.001 0.000 53.63 15 May 1974 (light) 16 May 1974 (dark) mean S.E. C.V.% 276.80 46.00 23.50 0.000 0.002 0.000 2.85 0.002 0.000 36.41 17 Oct. 18 Oct. 1 9 7 4 (light) 1 9 7 4 (dark) mean S.E. C.V.% 592.00 39.90 11.67 0.000 0.001 0.000 20.28 0.000 09 Jan. 08 Jan. 1975 (light) mean 1975 (dark) S.E. c.v.% 176.98 54.15 53.00 0.000 0.002 0.001 104.38 0.001 16 April 1 9 7 5 (light) mean 17 April 1975(dark) S.E. C.V.% 144.97 19.96 23.85 0.001 0.001 133.42 0.004 0.001 29.10 14 July 1975 (light) mean 15 July 1975 (dark) S.E. C.V.% 330.2 41.12 21.57 0.000 0.003 NCP -1 -1 G 0 2 /G D • • • • • CR -1 -1 G 0 2 /G D 0.000 20.86 GCP -1 -1 G 0 2 /G D • • • • • NDM g o 2 /g- i e - 0.001 0.000 25.05 0.000 • • • • • - 0.001 0.000 30.34 99.88 - 0.001 0.001 106.34 0.003 0.001 40.49 - 0.001 0.001 79.82 0.002 - 0.001 • * • • • 0.000 23.94 0.000 15.71 266 Dates Table B - 8. Summary of community m e t a b o l i s m estimates on a detrital A F D W basis for Upper 43 R D Site Augusta Creek, Michigan. Dates NDM g o 2/g" NCP -1 -1 G 0 2 /G ad CR -1 -1 g o 2/g GC P -1 -1 g o 2 /g ad 218.35 44.75 28.98 0.008 0.003 59.99 0.011 0.002 29.59 0.013 0.004 48.25 0.001 0.002 197.59 0.001 0.000 16.97 0.002 0.000 29.09 0.001 0.000 19.05 0.000 c.v.% 235.03 35.48 26.15 27 May 1975 (light) 28 May 1975 (dark) mean S.E. C.V.% 179.10 2.53 2.45 0.011 0.004 55.90 0.011 0.001 12.54 0.018 0.004 38.18 0.007 0.004 82.46 21 July 1 9 7 5 (light) 22 July 1975(dark) mean S.E. C.V.% 328.03 34.91 18.44 0.003 0.001 55.54 0.008 0.002 39.57 0.007 0.001 29.11 0.001 • ■«•« 30 Oct. 31 Oct. 1974 (light) mean 1974 (dark) S.E. C.V.% 21 Jan. 1975 (light) mean 22 Jan. 1975 (dark) S.E. 267 AFDW G/ M Table B - 9. Summary of community m e t a b o l i s m estimates on a detrital A F D W basis for Nagle Site Augusta Creek, Michigan. Dates NCP G 0 2 /G CR G02 /G- A D-i GCP G 0 2 /G 1973(light) mean 1 9 7 3 (dark) S.E. C.V.% 164.80 34.10 29.26 0.018 0.002 13.96 0.017 0.004 33.12 0.027 0.004 20.36 0.011 0.000 0.00 04 Oct. 03 Oct. 1973 (light) mean 1973 (dark) s .e ; C.V.% 223.75 3.85 2.43 0.006 0.000 7.42 0.006 0.001 12.67 0.009 0.001 9.08 0.003 0.000 2.29 12 Nov. 13 Nov. 1973 (light) m e a n 1973 (dark) S.E. C.V.% 286.45 47.95 23.67 0.009 0.001 19.73 0.006 0.000 6.29 0.011 0.001 16.79 0.005 0.000 28.61 18 Dec. 19 Dec. 1973 (light) mean 1973 (dark) S.E. C.V.% 297.70 44.20 21.00 0.003 0.000 21.41 0.002 0.000 7.03 0.004 0.001 18.19 0.002 0.000 34.29 31 Jan. 01 Feb. 1974 (light) mean 1974 (dark) S.E. C.V.% 299.50 80.40 37.96 0.002 0.001 81.85 0.002 0.000 19.72 0.002 0.001 60.98 0 .Q00 0.001 235.53 20 M a r c h 1 9 7 4 (light) mean 21 March 1974(dark) S.E. C.V.% 360.40 137.30 53.88 0.007 0.003 67.03 0.005 0.003 71.57 0.009 0.004 68.34 0.004 0.002 64.02 23 April 1974(light) mean 22 April 1974(dark) S.E. C.V.% 321.35 55.95 24.62 0.005 0.001 20.05 0.011 0.003 36.67 0.011 0.001 11.59 0.000 • • • •4 04 Sept. 05 Sept. D-1 D-i NDM GO^G AD 1 268 AFDW, G/M Table B - 9 (cont.). NDM , j g o 2 /g_ d" o > Dates NCP -1 -1 G 0 2/G d 24 A pril 1 9 7 4 (light) m e a n 22 April 1974(dark) S.E. C.V.% 321.35 55.95 24.62 0.008 0.001 24.46 0.011 0.003 36.67 0.014 0.000 2.70 0.003 0.003 113.82 29 M a y 1974 (light) 29 M a y 1974 (dark) mean S.E. C.V.% 230.75 75.65 46.36 0.010 0.002 26.24 0.013 0.004 42.97 0.019 0.004 33.71 0.005 0.000 10.55 01 July 1974(light) (dark) 05 Aug. 1 9 7 4 (light) 05 Aug. 1974(dark) mean 283.00 0.007 0.013 0.015 0.002 mean S.E. C.V.% 235.40 37.88 27.87 0.014 0.000 0.649 0.008 0.001 12.127 0.019 ••• • • 0.011 • •• •• 1974 (light) mean 1974 (dark) S.E. C.V.% 388.23 49.91 22.27 0.003 0.001 29.40 0.002 0.000 30.00 0.004 0.001 28.80 0.002 0.000 33.68 G 0 2 /G d 269 06 Nov. 07 Nov. CR -1 -1 G 0 2 /G d 30 J a n . 1975 (light) 31 J a n . 1975 (dark) mean S.E. C.V.% 393.33 7.79 3.43 0.001 0.000 16.86 0.002 0.000 18.97 0.001 0.000 0.89 0.000 • •• • • 06 M a y 1975 (light) 07 May 1975 (dark) mean S.E. C.V.% 325.60 63.12 33.58 0.004 0.001 45.08 0.003 0.001 42.63 0.006 0.001 44.12 0.002 0.001 46.94 24 July 1 9 7 5 (light) 25 July 1 9 7 5 (dark) mean S.E. C.V.% 265.40 36.45 23.79 0.005 0.002 53.69 0.011 0.003 41.00 0.012 0.003 44.20 0.001 0.001 170.38 Table B - 10. Summary of community m e t a b o l i s m estimates on a detrital A F D W basis for Kell o g g Forest Site Augusta Creek, Michigan. Date AFDW, G/M NCP -1 -1 G 0 2 /G d CR -1 -1 G 0 2 /G d G CP -1 -1 G 0 2 /G d NDM g o 2 /g 1974 (light) mean 1974 (dark) S.E. C.V.% 214.80 47.24 38.10 0.004 0.001 47.01 0.002 0.001 61.05 0.005 0.001 48.38 0.002 0.001 48.75 06 Feb. 05 Feb. 1975 (light) m e a n 1975 (dark) S.E. c.v.% 191.33 114.32 103.49 0.002 0.000 40.24 0.001 0.001 57.36 0.002 0.001 44.96 0.001 0.000 0.000 13 May 1975 (light) 14 May 1975 (dark) mean S.E. C.V.% 355.47 12.84 6.26 0.004 0.000 12.72 0.002 0.001 42.19 0.005 0.001 18.82 0.003 0.000 14.17 28 July 1 9 7 5 (light) 29 July 1 9 7 5 (dark) mean S.E. C.V.% 272.60 13.82 8.78 0.005 0.001 20.67 0.005 0.002 51.32 0.008 0.001 13.60 0.003 0.001 72.61 270 26 Nov. 27 Nov. APPENDIX C Table C— 1. Organic sediment composition (g AFDV) for selected sites on Angnsta Creek, Michigan. 24 October 1974 (light), 25 October 1974 (dark), SMITH SITE, STANDARD RUN DETRITUS g AFD W / m 2 561.7 291.6 586.2 2002. 2725. 1952. 2563. 3016. 2538. me a n se c v (3) 479.8 94.38 34.06 2226. 2 4 9.7 19.43 2706. 155.3 9.94 1m m 250um 75um 0.45nm 11.77 10.65 20.36 57.58 24.51 67.02 198.3 54.41 205.6 122.4 77.74 80.01 169.0 105.9 2 07.0 mean se c v (J5) 9.0 7 4 .74 90.62 14.26 3.06 37.25 49.70 12.88 44.91 152.7 4 9.22 55.81 93.38 14.52 2 6.93 160.6 29.48 3 1.79 4mm 1mm 250um 75 urn Rep 1 Rep 2 Rep 3 7 22.9 7 5 9.0 483.8 30. S B « • • • • • a • • » • • • • • a a a a a 5.56 825.7 839.6 2 2 2.4 58.39 5.56 mean se c v ( JS) 7 7 4.3 51.39 9.33 799.3 40.29 7. 13 353. 1 130.7 5 2 .34 4 4 .48 13.90 4 4.20 5.56 16187 4 mm 1mm 250um 75nm 725.3 . . . . . Rep 1 R ep 2 Rep 3 770.7 5 4 1.3 2 28.8 127.9 a a a a a * a a • • « a a a a • a • a a .......... 831.9 359.9 2 39.4 2 63.9 8 5.57 mean se c v (JI) 7 7 8.7 5 3 .22 9.66 315.3 44.39 7.7 3 4 15.4 125.9 42.88 2 46.4 17. oo 10.07 106.7 21. 19 28.07 271 2.62 18.34 6.26 16mm g T O T A L / m2 BY PA R T I C L E S I Z E 4mm R ep 1 Rep 2 Rep 3 16mm g EPILITH0N/m2 BY PARTICLE SIZE TOTAL Rep 1 Rep 2 Rep 3 16mm g D E T R I T U S / m2 BY PARTICLE SIZE EP1L1THON Table C-l (con’t ) . Organic sediment composition (g AFDW) for selected sites on Angusta Creek, Hichlgan. 15 January 1975 (light), 16 January 1975 (dark), SPIITH SITE, STANDARD RUN DETRITUS g A F D W / m2 / * 51 7.2 423. 1 379.3 1732. 1337. 1868. 2249. 1760. 2247. mean se c v ( %) 439.8 40.68 16.01 1645. 159.2 16.76 2085. 162.7 13.51 1mm 250um 75 um 0.45um 59. 15 15.84 25.70 86.70 50.59 59.66 162. 1 170.5 118. 1 56. 19 6 4 .24 29.84 153.0 121.9 106.6 me a n se c v ( J5) 13. 11 13. 11 173.2 33.56 13. 10 67 .63 65.65 10.84 28.61 150.2 16.24 18.73 50.09 10.38 3 5.92 127. 1 13.65 18.59 4mm 1mm 250um 75ura Rep 1 Rep 2 Rep 3 989.8 422.6 586.6 6 3 6.7 125. 1 191.8 2 7.80 7 5.07 2.78 11. 12 mean se c v (J5) 70 6.2 283.6 56.79 611.6 25.04 5.79 158.4 33.35 29.76 5 1.43 2 3 .63 64.98 6.95 4. 17 84.85 4mm 1mm 250um 75 u m ReT> 1 Rep 2 Rep 3 989.8 422.6 6 4 o .* 652.5 211.8 2 4 2.3 189.9 245.5 58.97 75.36 mean se c v ( 7S) 706.2 283.6 56.79 649. ! 3.3 3 .73 227. 1 15.29 9. 5 2 2 1 7.7 2 7.03 18.07 67. 16 8. 19 17.25 272 .00 .00 39.34 16mm g T O T A L / m2 BY PAR T I C L E SIZ E 4mm Rep 1 Rep 2 Rep 3 16mm g EPILITHON/ra2 BY PARTICLE SIZE TOTAL Rep 1 Re p 2 Rep 3 16mm g DETRITUS/m2 BY PARTICLE SIZE EPILITHON Table C— 1 (con't). Organic sediment composition (g AFDV) for selected sites on Angusta Creek, Hichlgan, 20 May 1975 (light), 21 May 1975 (dark), SMITH SITE, STANDARD RUN DETRITUS g AFDW/m2 86.63 42 6.3 247. 1 1170. 1387. 1885. 1256. 1813. 2132. me a n se c v (J5) 2 5 3.3 98. 10 6 7 .07 1480. 211.6 24.75 1734. 255.8 25.55 Rep 1 Rep 2 R ep 3 lmro 250um 75um 0.45um » « ■ • • * 4 • • * .00 .00 47.08 5.67 44.04 17.67 6 4.27 70.35 93.06 54.76 177! 8 98.63 .00 26.37 20.70 111.0 3 0.85 13. 18 60.43 67.31 3.04 6.38 73.91 19. 15 36.64 138.2 39.58 4 0 .50 16mm 4mm 1nun 250um 75 u m • • • • • Rep 1 Rep 2 Rep 3 380.9 6 3 6.7 2 B 9 .1 392.2 425.4 1348. 144.6 247.4 144.6 5 2.82 72.29 69.51 .00 5.56 3 3.36 mean se c v (75) 4 35.5 104.0 41.35 7 88.5 283.8 62.34 178.8 3 4 .26 33. 18 64.87 6.08 16.23 12.97 10.31 137.7 4mm 1mm 250um 75um 16mm g T O T A L / m2 BY P A R T I C L E SIZ E 4mm Rep 1 Rep 2 Rep 3 6 3 6.7 289. 1 472! 4 1353. 291! 4 162.2 136.5 139.8 9s! 62 88. 12 mean se c v ( 75) 462.9 173.8 53.09 9 1 3.3 440.6 68.24 226.8 64.58 40.26 138.2 1.64 1.68 9 3 . 37 5.25 7.95 273 mean se c v ( 75) g EPILITH0N/m2 BY PARTICLE SIZE TOTAL Rep 1 Rep 2 Rep 3 16mm g D E T R I T U S / m2 BY PARTICLE SIZE EPILITHON Table C-l (con't). Organic sediment composition (g AFDV) for selected sites on Angusta Creek, Michigan, 17 July 1975 (light), 18 July 1975 (dark), SMITH SITE, STANDARD RON DETRITUS 497.5 286.1 355.8 1743. 2002. 1362. 2240. 2288. 1717. mean se c v (5S) 379.8 62. 19 28.36 1702. 185.8 18.91 2082. 182.6 15. 19 1mm 250nra 75 u m 0.45um 77777 "i! 99 27.35 15.81 31. 12 4 7.7 0 5 7.33 55.54 8 7 .92 • • r• • 162.1 138.6 mean se c v (J5) 6.7 3 6.7 3 141.4 16. 17 11. 17 9 7.72 23.46 7.65 46. 13 52.51 4.81 12.96 71.73 16. 19 31 .92 150.3 11.75 11.05 4mm 1mm 250um 75 u m Rep 1 Rep 2 Rep 3 525.5 586.6 172.4 784. 0 853.5 697.3 3 3 6.4 3 6 1.4 344.7 91.75 180.7 136.2 5.56 19.46 11. 12 mean se c v (51) 428. 1 129.0 5 2 .22 778.4 45.03 10.02 347.5 7.35 3.66 136.2 2 5.67 32.65 12.04 4.03 5 8.07 4mm 1mm 250um 75um Rep 1 Rep 2 Rep 3 * • • •« 586.6 185.8 .777. .777. 853.5 725. 1 3 7 7 .2 3 7 5 .8 2 28.4 193.5 7 5 .0 0 .7777 mean se c v (J.) 386.2 2 0 0.3 7 3.36 791.8 66.67 11.90 376.5 .70 .26 210.9 17.43 11.68 87.02 12.02 19.53 99.04 274 .00 13.46 16mm g T O T A L / m2 BY PARTICLE SIZE 4mm R ep 1 R ep 2 R ep 3 16mm g EPILITH0N/m2 BY P A R T I C L E SIZ E TOTAL Rep 1 Rep 2 Re p 3 16mm g DETRITUS/'m2 BY PARTICLE SIZE EPILITHON Table C-l. Organic sediment composition (g AFDW) for selected sites on Angnsta Creek, Michigan. 17 October 1974 (light), 18 October 1974 (dark), B AVENUE, STANDARD RUN DETRITUS g AFD W / m 2 623.9 6 3 9.4 512.7 2049. 2452. 2260. 2672. 3091. 2772. mean se c v (75) 5 9 2.0 3 9 .90 11.67 2253. 116.3 8. 9 4 2845. 126.2 7.68 1mm 250nm 75 n n 0.45um Rep 1 R ep 2 R ep 3 .00 .00 .00 32. 16 5 5 .90 55.09 127.6 166.7 79.75 165.0 103.4 126.2 122.6 143.5 120.4 176.7 170.8 131.3 mean se c v (55) .00 47.41 7.62 27.86 124.6 25. 14 3 4 .92 131.5 17.98 2 3 .67 128.8 7.36 9.89 159.6 14.25 15.46 16mm 4mm 1mm 259um 75um Rep 1 Rep 2 Rep 3 925. B 8 2 5.7 180.7 111.2 5.56 ■ • • « • • • • • • • • » • ■ • • • • • • • • • • 1251. 786.8 139.0 8 0 .63 2. 7 8 mean se c v (75) 1088. 162.6 21. 12 80 6.2 19.44 3.41 159.8 20.85 18.44 95.91 15.28 2 2 .53 4.17 1.39 47. 14 4mm limn 25Qnm 75 a m 128. 1 16 m m g T O T A L / m2 BY PARTICLE SIZE 4mm Ren 1 Rep 2 Rep 3 925.8 857.0 308.3 2 76.2 • ■ • • . • • • • • * - . • * .......... 1251. 841.8 218.7 206.8 123. 1 mean se c v (75) 1088. 162.6 21. 12 849.8 7.98 1.32 263.5 44.77 24.02 241.5 3 4 .68 20.31 125.6 2.49 2.80 .......... 275 g EP I L I TH O N / m 2 BY PARTICLE SIZE TOTAL Rep 1 Rep 2 Rep 3 16mm g DETRITUS/m2 BY PARTICLE SIZE EPILITHON Table C— 1 (con’t). Organic sediment composition (g AFDV) for selected sites on Angnsta Greek* Michigan. 09 January 1975 (light), ©8 January 1975 (dark), B AVENUE, STANDARD RUN D ETRITUS g AF D W / m 2 166.9 88.63 275.4 1646. 2369. 1910. 1812. 2457. 2185. mean se c v ( «) 176.9 54. 15 52.99 1975. 2 11.2 18.52 2152. 186.8 15.04 1mm 250um 75um 0.45um Rep 1 Rep 2 Rep 3 .00 .00 .00 6 .6 2 9. 10 55. 17 15.84 12.79 28.31 21.88 11.07 40.39 57.69 8.67 64.42 64.90 46.99 87.07 mean se c v (55) .00 2 3 .63 15.78 115.7 18.98 4.74 43 .32 2 4 .44 8.56 60.65 43.59 17.56 69.80 66 .32 11.59 30 .27 16mn 4mm 1mm 250um 75um Rep 1 Rep 2 Rep 3 5 1 9.9 1082. 109O. 8 6 1.9 422.6 650.6 155.7 725.6 114.0 105.6 139.0 52.82 2. 7 8 .00 2.78 mea n se c v (55) 8 97.3 188.7 36.42 645.0 126.8 34.06 33 1.7 197.2 103.0 99. 14 25.08 4 3.82 1.85 .92 8 6.60 4mm 1mm 250um 75 um 16mm g T O T A L / m2 BY PARTICLE SIZE 4mm Re p I Rep 2 Rep 3 519.9 1082. 1090. 868.5 431.7 7 0 5.7 171.5 7 3 8.3 142.3 127.4 150.0 93.21 60.47 8. 6 7 67.20 mean se c v (55) 897.3 183.7 3 6 .42 663.6 127.4 33.01 3 5 0.7 194.0 95 .80 123.5 16.52 23. 16 45 .44 18.49 70.46 276 g E P I L I T H 0 N / m2 BY PARTICLE SIZE TOTAL Rep 1 Rep 2 Rep 3 16mm g D E T R I T U S / m2 BY , PA R T I C L E S I Z E EP I L I T H O N Table C - 1 (con’t). Organic sediment composition (g AFDV) for selected sites on Angusta Creek, Michigan, 16 April 1975 (light), 17 April 1975 (dark), B AVEHUE, STANDARD RUN DETR I T U S g A FD W / m 2 135.8 115.9 163.2 2508. 1777. 2891. 2643. 1892. 3074. mean se c v ( 218.3 44.75 28.98 EPILITHON TOTAL 4409. 5260. 4672. 5433. 4834. 425.5 12.44 5052. 380.7 10.65 • 16mm g DETR I T U S / m 2 BY PARTICLE SIZE 250um 75um 0.45um 34.68 9.51 39. 19 17.30 5 1 . 32 38.81 77.51 36.55 60.42 71.41 mean se c v (55) .00 22 .09 12.58 3 0 .54 28.24 10.94 5 4 .80 45.06 6.25 19.62 57.03 2 0.48 50.78 65.91 5.49 11.78 4mm lram 250um 75um 4624. 514.3 5 5.60 55.60 11. 12 4624. 514.3 55.60 55.60 11. 12 4mm 1mm 250um ?5um 4624. 523.0 7 2 .90 94.41 47.67 4624. 523.8 72.90 94.41 4 7 .67 Rep 1 Rep 2 16mm Rep 1 Rep 2 mean se c v ( 55) 279 .00 .00 mean se c v ( 55) g T O T A L / m2 BY P A RT I C L E S I Z E 1mm Re p 1 Rep 2 16mm g EPILITH0N/m2 BY P A R T I C L E SI Z E 4mm Table C-l (con’t). Organic sediment composition (g AFDV) for selected sites on Augusta Greek, Michigan. 21 January 1975 (light), 22 January 1975 (dark), UPPER 43, STANDARD RUN DETRITUS g AFDW / m 2 Rep 1 Re p 2 Rep 3 mean se c v (JS) 16mm g D E T R I T U S / m2 BY PARTICLE SIZE 301.4 180.1 223.6 2669. 4 4 07. 2970. 4587. 223.6 23 5.0 35.48 2 6 .14 3538. 2593. 1273. 85.05 ...... 4mm 1mm 250um 75um 0.45um 33.36 3.38 6.4 7 5 3.37 12.48 18. 11 67.43 3 9.62 45.29 64.58 54.91 7 0 .53 82.60 69.68 83. 19 m ea n se c v (%) .00 14.40 9.5 1 114.4 27.98 12.79 79. 18 50.78 8.48 2 8.93 63.34 4.55 12.44 78.49 4.40 9.72 4mra 1m m 25Gum 75um 1588. 3281. ..... 5 7 8.3 639.3 ..... 166.3 425.4 ..... 305.8 5 2 .82 ..... 11. 12 8.34 ..... 2434. 846.5 49. 17 6 0 8.9 3 0 .60 7. 10 305.8 119.5 55.27 179.3 126.4 99.76 9.73 1.38 20. 19 4mm 1mm 250nm 75um 1588. 3281. ..... 611.6 642. 8 ..... 239.6 437.8 ..... 3 73.2 9 2.44 ..... 7 5 .70 63.25 ..... 2434. 846.3 49. 17 627.2 15.61 3.51 3 3 8.7 99. 10 4 1 .37 232.8 140.4 8 5.27 6 9 .47 6.22 12.67 Rep 1 Rep 2 Rep 3 16mm Rep 1 Rep 2 Rep 3 mean se c v (71) 280 .00 .00 .00 mean se c v ( f.) g TOTAL/ra2 BY PARTICLE SIZE TOTAL Rep 1 Rep 2 R ep 3 16mm g EPILITH0N/ra2 BY PAR T I C L E S I Z E EPILITHON Table C-l (con’t). Organic sediment composition (g AFDW) for selected sites on Angusta Creek, Michigan. 27 May 1975 (light), 28 May 1975 (dark), UPPER 43, STANDARD RUN D ETR I T U S g AFD W / m 2 182.3 180.9 174. 1 2992. 608.9 1896. 3174. 789.8 2070. me a n se c v ( JI) 179. 1 2.53 2.44 1832. 688.6 65. 10 2011. 688.9 59.32 Rep 1 Rep 2 Rep 3 4mm 1mm 2 5 0 um ?5um 0.45um .00 .00 19.07 32 .70 27.03 28.50 2 4 .72 41.34 28.71 5 6 . 92 81.35 14.67 .00 25.88 6.31 3 7 .23 27.76 .73 3. 7 4 3 3 .03 8.31 35.58 42.81 14. 10 46.59 48.01 33.34 98.20 4mm 1mm 250um 75um 16 m m Rep 1 R ep 2 Rep 3 2344. 405.9 1499. 408.7 47.26 205.7 2 0 0.2 133.5 172.4 36. 14 2 2.24 19.46 2.78 .00 .00 m ea n se c v (%) 1416. 5 6 1.0 6 8.60 220.3 104.6 82. 14 168.7 19.34 19.86 2 5.94 5. 15 3 4 .44 .92 .92 173.2 4mm 1mm 250um 75nm 16 mm g T O T A L / m2 BY PARTICLE SIZE Rep 1 R ep 2 Rep 3 4 05.9 1499. 66! 33 238.4 160.5 200.9 46! 96 6 0.80 2 b !71 56.92 mean se c v (%y 9 5 2.4 5 46.5 81. 15 152.0 86.93 79.85 180.7 20. 18 15.79 5 3 .88 6.92 18. 16 42.81 14. 10 46.59 281 mean se cv ( g EPILITH0N/m2 BY PARTICLE SIZE TOTAL Rep 1 Re p 2 Re p 3 16mm g D E T R I T U S / m2 BY PA R T I C L E S I Z E EPILITHON Table C-l Ccon’t). Organic sediment composition (g AFDV) for selected sites on Augusta Creek, Michigan. 21 July 1975 (light), 22 July 1975 (dark), UPPER 43, STANDARD RUN DETRITUS g AFDW/m2 347.2 3 7 6.6 26 0.3 5438. 2330. 4323. 5785. 2706. 4583. me a n se c v (55) 328.0 34.91 18.43 4030. 9 0 9.0 39.06 4358. 8 95.8 35.60 1mm 250um 75 u m 0.45um 86.28 ..... 38.01 15.20 29.20 67.89 2 0 1.0 ••• •• 140. 1 .00 5.49 2.42 6 2 .37 16.99 1.79 14.89 3 3.60 4.40 18.53 77.08 9. 19 16.86 170.5 30.45 25.24 16mm 4mm 1mm 250nm 75um • • • .00 ■ • • • ..... ..... Rep 1 Rep 2 Rep 3 4023. 1073. 3873. 989.8 861.9 283.6 35 8.7 339.2 122.3 5 2 .82 44.48 4 1 .70 13.90 11. 12 2. 7 8 mean se c v (55) 2989. 9 5 9.3 5 5 .57 7 1 1.7 2 1 7.2 52.86 2 7 3.4 75 .75 4 7 .99 4 6.33 3. 3 4 12.49 9.26 3.34 62.45 4mm 1mm 256um 75um Rep I Rep 2 Rep 3 4023. ..... 3873. 992.8 377.4 90.83 ..... ..... 291.5 137.5 7 0.90 7 0 .67 mea n se c v ( 3) 3948. 75 .00 2.68 642. 1 350.6 77.22 2 57.4 119.9 6 5 .90 80.80 9.96 17.42 85.42 14.75 24.42 ..... ..... ieo. i 282 3.0 7 • 7.91 .00 •• 16mm g T O T A L / m2 BY P A R T I C L E SI Z E 4mm 00 1N 1 • 1 (0 Rep 1 Re p 2 Rep 3 mean se c v (J!) g EPILITH0N/m2 BY P A R T I C L E SIZ E T OTA L Rep 1 Rep 2 Rep 3 16mm g D E T R I T U S / m2 BY PARTICLE SIZE EPILITHOH 4 Table C - 1 (con’t). Organic sediment composition (g AFDV) for selected sites o n Augusta Greek, Michigan. 06 November 1974 (light). 07 November 1974 (dark). NAGEL SITE. STANDARD RON DETRITUS g AFDW/ra2 g DETRITUS/m2 BY PARTICLE SIZE Rep 1 Rep S Re p 3 312.3 482.3 370. 1 2808. 2224. 3951. 3120. 2706. 4321. mean se c v ( J5) 38 8.2 49 .90 22.26 2994. 507. 1 29 .33 3382. 4 8 4 .2 24.79 16ram 4mm 1mm 250um 75ttm 0.45am Rep 1 Rep 2 Re p 3 .00 .00 .00 5 1 .97 52.20 4 0 .57 33.98 8 1 .07 41.67 6 2 .60 151.8 128.3 73.87 88.02 6 6.62 89.91 109.2 92.96 me a n se c v (55) .00 48.24 3. 8 3 13.78 52 .24 14.58 48 .35 114,2 26.fe9 40.47 76. 17 6. 2 8 14.28 97.35 5. 9 8 10.65 16mm 4mm 1mm 250am 75am Rep 1 R ep 2 R ep 3 1607. 3292. 547.7 417.0 16.68 136.2 5 2 .62 105 :.6 !oo mean se c v ( 55) 2449. 842.5 48.64 482.3 65.35 19. 16 76.44 59.76 110.5 79.21 26.39 47.11 .OO 4mm 1mm 250nm 75um 16mm g T O T A L / m2 BY PARTICLE SIZE TOTAL .00 Rep 1 Rep 2 Rep 3 .1607. 3292. .777. .777. 77777 5 99.9 457.5 97.75 177.8 2 0 4.6 2 3 3.9 8 8.02 6 6.62 mean se c v (3) 2449. 842.5 48.64 5 2 8.7 71. 16 19.03 137.8 40.06 41. 11 2 1 9 .2 14.64 9.44 7 7.32 10.70 19.57 283 g EPILITH0N/m2 BY P AR T I C L E SI Z E EPILITHON Table C-l (con’t). Organic sediment composition (g AFDV) for selected sites on Augusta Greek, Michigan. 30 January 1973 (light), 31 January 1975 (dark), NACEL SITE, STANDARD RUN DETRI T U S g AFDV/m2 384.0 408.8 3 8 7.2 4145. 4946. 2967. 4529. 5354. 3354. mean se c v ( B) 3 9 3.3 7.78 3.43 4019. 574.7 24.76 4412. 5 8 0.4 22.78 1mm 250am 75um 0.45um 32.89 9 0.93 117.8 6 7.97 7 4.83 60. 14 109. 1 77.55 8 1 .72 86.41 63.48 3 6 .47 87.65 102.0 5 7.47 mean se c v (J?) 11. 19 11. 19 173.2 80.54 25.05 53.88 6 7 .64 4.24 10.86 89.45 9.89 19. 15 62. 12 14.43 4 0.24 82.37 13. 12 27.59 4mm 1mm 250um 75 u m Rep 1 Rep 2 Rep 3 3675. 4607. 2638. 3 5 8.7 166.3 166.8 4 1 .70 8 0.63 63.95 63.95 91.75 8 8 .97 5.56 .00 8.34 mean se c v ( ") 3640. 568.6 27.06 230.7 6 3 .96 48.01 62.09 11.27 31.45 81.55 8. 3 4 18.77 4.63 2.45 91.65 4mm 1mm 250um 75um Rep 1 Rep 2 Rep 3 3675. 4607. 2671. 39 1.5 257.7 2 34.6 109.6 155.4 124.0 173.0 169.3 170.6 9 1 .97 63.48 44.81 mean se c v (P.) 3651. 5 5 3.8 26.51 311.3 40. 3B 22.74 129.7 13.51 18.04 171.0 1.09 1.11 66.75 13.71 35.57 284 .00 .00 33.58 16mm g T O T A L / m2 BY PA R T I C L E S I Z E 4mm Rep 1 Rep 2 Rep 3 16mm g EPILITH0N/ra2 BY PARTICLE SIZE TOTAL Rep 1 Rep 2 Rep 3 16mm g DE T R I T U S / m 2 BY P AR T I C L E S I Z E EPILITHON Table C-l (con’t). Organ1c sediment composition (g AFDV) for selected sites on Augusta Greek, Michigan. 06 May 1975 (light), 07 May 1975 (dark), NAGEL SITE, STANDARD RUN DETR I T U S g AFDW/m2 212.9 332.7 43 1.2 4849. 5421. 4846. 5061. 5753. 5277. mean se c v (5S) 325.6 63. 11 33.57 5038. 191. 1 6.57 5364. 204.3 6.60 Rep 1 Rep 2 Rep 3 4mm 1mm 250um 75um 0.45um .00 7 0.32 33.53 8.03 25.68 18. 10 4 7.44 47.06 35.78 99.35 7 0.52 89.81 35. 16 35. 16 141.4 20.78 12.74 86.72 2 1.89 3.79 24.48 47.25 . 19 .56 67.56 31.78 66 .53 80. 16 9. 6 4 17.01 4mm 1mm 250um 75um 16mm Rep 1 Rep 2 Rep 3 4104. 4921. 4265. 5 61.6 358. 7 4 3 6.5 6 3.95 83.41 4 7.26 114.0 5 5 .60 9 4 .53 5.56 2.78 2.78 mean se c v (%) 4430. 249.3 9.76 452.2 59. 10 22.63 64.87 10.44 27.88 8 8 .04 17. 16 3 3 .77 3.70 .92 4 3.30 4mm 1mm 250um 75um 16mm g T O T A L / m2 BY PAR T I C L E S I Z E Rep 1 Rep 2 Rep 3 4104. 4991. 595. 1 3 6 6.7 3 9.63 101.5 161.4 102.6 4 1.34 102. 1 m ea n se c v ( ") 4547. 443.6 13.79 4 89.9 I 14.2 33.58 9 5.57 5.9 4 8.79 132.0 2 9.39 3 1.47 7 1.73 30.39 59.92 285 mean se c v (3) g EPIHTH0N/m2 BY PA R T I C L E S I Z E TOTAL Rep 1 Rep 2 Rep 3 16mm g DETR l T U S / m 2 BY PARTICLE SIZE EPILITHON Table C— 1 (con’t). Organic sediment composition (g AFDV) for selected sites on Augusta Creek, Michigan. 24 July 1975 (light), 25 July 1975 (dark), BAGEL SITE, STANDARD RUN DETRI T U S g AFDW / m 2 g D E T R I T U S / m2 BY PARTICLE SIZE 193.9 288.8 313.5 3286. 1151. 1501. 3479. 1439. 1814. mean se c v (?S) 265.4 36.45 23.79 1979. 661. 1 57.85 2244. 626.9 4 8 . 37 Re p 1 Rep 2 Rep 3 16 m m 4mm 1mm 250um 75um 0.45um ’ !©o 25.61 16.86 6.51 36.80 26.38 98.68 67.27 77.08 163! 0 8 3.85 .00 2 1 .23 4.37 29. 13 21.65 15. 14 9 8 .89 6 2.63 36.25 81.85 72.17 4.90 9.61 123.4 3 9.57 4 5.34 16mm 4mm 1mm 250um 75um •esse .90 (n) Rep 1 Re p 2 Re p 3 2402. 1029. 8 0 3 .o soa.a 52.82 58 6.6 316.9 33.36 80.63 5 2 .82 30.58 27.80 5.56 5.56 2.78 mean se c v ( %) 1 4 1 1. 499.5 61.29 382.7 166.4 75.34 143.6 87.70 105.7 37.06 7.91 36.99 4.63 .92 3 4 .64 4mm 1mm 250um 75ntn 16ntm Rep 1 Rep 2 Rep 3 1029. 803.5 78 .43 603.4 39.87 117.4 56! 96 126.6 72! 83 79.86 me a n se c v (%) 916.2 112.7 17.40 34 0.9 262.3 108.8 78.65 3 8.77 6 9.72 9 1 .82 34.86 53.69 76.34 3.51 6.51 286 CV g T O T A L / m2 BY PARTICLE SIZE TOTAL Rep 1 Rep 2 Rep 3 mean se g EPILITH0N/m2 BY PAR T I C L E S I Z E EPILITHON Table C - 1 (con’t). Organic sediment composition (g AFDV) for selected sites on Augusta Greek, Michigan. 26 November 1974 (light), 27 November 1974 (dark), KELLOGG FOREST, STANDARD RON DETR I T U S g AFDW/m2 120.8 270. 1 253.5 4429. 2513. 3220. 4549. 2783. 3473. me a n se c v (3) 214.8 47.24 38.09 3387. 5 59.3 2 8 .60 3602. 51 4.0 24.71 1mm 250am 75am 0. 4 5 a m Rep 1 Rep 2 Rep 3 .00 .00 .00 3.48 12.84 23.05 11.59 21.62 15.80 14.66 8 0 .22 3 1 . 20 39.81 48.03 102. 1 5 1 . 26 107.4 8 1.32 mean se c v ( 55) .00 13. 12 5.64 7 4 .35 16.33 2. 9 0 3 0 .82 4 2 . 02 19.68 81. 12 63.31 19.53 5 3.45 7 9.9 9 16.22 35. 12 16um 4mm 1mm 250am 75am Rep 1 Rep 2 Rep 3 •« •• « 1248. 1799. 1051 ! 870.2 15o! 1 364.2 5 2.82 177.9 11. 12 8.34 mean se c v ( 55) 1323. 275.5 2 5.57 960.6 9 0 .40 13.30 257. 1 107.0 58.87 115.3 62.54 7 6.66 9.73 1.38 20. 19 4mm 1mm 250am 75am 16mm g T O T A L / m2 BY P A R T I C L E SI Z E 4mm Rep 1 Rep 2 Rep 3 1248. 1799. 1063. 8 9 3.2 171.7 3 8 0.0 133.0 209. 1 59. 15 110.4 mean se c v (55) 1523. 275.5 2 5.57 978.5 85.29 12.32 275.8 104. 1 53.38 171.0 38.03 31.43 84.79 2 5.64 4 2.77 287 g EPILITH0N/m2 BY PARTICLE SIZE TOTAL Rep 1 Rep 2 Rep 3 16mm g DETRITUS/m2 BV PARTICLE SIZE EPILITHON Table C-l (con’t). Organic sediment composition (g AFDV) for selected sites on Angusta Creek., Michigan. 13 May 1975 (light), 14 May 1973 (dark), KELLOGG FOREST, STANDARD RON DETRITUS g AF D W / « 2 335. B 379.6 351.0 1105. 2174. 1579. 1440. 2553. 1930. mean se c v (?S) 355.4 12.83 6.25 1619. 3 0 9.2 33.07 1974. 322.0 28 .24 Rep 1 Rep 2 Rep 3 1mm 250um 75 u m 0. 4 5 u m 7.39 29.03 8 5.93 74.65 ..... .... .... .00 12.91 43.79 130. 1 64.82 138.8 ..... 99.32 .00 lO. 15 2.75 38.39 36.41 7.38 28.66 108.0 2 2.08 28.91 6 9.73 4.91 9.96 119.0 19.74 2 3.44 16mm 4mra 1mm 25Gum 75ura .00 ••• • • •• » • • Rep 1 Rep 2 Rep 3 4 7 1.3 1207. 1065. 508.8 825.7 350.3 97.31 83.41 125. 1 2 7 .80 5 0.04 36. 14 .00 8.34 2. 7 8 mea n se c v (JJ) 914.4 2 25.3 42.68 561.6 139.7 43. lO 101.9 12.25 2 0 .82 37.99 6. 4 8 29 .57 3.70 2.45 114.5 4mm 1mm 250um 75um 1065. 516.2 •9*• • 3 6 3.2 126.3 •• •• • 168.8 113.7 •• • • « 166.2 74.65 • • ■• • 67.60 768. 1 296.8 54.63 439 •7 76.49 24.60 147.6 21.27 2 0 .38 139.9 26.25 26.52 71. 12 3.52 7.00 16irm g T O T A L / m2 BY PAR T I C L E S I Z E 4mm Rep 1 Rep 2 Rep 3 mean se c v (?•) 47 1.3 ..... 288 mean se c v (JS) g EPILITH0N/m2 BY PARTICLE SIZE TOTAL Rep 1 Rep 2 Rep 3 16mm g D E T R I T U S/ m 2 BY PAR T I C L E S I Z E EPILITHON Table C— 1 (con’t). Organic sediment composition (g AFDV) for selected sites on Augusta Greek, Michigan. 06 February 1975 (light), 05 February 1975 (dark), KELLOGG FOREST, STANDARD ROW EPILITHON TOTAL *178.6* 395.4 2241. 3175. 2419. 3570. 287.0 1G8.4 53.4 2708. 46700 24.38 DETRITUS g AFDW / m 2 Rep 1 Rep 2 Rep 3 mean se c v (JJ) 16mm g DETRITUS/m2 BY PARTICLE SIZE Rep 1 R ep 2 Rep 3 75um 0.45um .00 7.68 20.06 2 7.29 5 7 .48 109.8 45. 11 109.8 56.00 140.8 .00 j 3. 8 4 3. 8 4 141.4 2 3 .67 3.61 21.59 8 3.64 26. 16 4 4 .23 7 7.45 3 2.34 5 9.05 98.40 4 2.4 0 60.93 4mm lntm 250nm 75um Rep 1 Rep 2 Rep 3 1479! 2388. 4 94.9 614.4 183! 5 91.75 8 3 ’41 7 2.29 .00 8.34 mean se c v .(JJ) 1933. 454.5 33.24 554.6 59.75 15.23 137.6 4 5.87 47. 14 7 7.85 5.56 10. 10 4. 17 4. 17 141.4 4mm 1mm 250um 75 um 16mm g T O T A L / m2 BY P A R T I C L E SIZ E 250um .00 .00 16mm g EPlLlTHOW/'m2 BY P A R T I C L E SIZ E 1mm Rep 1 Rep 2 Rep 3 • * * * * 494.9 6 2 2.0 .777. 203.5 119.0 • • • • • 1479! 2388. 140.8 102.0 45. 11 118. 1 mean se c v (5i) 1933. 454.5 33.24 5 5 3.4 63.39 16. 10 161.3 42.26 37.05 161.4 20.60 18.04 8 1 .62 36.51 63.26 289 mean se c v (3) 4mm 2994. 575.5 27.17 Table C - 1 (con’t). Organic sediment composition (g AFDV) for selected sites on Augusta Creek, Michigan. 28 July 1975 (light), 29 July 1975 (dark), KELLOGG FOREST. STANDARD RUN EPILITHON TOTAL 246.3 293. 1 278.4 2789. 1474. 4696. 3035. 1767. 4974. 272.6 13.81 8.78 2986. 935.3 54.24 3258. 932.5 49.56 DETRITUS g AFDV/'m2 Rep 1 Rep 2 Rep 3 mean se c v ( a) g DETR.ITUS/m2 BY PARTICLE SIZE Rep 1 Rep 2 Rep 3 4mm .00 .00 12.07 .87 .00 "77777 77777 110.0 97.48 6. 4 7 5.59 12°. 2 11.79 .71 8.57 108. 1 .79 1.04 5 5.62 4.44 11.30 103.7 6.26 8.53 4mm 1mm 250um 75um 586.6 1721. 1176. 689.5 2619. 355.9 164.0 322.5 4 1 .70 33.36 30.58 2.78 .00 2.78 1173. 328.0 4 8 .43 1494. 57 9.3 67. 13 280.8 59. 19 36.51 35.21 3.34 16.43 1.85 .92 86.60 4mm lmm 250um 75ura 16 inn g T O T A L / m2 BY P A R T I C L E SIZ E 0.45um 11.08 1• 11S!C*■)* mean se c v (55) 75um 51. 18 6 0.07 16 m m Rep 1 Rep 2 Rep 3 12ls i 250um 107.3 108.9 ( g EPILITH0IT/'in2 BY PARTICLE SIZE 1mm .777. .7777 Rep 1 Rep 2 Rep 3 586.6 1721. 701 !5 2619. 176.5 333.5 140.6 139.4 51. 18 62.85 mean se c v (55) 1153. 5 6 7.2 69 .52 1660. 959. 1 81.67 2 5 5.0 7 3 .53 43 .54 140.0 .59 .59 57.01 5.83 14.47 290 mean se c v J5) 16ram APPENDIX D Table D — 1. Summary of physical paramete Augusta Creek, Michigan. D AT E 5. 1 - 11.8 8.0 - 9.9 8.2 8.6 .2 .9 - 1.7 1.9 1.0 1.3 20 M a y 1975 (light! 21 H a y 1975 (dark) 13.0 15.5 - 19.0 19.2 16. 17. O' 01 17 J u l y 1975 ( 1 ight) 18 J u l y 1975 (dark) 12.8 - 19.9 13.6 - 16.3 16. 15. « CO 24 0c t . 1974 (light) 25 0c t . 1974 (dark) TE M P E R A T U R E (C) me a n range 15 Jan. 16 Jan. 1975 ( 1 ight) 1975 ( dark) for selected riffle sections of SMITH SITE, O X Y G E N (mg/liter) range mean • •• o • o • o o o o » « t i t » » » • » • • 9 • • 9 9 ft 9 ftft9 ft9 ftft ftft ft ft ft 9 ft ft 0009 OOOO 0 0 9 9 000 OOOO *009 0*0 f tf tf tf t *999 • •• “ ft ft • 090 "" INCIDENT L I G H T ( lux) range mean 161 - 2 5 8 00 1080 - 2 4 800 10940 12330 181 - 2 3 700 6 4 6 - 19400 7926 9846 42 7970 7 7 5 00 49500 19083 2335S 99 - 4 0 900 2 5 8 0 - 11800 6268 6680 - 291 Table D-2. S u m m a r y of physical parameters A ugusta Creek, Michigan. D AT E T E M P E R A T U R E (C) range mea n 26 Jan. 25 Jan. 1973 (light) 1973 (dark) • •« • .... — .... .... 07 June 06 June 1973 (light) 1973 (dark) _ 15! 6 - .... 07 June 1973 ( 1 ight) .... .... sele c t e d riffle sections of B AVENUE, O X Y G E N (ms/liter) range mea n --- . ... — . . . . - --- 17.3 16.3 - .... .... I NCIDENT L I G H T ( lux) range mean .... 538 — 2 3700 502 — 26900 12320 13560 .. . . — . . . . — --- ---- 147 — 2 1 5 0 0 5 3 8 — 19400 7102 6324 . .. . - .... 147 - 2 1500 7102 61 0 7 7782 • * • • 13. 1 — 18. 1 14.3 — 17.2 15.8 15.5 9 . 1 - 10.3 9.5 - 10.0 9.7 9.9 108 — 4 3100 861 — 3 4400 02 Aug. 01 Aug. 1973 ( light) 1973 (dark) 15.2 - 17.7 15.9 - 19. 1 16.4 17.0 9. 2 - 10.0 9.0 9.8 9.6 9.5 108 _ 64600 1610 — 91500 80 1 7 14607 29 Aug. 30 Aug. 1973 ( 1 ight) 1973 (dark) 15.2 — 15.3 — 19.9 17.2 17.6 16.2 7 . 8 - 10.0 8.3 9.8 9.2 9.5 161 — 38 800 9900 861 — 9445 5490 27 Sep. 26 Sep. 1973 ( light) 1973 (dark) 14.3 - 17.1 14. 1 — 17.7 Id .9 15.8 6.8 6.7 - 8.0 9.2 7.9 7.4 108 — 15100 2488 — 2 3 700 4795 9799 05 Nov. 06 Nov. 1973 ( light) 1973 1 significant, higher more so 2 2 M 2Reg= mean ^of deviation due to regression M Res= mean of deviation of residuals P= level of significance (P=0.05 significant) Table E - 1. Regression parameters for average NCP, CR, GCP a nd N D M versus various independent variables from data collected in Augu s t a Creek, M ic h i g a n riffle sections. Dependent Independent Variable V a r iable N R S.E. M 2 Reg M 2 Res F Ratio P NCP Order X Light X Temperature X AFDW X EAFDW X TAFDW 103 101 102 96 85 85 0.740 0.517 0.328 0.117 0.419 0.410 0.056 0.071 0.079 0.077 0.064 0.065 0.380 0.183 0.075 0.008 0.073 0.070 0.003 0.005 0.006 0.006 0.004 0.004 121.9 36.0 12.0 1.3 17.6 16.8 0.000 0.000 0.001 0.255 0.000 0.000 X CR Order X Temperature X AFDW X EAFDW X TAFDW 104 103 96 85 85 0.519 0.640 0.081 0.306 0.302 0.033 0.030 0.039 0.038 0.384 0.042 0.063 0.001 0.013 0.012 0.001 0.001 0.002 0.001 0.001 37.5 70.0 0.6 8.6 8.3 0.000 0.000 0.435 0.004 0.005 X GCP Order X Light X Temperature X AFDW X EAFDW X T AFDW 101 100 100 94 83 83 0.722 0.565 0.414 0.100 0.393 0.386 0.078 0.093 0.103 0.107 0.094 0.094 0.660 0.401 0.216 0.011 0.130 0.125 0.006 0.009 0.011 0.012 0.009 0.009 108.1 46.0 20.3 0.9 14.8 14.2 0.000 0.000 0.000 0.335 0.002 0.000 X NDM Order X Light X Temperature X AFDW X EAFDW X TAFDW 101 100 100 94 83 83 0.655 0.534 0.339 0.121 0.296 0.286 0.639 0.717 0.800 0.735 0.673 0.675 30.323 20.146 8.123 0.742 3.511 3.274 0.409 0.514 0.639 0.540 0.453 0.456 74.2 39.2 12.7 1.4 7.8 7.2 0.000 0.000 0.001 0.244 0.007 0.009 298 X Table E - 2. Regression parameters for individual rates of NCP and C R versus various independent variables from data collected in A u g u s t a Creek, M i c h i g a n riffle sections. Dependent Independent Variable Variable NCP R S.E. M 2Reg M^Res F Ratio P Light Temperature 666 1957 0.598 0.182 0.086 0.105 8.014 0.744 0.007 0.011 1089.7 67.1 0.000 0.000 T + L 1957 0.598 0.086 4.007 0.007 544.6 0.000 * L 0.597 T 0.005 299 CR N Temperature 1189 0.523 0.035 0.558 0.001 447.1 0.000 T + L 1189 0.567 0.034 0.328 0.001 281.5 0.000 L 0.241 T 0.425 * = multiple regression - temperature + light. Table E - 3. Regression analyses for various sites of Augusta Creek, Mi c h i g a n SMITH Dependent Independent Variable Variable N R S.E. 2 M Reg M 2Res F Ratio P NCP X Light X Temperature X AFDW X EAFDW X TAFDW 12 12 12 12 12 0.270 0.0003 0.131 0.759 0.728 0.008 0.009 0.009 0.006 0.006 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.786 0.000 0.175 13.601 11.244 0.396 0.999 0.684 0.004 0.007 X CR X Temperature X AFDW X EAFDW X TAFDW 12 12 12 12 0.013 0.354 0.381 0.456 0.010 0.009 0.009 0.009 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.002 1.434 1.699 2.618 0.968 0.259 0.222 0.137 X GCP X Light X Temperature X AFDW X EAFDW X T AFDW 12 12 12 12 12 0.255 0.022 0.316 0.405 0.268 0.007 0.007 0.007 0.007 0.007 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.695 0.005 1.112 1.966 0.771 0.424 0.946 0.316 0.191 0.401 X Light X Temperature X AFDW X EAFDW X TAFDW 12 12 12 12 12 0.233 0.133 0.318 0.660 0.696 0.203 0.207 0.198 0.157 .150 0.024 0.008 0.044 0.189 0 . 211 0.041 0.043 0.039 0.025 0.022 0.573 0.180 1.126 7.696 9.393 0.467 0.680 0.314 0.020 0.012 X NDM 300 X Table E - 3. cont. B AVENUE Dependent Independent Variable Variable N R S.E. M 2 Reg M 2 Res F Ratio P X NCP X Light X Temperature X AFDW X EAFDW X TAFDW 33 35 30 24 24 0.160 0.047 0.024 0.206 0.174 0.009 0.008 0.008 0.007 0.007 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.812 0.072 0.016 0.979 0.686 0.374 0.790 0.900 0.333 0.416 X CR X Temperature X AFDW X EAFDW X TAFDW 34 29 23 23 0.785 0.172 0.191 0.271 0.007 0.010 0.010 0.010 0.002 0.000 0.000 0.000 0.000 0.000 0.000 0.000 51.292 0.826 0.796 1.668 0.000 0.372 0.382 0.211 X GCP X Light X Temperature X AFDW X EAFDW X TAFDW 33 34 29 23 23 0.072 0.600 0.117 0.239 0.277 0.014 0.011 0.013 0.014 0.013 0.000 0.002 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.000 0.162 17.861 0.372 1.270 1.739 0.690 0.000 0.547 0.273 0.201 X NDM X Light X Temperature X AFDW X EAFDW X TAFDW 33 34 29 23 23 0.121 0.368 0.181 0.101 0.022 0.149 0.138 0.146 0.112 0.119 0.010 0.096 0.019 0.003 0.000 0.022 0.019 0/021 0.014 0.014 0.457 5.002 0.917 0.215 0.011 0.504 0.032 0.347 0.648 0.919 Table E - 3. cont. U P P E R 43RD Dependent Independent Variable Variable N R S.E. M 2 Reg M 2 Res F Ratio P NCP X Light X Temperature X AFDW X EAFDW X TAFDW 10 10 10 9 9 0.405 0.547 0.281 0.039 0.048 0.070 0.064 0.074 0.076 0.076 0.008 0.014 0.004 0.000 0.000 0.005 0.004 0.005 0.006 0.006 1.569 3.413 0.687 0.007 0.016 0.246 0.102 0.431 0.935 0.902 X CR X Temperature X AFDW X EAFDW X TAFDW 11 11 10 10 0.898 0.194 0.159 0.166 0.018 0.041 0.039 0.038 0.012 0.001 0.000 0.000 0.000 0.002 0.001 0.001 37.649 0.351 0.206 0.226 0.000 0.568 0.662 0.648 X GCP X Light X Temperature X AFDW EAFDW X X TAFDW 10 10 10 9 9 0.484 0.677 0.135 0.004 0.005 0.095 0.080 0.108 0.105 0.105 0.020 0.043 0.002 0.000 0.000 0.009 0.006 0.012 0.011 0.011 2.444 6.768 0.149 0.000 0.000 0.157 0.032 0.710 0.992 0.990 X NDM X Light X Temperature X AFDW EAFDW X X T AFDW 10 10 10 9 9 0.609 0.519 0.340 0.372 0.388 0.711 0.766 0.843 0.861 0.855 2.381 1.728 0.742 0.835 0.905 0.505 0.587 0.710 0.742 0.732 4.713 2.944 1.045 1.126 1.237 0.062 0.125 0.337 0.324 0.303 302 X Table E - 3. cont. NAGEL Dependent Independent Variable Variable N R S.E. M 2 Reg M 2 Res F Ratio P NCP X Light X Temperature X AFDW X EAFDW X TAFDW 35 35 33 29 29 0.241 0.377 0.179 0.083 0.080 0.078 0.075 0.071 0.063 0.063 0.012 0.030 0.005 0.001 0.001 0.006 0.006 0.005 0.004 0.004 2.035 5.453 1.031 0.189 0.175 0.163 0.026 0.318 0.667 0.679 X CR X Temperature X AFDW X EAFDW X TAFDW 34 33 29 29 0.742 0.308 0.198 0.213 0.029 0.042 0.045 0.045 0.034 0.006 0.002 0.003 0.001 0.002 0.002 0.002 39.298 3.243 1.104 1.286 0.000 0.081 0.303 0.267 X GCP X Light X Temperature X AFDW X EAFDW X TAFDW 34 34 32 28 28 0.383 0.529 0.254 0.066 0.074 0.099 0.091 0.098 0.095 0.095 0.054 0.102 0.020 0.001 0.001 0.010 0.008 0.010 0.009 0.009 5.496 12.417 2.062 0.113 0.141 0.025 0.001 0.161 0.740 0.710 X NCP X Light X Temperature X AFDW X EAFDW X TAFDW 34 34 32 28 28 0.263 0.417 0.131 0.181 0.182 0.930 0.876 0.740 0.705 0.705 2.050 5.171 0.287 0.437 0.440 0.864 0.766 0.548 0.496 0.496 2.373 6.747 0.524 0.881 0.887 0.133 0.014 0.475 0.357 0.355 303 X Table E - 3. cont. KELLOGG FOREST Dependent Independent Variable Variable N R S.E. M 2 Reg M 2 Res F Ratio P NCP X Light X Temperature X AFDW X EAFDW X T AFDW 11 10 11 11 11 0.425 0.361 0.271 0.381 0.372 0.030 0.031 0.032 0.031 0.031 0.002 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 1.984 1.200 0.711 1.523 1.447 0.193 0.305 0.421 0.248 0.260 X CR X Temperature X AFDW EAFDW X X TAFDW 12 11 11 11 0.765 0.056 0.168 0.177 0.016 0.024 0.024 0.024 0.003 0.000 0.000 0.000 0.000 0.001 0.001 0.001 14.079 0.029 0.260 0.291 0.004 0.869 0.510 0.603 X GCP X Light X Temperature X AF D W X EAFDW X TAFDW 11 10 11 11 11 0.531 0.660 0.225 0.186 0.175 0.039 0.034 0.052 0.046 0.046 0.005 0.007 0.001 0.001 0.001 0.002 0.001 0.002 0.002 0.002 3.533 6.157 0.478 0.321 0.283 0.093 0.038 0.507 0.585 0.608 X NCP X Light X Temperature X AFDW X EAFDW X TAFDW 11 10 11 11 11 0.637 0.512 0.473 0.564 0.625 0.353 0.398 0.403 0.352 0.357 0.765 0.500 0.422 0.773 0.737 0.124 0.158 0.168 0.124 0.128 6.146 2.838 2.598 6.254 5.769 0.035 0.131 0.141 0.034 0.040 304 X APPENDIX F 305 COMBINED SITES 0.60 1 1 1 1 0.45 1 1 1 11 12 0.30 *1 <§-0.15 O. £ 0.0 1 1 1 1 1 11 1 1 1 22 1 111 1 2 12 11 23 1 121 31221 1221 12 1 1 21 1 311 1 1 1 11 12 1 11 2 311 131 11 21 2 21 2311 111 11 132153441111 1 311 413 2 1141 213 1211 22121 1 234114 131 132411 122211 11122 2 1 111211 1 11 1 24841732 73 222 121211 212112 111 11 112 321 31634351332212 3 1 1 1 1 1 11 1 11 141 3 1 1 11 25144D26242131 3221 1 21 1212 1 24 121 13 1 2 132 11 I 18665764437131 3 3 11 1111 1 3 2 1 111 1 1 111 II 3ANC6CA457B34426163 2 1 2 3113 3 1 12 7flfUSGFFFDC77628S1721 12 2112131 4 111 GMMSSIL7783784417 22 1 1322413 121 1 2 1 0ttSRJ4GSA?615 2 2 2 1 13 11 1 1 411785211 1 1 1 1 1 1 212? 1 11 821 -0.15 1 -0.30 N = 1968 -0.45 .4....4. . . . 4 . . . . 4 . . . . 1 . . . . 4 . . . . 4 . . . . 4 . . . . 4 . . 4 . . . . 1 . . 4 . . . . 4 . . . . 4 . 01 20 m 60 80 100 120 W LIGKT (LUX X 103) Figure F - i. Net community productivity at various light intensities for combined sites of Augusta Creek, Michigan (1973-1975), 306 SMITH 0.60 0.45 0.30 0.15 i < 5* o i 0.0 i 1 i 122311111 1 1 42DJHBE5A4311 43 2 5IGBD7554915 3 4 11 1 1 2 1 12 3 1 2 2 1 -0.15 -0.30 n = 258 -0.45 ♦ .4....4....4....4....4....4....4....4....4....4....4....4....4...,4....4....4. 0 20 40 60 80 100 120 140 LIGHT (LUX X 10^) Figure F - 2. Net community productivity at various light intensities for Smith site, Augusta Creek, Michigan (1975 - 1975). 307 B AVENUE 0.60 0.45 0*30 'I 0.15 1 i l l 0.0 2 1 444 42 214 1 2 3 7IXGC551545434144 6 4ttUTPL0BA47744824 4 JMC74 21 1112 1 2 47 3 1 3 1 31 1 11 2 2 2112 121 1 2 12 11 1 1 -0.15 -0.30 N = 667 -0.45 ♦ ,4....4.,..4....4..m 4.„,4...,4m ..4....4....4....4.. 4....4....4....4....4. 0 20 40 60 80 100 120 140 LIGHT (LUX X 10^) Figure F - 3. Net community productivity at various light intensities for B Avenue site, Augusta Creek, Michigan (1973 - 1975). 308 43rd Up p e r 0.60 i1 0.45 i n i i 1 0.30 1 1112 1 1 1 31 1 1 1 2 12 11 11 1 1 1 1 111 1 1 i 1 2 2 2 1 1 *1 o_ g 0.15 0.0 111222 2 1 1 1 1 122344E4 4 31 1 1 7 312241 1221113 1 3421 1 2212 1 1 1 1 1 1 11 2 1 1 2 i i 1 i i 1 11 11 1 1 1 1 2 1 2 1 21 1 11 11 11 i i 1 i 2 -0.15 1 n = 2 20 -0.30 -0.45 * .4....4....4... . 4 . . . . 4 , . . 4 . . . . 4 . . . . 4 . . . . 4 . . . . 4 , . . . 4 . . 4 . 0 Figure F - 4. 20 40 60 80 100 LIGHT (LUX X 10^) 120 140 Net community productivity at various light intensities for Upper 43rd site, Augusta Creek, Michigan (1974 - 1975), 309 NAGEL 0.60 0.45 i 1 1 1 1 i 1 12 l 1 1 2112 11 1 111 2 1 1 1221 0.30 i < r 1 1 2 1 11 1 2 2 1 1 2 11 11 2 1 1 2 311 21 32142411 11 1 311 311 1 1 4 1 133113 31 2221 11 1 11 2 1 2451 411 43 121 12111 2 21 31534351331212 1 1 1 1 1 1 1 2411382213 121 2221 1 11 2 2 1445123 3 3111 3 2 1 1111 11 356321211261422313 1 1 1 22 1 466S5113733 1114 1 1 11 8A342 1 2 11 1 16243 34321 532 1 1 42 1 i o.i5 C l. 0.0 -0,15 1 11 112 11 1 11 311 1 1 1 2111 1 212 11 1! 11 11 1 1 11 121 11 1 1 1 1 3 1 I II 1 n = 571 -0.30 -0.45 ♦ .4....4....4. 0 20 4....4....4....4....4. .4..,.4,...4....4.,..4....4....4. 40 60 80 100 120 140 LIGHT (LUX X 105) F i g u r e F - 5. Ne t co mmun it y p r o d u c t i v i t y a t v a r i o u s l i g h t i n t e n s i t i e s f o r Nagel s i t e , A ug u sta C re e k , M ich igan (1973 - 1975). 310 KELLOGG FOREST 0,60 0.45 12 0.30 4 1 11 1 1 32 31 4 12 1 . 1 2 1 3112 . 2114234133 . 1923241221 . 2E6212121 4 3423 1 . 14112 1 . 1211 *1 0.15 fe 0.0 1 1 2 1 2 1 1 1 1 1 1 12 2 1 1 1 1 1 1 1 1 1 1 1 11 1 1 2 1 1 1 2 1 1 1 1 1 1 1 11 11 2 3 1 1 1 1 1 11 1 31 11 1 1 11 1 21 1 1 1 1 1 1 241 2 1 1 -0.15 N = 242 -0.30 -0.45 ; .4....4....4....4....4....4....I....4....4....4....4..,.4..,.4....4....4,...4. 0 20 40 60 80 100 120 140 LIGHT (LUX X ICft Figure F - 6. Net community productivity at various light intensities for Kellogg Forest site, Augusta Creek, Michigan (1974 - 1975). 311 COMBINED SITES 0.60 0-5C 0.45 0.30 li 2 *1 U3 21 11 0.15 0.0 3 11 1 11 1 11 2 i i 1 11 1 12112131135 1 116224532361322113 26J9A68B6584 112 4*LLDE5BA52311 1B281821212 2 12 i l 11 1 1 1 1 1 1 1 1 -0.15 -0.30 n = m -0.45 .4....4....4....4....4....1....4....4....4....4...4.,.4....4. Q 20 40 60 light Figure F - 7. 8Q JLQQ 120 M (l u x x 1 0 5) Net community productivity w i thin the 0 - 5 C temperature range for combined sites of Augusta Creek, Michigan (1973 - 1975). COMBINED SITES 0.60 5 -10C 0.45 1 0.30 cS^ CD 0.15 C l. 1 2 11 I ll 1 2 1 1 1 121 11 2 131112 2 1 1 1 1 21 1 222121 111 13 l&U 3 1 1 11 131112 1 1 1 122 1 113 1 1 434422112 122 1 47 42&24C212223 2 1 2BB451 428 512 1 32 1 5=2 0.0 1 1 1 1 1 1 1 1 1 1 1 1 1 2 11 2 2 1 12 1 1 2 2 1 1 1 -0.15 N = 284 -0.30 -0.45 .4....4....4....4....4....4. ,4....4....4. 0 20 40 60 80 100 4....4....4....4....4. 120 140 LIGHT (LUX X 10^) Figure F - 8. Net community productivity within the 5 - 10 C temperature range for combined sites of Augusta Creek, Michigan (1973 - 1975). 313 0.60 COMBINED SITES 10 - 15 C 0.45 1 11 1 1 11121 1 1 1 1 1 1 12 11 0,30 21 1 1 11 1 1 11 2 'I'1 3 111 2 21211 41 1 1 0.15 131 11 3 1 17 3 2 1 2 1 1 12 3 1 1 2 44312 211 11 2 12 5AI9223 234 1 74 2 0.0 2XHAC7541313315342 4 2 1 3PI573232 2 2 2721 1 1 1 11 1 1 1 1 111 1 1 1 -0 .1 5 n = 418 -0.30 -0.45 .*....1....4....4,...4.........4....4. ,.4....4....4....4.M.4.M.4....i....t. 0 20 40 60 80 100 120 140 LIGHT (LUX X ] * Figure p “ 9. Net community productivity within the 10 - 15 C temperature range for combined sites of Augusta Creek, Michigan (1973 - 1975). 314 COMBINED SITES 0.60 15 - 20 C 0,45 1 1 1 11 1 12 12 1 1 I 11 1 0.30 i CD 0.15 0,0 1 1 1 1 11 2 2 311 1 I 1 11 234 31 131321 1 1 1 1 221 1 2 2 41221 1 2 5AA3531 212 !5J?341133 2321 +2K0HFF94112224 2 ISL331 1 1 35112 1 24 1 1 1 1 11 1 1 1 1 11 11 1 1 1 1 13 1 1 1 11 1 1 22 1 2 12 11 1 1 1 1 1 1 1 1 1 1 11 1 1 11 1 1 1 2 1 1 1 1 11 2 11 1 1 1 22 3 1 31 12 2111 1 3 1 12213 1 1 2 1 1 1 1111 1 22 1 1 11 1 121 1 2 1 1 1 11 1 1 -0.15 N = 567 -0.30 -0.45 .i. 0 20 40 60 80 100 120 140 LIGKT (LUX X 10^) Figure F - 10. Net community productivity within the 15 - 20C temperature range for combined sites of Augusta Creek, Michigan (1973 - 1975). 315 0.60 COMBINED SITES 20 - 25 C 11 1 0.45 1 11 I 0.30 1 1 1 11 *1 S1 0.15 4 & 1 112 11 1 t 1 12 231111 15444472 152662111 6632 1644 2 621 1 £s 0 .0 -0.15 1 1 1 1 1 1 11 1 12 1 1 11 21 11 1 1 3 1 12 1 1 11 1 11 1 1 11 1 12 1 1 1 1 1 11 1 12 11 11 111 1 11 1 1 1 11 1 1 1 1 1211 11 111 11 2 1 11 1 1 1 31 1 11 11 2 1 1 1 1 111 1 11 1 1 1 1 (1 i 1 1 1 -0.30 N - 269 -0.45* , i . .. . i . . t , . 0 20 40 . . 1 60 . . . , 4 80 , 100 a, 120 140 LIGHT (LUX X 103) Figure F - 11. Net community productivity within the 20 - 25 C temperature range for combined sites of Augusta Creek, Michigan (1973 - 1975). APP E N D I X G 316 l 0.175 i COMBINED SITES 14 1 1 2 11 2 4 22 11 2 2 2 24 2 11 121 0.M0 1 1111 11 *1 o on 4 22 1 2 2 1 1 11 4232 1 1 112122 22 111241 0.105 31 22 1 21 322312 311 1 1 111 2 3 1 11 1 12 32111222134 2 t i l l 112 1 11 13 11 4 1121 411 111 1 1 1123411 3362 1 11 1 3 3 23223 3542 24211211 2 2211 32 0.070 1 111 4222 11 1 11 1 2 112 31121311 2 2 1 121 22425 113433461A531 3 1 111 1 6 3 1 56123 11 3 1 412 21 2 0.035 4A54U1326S442172 2 1314211356BED833141 140331 2 142 3 213 1 11113 5445AS74312 111 1 11 3 A3? 1 171221 1 31 1 07RBM32343B937 233 1 22 3132794EH865322 111132 1 1 E 436 31 0.000 412ECK54221161 1111 1 11 13217234 113 1 111 1 22 1 1 11 1 1 1 1 -0.035 1 n -1189 i .4....4. 0 5 .4....4....4. 10 .4....4....4., 15 20 ...4. I.4.. 25 30 ..4.1 .4. 35 TEMPERATURE °C Figure G - 1. Rates of community respiration at various temperatures for combined sites of Augusta Creek, Michigan (1973 - 1975). 317 Sm i t h 0.175 0,140 0.105 i sT ai 0.070 1 1 3 32 141 0.035 11 3 3 52 212 13 71 43 44 0.000 22 2 1 1 3 1 2 11541 1 1 1 1 11 13 31 1 2 1 1131 1 1 1 114 2 11 111 11 -0.035 N = 138 i .4....4. 0 5 .4....4....4....4....4....4....4....4....4....4....4....4....4, 10 15 20 25 30 35 TE MPERATURE °C Figure G - 2. Rates of community respiration at various temperatures for Smith site, Augusta Creek, Michigan (1974 - 1975). 318 B AVENUE 0.175 0,140 0.105 *1 Q£ (_> 0.070 u i i 11 l 11 1 5 1 0.035 1 1 1 1 i 1 11231 341722 22112 1 1 11113142111 12 1311 1 457DA42213 2 21 2 1 2 1 1 11113 123357742 1 151221 1 332G323437754 122 1 22 21 2438GS41111 31 1 1K54211151 1111 1 11 1 2232 1 1 0.000 -0.035 1 1 22 1 1 1 1 1 1 N = 426 .4....4. ,.4....4..4.4....4....4....4. .I....*....*. 0 io 15 20 25 30 35 TEMPERATURE ° C Figure G - 3. Rates of community respiration at various temperatures for B Avenue site, Augusta Creek, Michigan (1973 - 1975). 319 UPPER 43rd 0.175 0.140 1 1 1 1 1 1 1 1 0.105 11 21 11 2 111 31 2 21 211111 1 2 1 1 1 1 1 1111 2 11 11 2 111 S' 2 1 1121 1 0.070 11 311 £5 1 1 12 11 0.035 1 1 1 1 i 43 1 33 444 0.000 133 -0.035 n = 125 .4 ....4 ....4 ....4 ....4 ..4 ,...4 .,,,4 ....4 ....4 ....4 ....4 ....4 ....4 . 0 5 10 15 20 25 30 35 TEMPERATURE ° C Figure G - 4. Rates of community respiration at various temperatures for Upper 43rd site, Augusta Creek, Michigan (1974 - 1975), 320 1 1 i 4 • 0,175 4 1 2 * » 2 ♦ • 4 22 11 2 2 24 2 1 12 111 1 4 2 1 2122 1 2122 2 1 24 2 1112 1 1 1 2 221 11 34 1 2 11 1 311 1 2111 2141 2111 323 1 1 21 1 12 11 2 11 11 11 12 13 11 32 121113111 1 I i i 0.M0 • • » • 0.105 i i • a » I 0.070 i i u 112 1 1 11 1 3 3 1111 31 • <1 cS^ NAGEL 1 • • e ♦ . . 0.035 4 . • . . 0.000 4 » f • • 1 11 1113 424 743 1 2 21 12415 1 2 2 12333111 4 3 2 4 31 1 31 1 1 1 2 22 111 3 11 111 2 1 1 2 2 1 1 1 N N 1 = 349 -0.035 • • • • .4....4,...4....4....4. 0 5 10 15 20 2! 30 35 TEMPERATUR E °C Figure G - 5. Rates of community respiration at various temperatures for Nagel site, Augusta Creek, Michigan (1973 - 1975). 321 4....4....4....4....4,,,.4,.,.4..,.4....4....4....4...,4....4.,..4....4....i KELLOGG FOREST 0.175 0.140 1 1 0.105 1 1 3 1 11 1 32 1 22 11 23 11 1 0.070 2 0.035 0,000 1 51 1 3 A3? D 3 5 54 1 1 11 2 11 21 1 1 11 1 32 1 1 1 2 1 2 1 1 11 11 1 1 1 1 1 -0.035 N = 151 ...4....4,...4....4.,.,4....4....4....4.,..4,...4, 0 5 10 15 20 25 .4....4....4....4....4 30 35 TEMPERATURE °C Figure G - 6. Rates of community respiration at various temperatures for Kellogg Forest site, Augusta Creek, Michigan (1973 - 1975). 322 COMBINED SITES 0.60 1 0.4 5 1 111 1 1 11 1 11 1 0.30 i 0.15 Q. O 0.0 1 21 112 1 24 112 1 1 1 1 1 1 1 1 112 22 221 211 2 2 2222 11 311 1 12 111 2213 112 1 1 11 1 41 32 1 2212243211 21 1 2 11213 2 1 231 111 212352 243 113312313 1 2 1 1 2? 1 153 1 511 1 2 111231233 3133 212 113 1 13 22 11 21 3 1111254324741322 22132 2111 2 11 1 3211 1 1 3341 33 3 33226431 3212 2112 2 1 112 2 212353 129 1 22 331134524331131312112 22 11 1 335 1 11 1 11 221112 414635 1 2 1 1 3 122 3AL62 1223 3 2 112114 12 667743688652112211131111 1 60YI821142273314631387881588A97FA417GA574411 32 131 38SLtSF6953C6636443B6KJD13N6CCEI8E?ALtiCA7F311 12113 179A52121 9B23B562346S4 G094CI422Fr?A53A5224 6 211 2 1 21 1 34 1232 1 4 12112 1251 5 2 3 1 11 32 41 323122 11 1 1 3 1 312 1 -0.15 1 -0.30 1 N = 1968 -0.45 1. . . 4 . . , . i . . . . 4 . . . . 4 . . . . 4 . . . . 4 . . . . 4 . 4 . . 4 . . , . 4 , , , . 4 . . . . 4 . M , 4 , . „ 4 . . . . 4 , . . , i . 0 Table G - 7. 5 10 15 20 T EMPERATUR E °C 25 30 7<~ JO Net community productivity at various temperatures for combined sites of Augusta Creek, Michigan (1973 - 1975). 323 Sm i t h 0.60 0.45 0.30 CD 0.15 11 132 459B 4342 0.0 11 121 11212 13 77 2312123342 11 11 231111 735224251344A26B3 T4231 223742422 11 1 1 11 -0.15 n = 258 -0.30 -0.45 .K...!.. 0 . . f . . +... .f.........4....t. 5 10 15 TE MPER ATUR E Figure G - 8. 20 25 30 35 OrC Net community productivity at various temperatures for Smith site of Augusta Creek, Michigan (1974 - 1975), 324 B AVENUE 0,60 0.45 % <§" 0.30 1 0.15 1 4 1 2 12111 5444333422 1 334 13227311343134748 444945E9312K 3 IW5M?53B433332256JGB13D36ACE?975rD 311 5432121 242 &341U241 G9529G4219A51 42 2 1 1 2 1 33 131 1 0.0 -0.15 N = 677 -0.30 -0.45 f ■4....4. 0 Figure G - 9. 5 10 15 20 TEMPERATURE °C 25 30 35 Net community productivity at various temperatures for B Avenue site of Augusta Creek, Michigan (1973 - 1975). 325 43rd upper 0.60 i i i i 0.45. i i 11 11 1 i 1 1 211 1 2 1 111 1 1 1 12 0.30 £ 1 1 0,15 .1 0.0 .232 .3H4 ♦472 .11 2 11 2 3 1 11 -0.15 l l l i l l 11 11 2 111 2 21 1 1 1 112 11 1 11 2 1 1 1 2 1 11 111111 22 2 1 21 11 21111 1 1 2 131 11 1 12 3 1 211 1 2 3 1 1 1 1122 1 1 1 1 1 1 1 1 11 1 1 1 1 1 1 1 n -0.30 = 2 20 -0.45 0 5 10 15 20 25 30' 35 TEMPERATURE ° C Figure G - 10. Net community productivity at various temperatures for Upper 43rd .site of Augusta Creek, Michigan (1974 - 1975). 326 0,60. NAGEL 0.45 1 111 2 1 11 1 1 1 21 112 0,30 1 24 112 1 i 13 0,15 0.0 3 1 11 1 11213 2 1 23 22 1 153 1 51 3 11 22 11 21 3 1 1 3341 33 2 212353 12? 1 1 1 2211 335 1 1 17A3 1223 1 2 1 3 I33C5 11 1 2 1 4 14 32 22 22 1 1 1 1 111 2 211 211 1 2 1112 12 111 11 2 112 1 1 1132111 21 3212112 1 11122 2 3 11 1 31113731121 21 1 11 1 I 22225311 1 2 11 2 12113311 21 131 1 211 2111 412222 2 1213151 5211 2 1 tl 12 21 2 3213 34 22 1 2 I 1351 32 II 11 2 1 1 2 1 1 21 3 1 12 II 1 4 3 -0,15. 111 1 11 2 1 12 211 -0.30 -0.45 N = 571 .4....4....4....4..,.l...,4....l....4..,.4..,.4....4....4....4.,..4..,.4.. 0 5 10 15 TE MPERATURE Figure G - 11. 20 25 30 35 °C Net community productivity at various temperatures for Nagel site of Augusta Creek, Michigan (1973 - 1975). 327 0.60 KELLOGG FOREST 0.45 1 1 1 1 11 11 1 21 1 2 1 111 1 21 1 1 2 11211 1 2 1 1211211 1 2 1 111 1 2 11 :112 111 1 22 1 141311 2 1 1 1 2 1 2 11 12 141 3 1 1 1 11 1 3 3 11 1 13 11 12 1 1 24 1 122 1 31 21 1 1 1 0.30 2 13 111 112 9431 032 CG41 5121 1 1 0.15 0.0 -0.15 -0.30 N = 242 -0.45 .4.,,.4....4.,..4....4....4....4....4.,,.4....4....4. 0 5 10 15 TEMPERATURE Figure G -12. 20 25 .4....4....4. ..4....4. 30 35 °C Net community productivity at various temperatures for Kellogg Forest site of Augusta Creek, Michigan (1974 - 1975). A PP E N D I X H NCP = NET COMMUNITY PRODUCTIVITY GCP = GROSS COMMUNITY PRODUCTIVITY CR = COMMUNITY RESPIRATION NDM = NET DAILY METABOLISM Table H - 1. SITE: Estimates of m e a n hourly rates of community m e t a b o l i s m b a s e d on G i lson Respirometery from five riffle sites of Augusta Creek, Michigan. SMITH DATE: 26 F e b r u a r y 1975 PARTICLE SIZE AFDW G/M-2 DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm jim urn urn 9.06 16.76 28.11 31.77 145.04 - - 27.8000 - 57.1000 - 118.5000 - 70.5000 - 128.3000 28.5000 40.4000 33.0000 27.3000 37.3000 4 1 250 75 mm mm jim jim 1883.83 426.76 95.40 13.67 7.5000 - 19.3000 - 102.4000 - 116.8000 8.7000 1.6000 0.0000 20.2000 COMBINED COMBINED COMBINED COMBINED DETRITUS 4 1 250 75 0.45 mm mm jim jim jim 1892.89 443.51 123.51 45.44 145.04 7.3310 - 20.7283 - 106.0640 - 84.4321 - 128.3000 8.7948 3.0661 7.5108 25.1635 37.3000 DETRITUS EPILITHON TOTAL TOTAL 230.74 2419.66 - 110.030 2.2622 35.2788 7.1697 COMBINED TOTAL 2650.40 - EPILITHON EPILITHON EPILITHON EPILITHON NCP X U102/G-1H-1 11.6443 CR X U102/G-1H-1 9.6169 328 SEDIMENT TYPE Table H - 1 (cont.). SITE: SMITH DATE: 09 June 1975 SEDIMENT TYPE PARTICLE SIZE AFDW G/M-2 X DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm jim jim jim 10.30 38.67 71.71 62.93 136.65 -110.8000 -113.8000 - 68.7000 - 73.2000 -128.6000 92.8000 72.6000 42.1000 46.7000 87.9000 EPILITHON EPILITHON EPILITHON EPILITHON 4 1 250 75 mm mm jim jim 1036.36 321.18 96.35 15.39 - 6.3000 - 11.8000 - 78.8000 - 27.3000 4.2000 19.1000 43.2000 87.6000 COMBINED COMBINED COMBINED COMBINED DETRITUS 4 1 250 75 0.45 mm mm jim jim jim 1046.67 359.85 168.06 78.32 136.65 - 7.3287 - 22.7608 - 74.4905 - 64.1803 -128.6000 5.0722 24.8490 42.7307 54.7372 87.9000 DETRITUS EPILITHON TOTAL TOTAL 320.27 1469.29 -101.943 - 12.4767 67.8598 10.8883 COMBINED TOTAL 1789.55 - 28.4880 21.0842 NCP U102/G_1H'-1 CR x uio2/g" 329 Table H - 1 (cont.). SITE: SMITH DATE: 13 August 1975 PARTICLE SIZE AFDW G/M“2 NCP x u i o 2/g ~ 1h - i CR X U102/G_ DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm pm pm pm 17.20 20.48 20.45 12.56 124.74 -263.1000 -114.6000 - 49.7000 -205.1000 -220.7000 108.4000 88.6000 51.3000 114.9000 156.5000 EPILITHON EPILITHON EPILITHON EPILITHON 4 1 250 75 mm mm um pm 1523.54 297.01 86.81 7.63 - 2.6000 - 25.1000 -189.1000 -321.0000 17.7000 15.8000 147.2000 159.0000 COMBINED COMBINED COMBINED COMBINED DETRITUS 4 1 250 75 0.45 mm mm pm pm pm 1540.74 317.49 107.26 20.19 124.74 - 5.5087 - 30.8730 -162.5260 -248.9050 -220.7000 18.7127 20.4958 128.9184 131.5677 156.5000 DETRITUS EPILITHON TOTAL TOTAL 195.43 1915.00 -194.4210 - 15.8134 131.4702 23.8392 COMBINED TOTAL 2110.43 - 32.3531 33.8062 330 SEDIMENT TYPE Table H - 1 (cont:). SITE: B A VENUE DATE: 24 F ebruary 1975 SEDIMENT TYPE PARTICLE SIZE AFDW G/M-2 DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm urn Aim Aim 10.02 27.03 26.71 30.56 82.45 - 42.9000 - 88.7000 - 85.5000 -108.9000 -160.8000 45.9000 41.7000 26.8000 39.8000 69.4000 EPILITHON EPILITHON EPILITHON EPILITHON 4 1 250 75 mm mm urn pm 1824.18 428.22 258.28 11.29 4.4000 8.8000 3.9000 - 97.4000 4.5000 1.7000 3.7000 4.7000 COMBINED COMBINED COMBINED COMBINED DETRITUS 4 1 250 75 0.45 mm mm pm Aim jum 1834.19 455.25 284.99 41.85 82.45 4.1417 - 13.5440 - 4.4794 -105.798 -160.8000 4.7261 4.0750 5.8651 30.3315 69.4000 DETRITUS EPILITHON TOTAL TOTAL 176.77 2521.96 -122.742 1.6518 52.2778 3.9435 COMBINED TOTAL 2698.73 - NCP x uio2/g~ lV T l 6.4960 7.1094 331 - CR x uio2/g_ Table H - 1 (cont.)* SITE: B AVENUE DATE: 02 June 1975 PARTICLE SIZE AFDW G/M-2 NC? _2 X U l O ^ G -1!! CR X U102/G“ DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm jum jum um 72.15 125.67 94.67 100.23 109.44 - 43.8000 - 75.9000 - 68.2000 - 90.1000 -159.5000 41.2000 56.5000 52.0000 78.2000 153.0000 4 1 250 75 mm mm Aim Aim 2541.46 284.29 125.29 9.20 - 7.6000 - 15.0000 -110.6000 -128.8000 19.5000 21.9000 125.0000 162.9000 COMBINED COMBINED COMBINED COMBINED DETRITUS 4 1 250 75 0.45 mm mm Aim 2613.61 409.97 219.96 109.44 109.44 - 8.5994 - 33.9461 - 92.3515 - 93.3544 -159.5000 20.0991 32.5065 93.5816 85.3227 153.0000 DETRITUS EPILITHON TOTAL TOTAL 502.17 2960.24 - 90.8897 - 13.0853 78.8152 24.6416 COMBINED TOTAL 3462.41 - 24.3697 32.4986 EPILITHON EPILITHON EPILITHON EPILITHON Aim Aim 332 SEDIMENT TYPE Table H - 1 (cont.). SITE: B AVENUE DATE: 11 August 1975 SEDIMENT TYPE PARTICLE SIZE AFDW G/M-2 NCP X U102/G H-1 x CR DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm pm urn pm 26.49 113.27 72.69 64.14 107.17 -108.8000 -108.7000 -245.4000 -191.4000 -319.7000 78.2000 93.7000 206.0000 117.5000 259.6000 EPILITHON EPILITHON EPILITHON EPILITHON 4 1 250 75 mm mm jum pm 2586.29 237.55 149.14 11.77 - 21.7000 52.4000 77.5000 62.7000 34.0000 56.9000 96.1000 144.3000 COMBINED COMBINED COMBINED COMBINED DETRITUS 4 1 250 75 0.45 mm mm pm pm pm 2612.78 350.82 221.84 75.91 107.17 - 22.5830 - 70.5781 -132.5200 -171.4510 -319.7000 34.4481 68.7819 132.1137 121.6542 259.6000 DETRITUS EPILITHON TOTAL TOTAL 383.76 2984.75 -207.346 - 27.0931 164.2083 39.3604 COMBINED TOTAL 3368.51 - 47.6287 53.5838 u i o 2/g " 333 Table H - 1 (cont.) SITE: U P P E R 43RD DATE: 28 F ebruary 1975 PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm jum jum jum 8.55 11.58 34.82 31.23 67.65 - 33.2000 10.2000 - 32.4000 - 52.2000 - 39.7000 78.6000 43.8000 38.3000 35.1000 77.4000 4 1 250 75 mm mm jum jum 4334.02 354.25 67.73 5.72 177.3000 59.4000 169.9000 404.2000 1.9000 4.6000 0.0000 0.0000 COMBINED COMBINED COMBINED COMBINED DETRITUS 4 1 250 75 0.45 mm mm jum jum um 4342.58 365.83 102.56 36.95 67.65 176.8853 57.8433 101.2121 18.4988 - 39.7000 2.0511 5.8403 13.0042 29.6628 77.4000 DETRITUS EPILITHON TOTAL TOTAL 153.83 4761.73 - 36.4687 168.6963 57.5003 2.0716 COMBINED TOTAL 4915.56 162.2759 EPILITHON EPILITHON EPILITHON EPILITHON AFDW G/M“ 2 NCP x u 1 o 2/g - 1h " 1 CR SEDIMENT TYPE x u i o 2/g ~ 3.8061 Table H - 1 SITE: (cont.). UPPER 43RD DATE: 12 June 1975 PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 ram mm um jum pm 30.97 23.12 10.43 16.43 32.53 4 1 250 75 mm ram pm pm 1162.61 27.03 27.03 2.23 COMBINED COMBINED COMBINED COMBINED DETRITUS 4 ram 1 ram 250 p m 75 pm 0.45 jum DETRITUS EPILITHON COMBINED EPILITHON EPILITHON EPILITHON EPILITHON AFDW G/M-2 NCP X UlO^G-J-H-1 CR X U102/G_1H-1 -130.7000 -105.8000 - 28.2000 - 49.4000 1687.6000 127.7000 172.7000 199.5000 257.5000 738.8000 72.8000 72.8000 111.1000 111.1000 205.9000 205.9000 211.0000 211.0000 1193.58 50.15 37.46 18.25 32.53 131.6118 29.4885 265.9547 32.6975 1687.6000 74.2247 139.4977 204.1180 251.8292 738.8000 TOTAL TOTAL 113.08 1218.89 423.6638 144.4912 337.7225 76.8533 TOTAL 1331.98 168.1921 99.0004 335 SEDIMENT TYPE Table H - 1 (cont.). SITE: U P P E R 43RD SEDIMENT TYPE DATE: 15 August 1975 PARTICLE SIZE AFDW G/M-2 4 mm 1 mm 20.00 250 jim 75 um 0.45 jum CR . X U102/G_1H 214.9000 119.1000 139.3000 96.6000 236.8000 mm mm um jim 4676.19 222.92 19.40 7.31 33.1000 20.7000 117.7000 130.1000 12.3000 13.4000 44.1000 119.9000 COMBINED COMBINED COMBINED COMBINED DETRITUS 4 mm 1 mm 250 jim 75 pm 0.45 jim 4682.36 242.92 51.90 78.93 81.31 32.9221 6.2492 - 47.2478 34.4671 -191.9000 12.5669 22.1034 103.7167 98.7591 236.8000 DETRITUS EPILITHON TOTAL TOTAL 211.60 4925.82 -105.366 33.0160 162.6104 12.6348 COMBINED TOTAL 5137.42 27.3164 18.8119 EPILITHON EPILITHON EPILITHON EPILITHON 4 1 250 75 336 32.50 71.61 81.31 -101.9000 -154.8000 -145.7000 24.7000 -191.9000 DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 6.17 NCP X U102/G-1H_1 Table H - 1 (cont.). SITE: NAGEL DATE: 03 M a r c h 1975 SEDIMENT TYPE PARITCLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 I 250 75 0.45 mm mm um um um 27.03 25.15 29.13 30.37 75.46 4 1 250 75 mm mm um um COMBINED COMBINED COMBINED COMBINED DETRITUS 4 1 250 75 0.45 mm mm um um um DETRITUS EPILITHON COMBINED EPILITHON EPILITHON EPILITHON EPILITHON AFDW G/M"2 _ NCP X U10_/G-1H-1 CR X UlO-ZG-1^ 1 6.0000 43.8000 38.0000 49.4000 52.1000 15.1000 19.8000 16.2000 20.5000 28.9000 7098.30 273.67 289.63 3.24 22.4000 11.5000 152.1000 193.4000 1.4000 0.2000 0.0000 0.9000 7125.33 298.82 318.76 33.61 75.46 22.2923 6.8451 134.7285 - 25.9699 - 52.1000 1.4520 1.8498 1.4804 18.6086 28.9000 TOTAL TOTAL 187.14 7664.85 - 41.6931 26.9842 22.3438 1.3040 TOTAL 7851.99 25.3474 1.8055 - Table H - 1 (cont.). SITE: NAGEL DATE: 04 June 1975 CR X U l O ^ G - l H -1 um jjm 35.17 20.64 32.18 10.53 85.89 - 51.9000 - 69.9000 - 85.7000 -124.5000 -159.6000 44.2000 48.5000 57.0000 103.2000 225.1000 4 1 250 75 mm mm jum ju® 4442.46 442.02 167.90 10.53 26.4000 59.3000 153.6000 211.4000 28.9000 32.9000 96.7000 137.8000 COMBINED COMBINED COMBINED COMBINED DETRITUS 4 1 250 75 0.45 mm mm um um um 4477.63 462.66 200.09 20.86 85.89 25.7850 53.5367 1.15.1112 44.9861 -159.6000 29.0202 33.5959 90.3147 120.6582 225.1000 DETRITUS EPILITHON TOTAL TOTAL 184.22 5062.91 -114.1100 33.8754 143.5729 31.7241 COMBINED TOTAL 5247.13 28.6799 35.3349 PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm EPILITHON EPILITHON EPILITHON EPILITHON AFDW G/M-2 338 NCP x u i o 2 /g _1 h -1 SEDIMENT TYPE Table H - 1 (cont.). SITE: NAGEL DATE: 18 August 1975 PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm um um pm 9.86 35.49 28.33 34.63 69.83 ■ 11.4000 ■ 7.1000 - 39.0000 -110.4000 - 40.0000 43.6000 34.0000 52.8000 90.7000 218.8000 4 1 250 75 mm mm um pm 3491.64 181.26 43.25 5.72 58.5000 - 10.7000 14.3000 212.6000 24.9000 39.7000 49.7000 241.9000 COMBINED COMBINED COMBINED COMBINED DETRITUS 4 mm 1 mm 250 um 75 um 0.45 um 3501.50 216.75 71.58 40.35 69.83 58.3032 - 10.1106 - 6.7974 - 64.5844 - 40.0000 24.9527 38.7667 50.9271 112.1468 218.8000 DETRITUS EPILITHON TOTAL TOTAL 178.14 3721.87 45.3895 54.8533 120.9857 26.2427 COMBINED TOTAL 3900.02 50.2744 30.5703 EPILITHON EPILITHON EPILITHON EPILITHON AFDW_ G/M NCP X U102/G“ 1H“ 1 CR X U H ^ / G - I h -1 339 SEDIMENT TYPE Table H - 1 (cont.). SITE: KELLOGG FOREST SEDIMENT TYPE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS DATE: PARTICLE SIZE AFDW G/M 4 mm 8.01 1 mm 21.37 43.50 55.94 116.60 250 jum 75 pm 0.45 Aim • 05 M a r c h 1975 _ NCP X U102/G-lH-l ■ ■ ■ - CR X UlO^G-lH"! 1.3000 33.8000 28.0000 47.1000 47.2000 93.7000 104.2000 41.5000 71.0000 142.7000 2174.17 120.08 31.48 3.18 57.3000 50.5000 111.5000 - 50.0000 22.2000 COMBINED COMBINED COMBINED COMBINED DETRITUS 4 mm 1 mm 250 pm 75 pm 0.45 jum 2182.18 141.45 74.98 59.12 116.60 57.0848 37.7640 30.5687 - 47.2560 - 47.2000 22.4626 44.8604 36.0420 69.0097 142.7000 DETRITUS EPILITHON TOTAL TOTAL 245.42 2328.90 41.1084 57.5356 103.4678 22.9251 COMBINED TOTAL 2574.33 48.1314 30.6037 4 1 25Q 75 34.3000 28.5000 34.0000 340 mm mm pm jum EPILITHON EPILITHON EPILITHON EPILITHON Table H - 1 (cont.). SITE: KELLOGG FOREST DATE: PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm jum *im jum EPILITHON EPILITHON EPILITHON EPILITHON 4 1 250 75 COMBINED COMBINED COMBINED COMBINED DETRITUS 4 1 250 75 0.45 DETRITUS EPILITHON COMBINED AFDW G/M-2 NCP x u i o 2/g -1 h -1 CR x u i o 2/g _ 5.98 34.79 52.57 40.55 61.69 - 81.8000 - 84.8000 - 63.6000 -117.7000 -102.8000 65.0000 78.7000 43.8000 105.8000 358.8000 mm mm jun jim 1884.47 154.55 36.57 34.41 97.0000 33.9000 333.9000 421.2000 41.6000 36.9000 190.6000 214.5000 mm mm jim jim jum 1890.45 189.34 89.14 74.95 61.69 96.4346 12.0898 99.4842 129.6864 -102.8000 41.6740 44.5804 104.0283 155.6996 358.8000 TOTAL TOTAL 195.57 2109.99 - 91.5089 101.7708 162.8758 46.6577 TOTAL 2305.56 85.3758 56.5159 341 SEDIMENT TYPE 06 Ju n e 1975 Table H - 1 (cont.). SITE: KELLOGG FOREST DATE; PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm jum jum Aim EPILITHON EPILITHON EPILITHON EPILITHON 4 1 250 75 mm mm jum jum COMBINED COMBINED COMBINED COMBINED DETRITUS 4 1 250 75 0.45 mm mm Aim Aim Aim DETRITUS EPILITHON TOTAL TOTAL AFDW G/M-2 NCP X U102/G“ 1H“ 1 CR X U102/G-1H-1 -239.0000 - 61.6000 - 96.2000 - 94.5000 -194.8000 489.6000 189.4000 107.5000 98.6000 284.5000 3136.43 146.28 30.21 10.49 - 12.5000 17.5000 7.2000 219.5000 19.1000 20.1000 103.7000 147.1000 3137.39 160.49 67.54 35.96 78.75 - 12.5689 10.4943 - 49.9480 - 2.8658 -194.8000 19.2431 35.0945 105.8002 112.7537 284.5000 156.71 3323.42 -143.2020 - 10.2679 204.7524 20.3172 0.95 14.21 37.33 25.47 78.75 342 SEDIMENT TYPE • 20 A u gust 1975 Table H — 2. SITE: Areal estimates of community m e t a b o l i s m based on Gilson Respirometery experiments from five riffle sites of Aug u s t a Creek, Mic h i g a n . SMITH DATE: 26 F e b r u a r y 1975 NCP GCP CR VR NDM SEDIMENT TYPE PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm jum um jum -0.0032 -0.0121 -0.0423 -0.0284 -0.2363 0.0001 -0.0036 -0.0305 -0.0174 -0.1676 0.0071 0.0187 0.0256 0.0239 0.1492 0.01 -0.19 -1.19 -0.73 -1.12 -0.0070 -0.0222 -0.0561 -0.0413 -0.3168 4 1 250 75 mm mm urn um 0.1794 -0.1046 -0.1240 -0.0203 0.3875 -0.0959 -0.1240 -0.0168 0.4520 0.0188 0.0000 0.0076 0.86 -5.09 0.00 -2.20 -0.0645 -0.1147 -0.1240 -0.0244 4 1 250 75 0.45 mm mm jum pm pm 0.1762 -0.1167 -0.1663 -0.0487 -0.2363 0.3876 -0.0995 -0.1545 -0.0342 -0.1676 0.4591 0.0375 0.0256 0.0315 0.1492 0.84 -2.65 -6.04 -1.08 -1.12 -0.0715 -0.1370 -0.1801 -0.0657 -0.3168 DETRITUS TOTAL EPILITHON TOTAL -0.3223 -0.0695 -0.2190 0.1508 0.2245 0.4784 -0.98 0.32 -0.4435 -0.3276 COMBINED -0.3918 -0.0682 0.7029 -0.10 -0.7711 COMBINED COMBINED COMBINED COMBINED DETRITUS TOTAL g o 2/m "2d _1 G02/M“2D_1 G02 /M-2 d - 343 EPILITHON EPILITHON EPILITHON EPILITHON G02/m "2D-1 SITE: (cont.). SMITH DATE: 09 June .1975 GCP CR g o 2/m “^ d - i VR NDM PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm urn urn jum -0.0199 -0.0769 -0.0860 -0.0805 -0.3069 -0.0032 -0.0278 -0.0333 -0.0291 -0.0971 0.0264 0.0774 0.0833 0.0810 0.3312 -0.12 -0.36 -0.40 -0.36 -0.29 -0I0296 -0*1052 -0.1166 -0.1102 -0.4284 4 1 250 75 mm mm nm jum -0.1140 -0.0662 -0.1326 -0.0073 -0.0380 0.0409 -0.0599 0.0162 .0*1200 0.1692 0.1148 0.0372 -0.32 0.24 -0.52 0.44 -0.1580 -0.1282 -0.1747 -0.0210 4 1 250 75 0.45 mm mm jum um -0.1340 -0.1430 -0.2186 -0.0878 -0.3069 -0.0412 0.0131 -0.0932 -0.0129 -0.0971 0.1464 0.2466 0.1980 0.1182 0.3312 -0.28 0.05 -0.47 -0.11 -0.29 -0.1876 -0.2335 -0.2913 -0.1311 -0.4284 DETRITUS TOTAL EPILITHON TOTAL -0.5702 -0.3202 -0.1906 -0.0408 0.5993 0.4412 -0.32 -0.09 -0.7900 -0.4819 COMBINED -0.8904 -0.2314 1.0405 -0.22 -1.2719 EPILITHON EPILITHON EPILITHON EPILITHON COMBINED COMBINED COMBINED COMBINED DETRITUS TOTAL Jim NCP G02/M-2 d ~1 g o 2 /m " 2d - i SEDIMENT TYPE . g o 2 /m -2d -1 <7»7£ Table H - 2 Table H - 2 SITE: (cont.). SMITH DATE: 13 August NCP G02/M-2D_1 1975 GCP G02/M-2D-! CR -2d - i VR NDM SEDIMENT TYPE PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm pm pm pm -0.0728 -0.0378 -0.0163 -0.0414 -0.4429 -0.0428 -0.0086 0.0005 -0.0182 -0.1288 0.514 0.0500 0.0289 0.0398 0.5383 -0.83 -0.17 0.02 -0.46 -0.24 -0.0942 -0.0586 -0.0284 -0.0580 -0.6672 EPILITHON EPILITHON EPILITHON EPILITHON 4 1 250 75 mm mm pm pm -0.0637 -0.1199 -0.2641 -0.0394 0.3701 -0.0444 -0.0585 -0.0199 0.7436 0.1294 0.3524 0.0335 0.50 -0.34 -0.17 -0.59 -0.3736 -0.1733 -0.4109 -0.0534 COMBINED COMBINED COMBINED COMBINED DETRITUS 4 1 250 75 0.45 mm mm pm pm pm -0.1365 -0.1577 -0.2804 -0.0809 -0.4429 0.3273 -0.0530 -0.0580 -0.0381 -0.1288 0.7951 0.1794 0.3813 0.0733 0.5383 0.41 -0.30 -0.15 -0.52 -0.24 -0.4678 -0.2324 -0.4393 -0.1114 -0.6672 DETRITUS TOTAL EPILITHON TOTAL -0.6112 -0.4871 -0.1979 0.2472 0.7085 1.2589 -0.28 0.20 -0.9064 -1.0117 COMBINED -1.0983 0.0493 0.03 -1.9181 1.9674 g o 2/m ~2d _1 345 TOTAL g o 2 /m Table H - 2 SITE: ( c o n t .) . B AVENUE DATE: 24 PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm jum pm pm -0.0054 -0.0302 -0.0288 -0.0419 -0.1671 0.0004 -0.0160 -0.0198 -0.0266 -0.0950 0.0127 0.0311 0.0197 0.0335 0.1578 0.03 -0.52 -1.00 -0.79 -0.60 -0.0123 -0.0471 -0.0395 -0.0601 -0.2527 4 1 250 75 mm mm pm jum 0.1011 -0.0475 0.0127 -0.0139 0.2046 -0.0383 0.0247 -0.0132 0.2264 0.0201 0.0264 0.0015 0.09 -1.91 0.94 -9.01 -0.0218 -0.0584 -0.0016 -0.0146 4 1 250 75 0.45 mm mm pm pm pm 0.0957 -0.0777 -0.0161 -0.0558 -0.1671 0.2050 -0.0543 0.0050 -0.0398 -0.0950 0.2390 0.0512 0.0461 0.0350 0.1578 0.86 -1.06 0.11 -1.14 -0.60 -0.0341 -0.1055 -0.0411 -0.0748 -0.2527 DETRITUS TOTAL EPILITHON TOTAL -0.2734 0.0525 -0.1570 0.1778 0.2548 0.2743 -0.62 0.65 -0.4118 -0.0964 COMBINED -0.2209 0.0209 0.5291 0.04 -0.5082 COMBINED COMBINED COMBINED COMBINED DETRITUS TOTAL GCP G 0 2/M"2D-i CR GO^M^D"1 P /R ^ NDM G02/M“2D-1 346 SEDIMENT TYPE EPILITHON EPILITHON EPILITHON EPILITHON NCP G02/M-2 d ~ 1 February 1975 TABLE H - 2 SITE: (cont.). B AVENUE DATE: 02 j Une 1975 NDM G02/M“2D~l 0.0820 0.1958 0.1358 0.2161 0.4617 -0.04 -0.22 -0.20 -0.10 -0.03 -0.0852 -0.2381 -0.1623 -0.2368 -0.4741 0.5241 0.0320 0.0313 0.0054 1.3666 0.1717 0.4319 0.0413 0.38 0.19 0.07 0.13 -0.8425 -0.1397 -0.4006 -0.0359 -0.3895 -0.2412 -0.3521 -0.1771 -0.3025 0.5209 -0.0102 0.0047 -0.0152 -0.0123 1.4486 0.3675 0.5676 0.2575 0.4617 0.36 -0.03 0.01 -0.06 -0.03 -0.9277 -0.3777 -0.5629 -0.2727 -0.4741 DETRITUS TOTAL EPILITHON TOTAL -0.7910 -0.6713 -0.1051 0.5929 1.0914 2.0115 -0.10 0.29 -1.1965 -1.4187 COMBINED -1.4623 0.4878 3.1030 0.16 -2.6152 PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm jum pm urn -0.0548 -0.1653 -0.1119 -0.1565 -0.3025 -0.0033 -0.0423 -0.0266 -0.0207 -0.0123 EPILITHON EPILITHON EPILITHON EPILITHON 4 1 250 75 mm mm jum pm -0.3347 -0.0759 -0.2402 -0.0205 COMBINED COMBINED COMBINED COMBINED DETRITUS 4 1 250 75 0.45 mm mm pm jum pm TOTAL NCP ~2i t :l g o 2/m GCP GOj/M^D-1 CR g o 2/m -2d -1 347 VR SEDIMENT TYPE Table H - 2. (cont.). SITE: B AVENUE DATE: 11 August 1975 VR NDM G02/M-2D_1 -0.0131 -0.0275 -0.0463 -0.0767 -0.1042 0.0571 0.2927 0.4130 0.2078 0.7672 -0.23 -0.09 -0.11 -0.37 -0.14 -0.0702 -0.3202 -0.4593 -0.2845 -0.8714 -0.9082 -0.2014 -0.1870 -0.0119 0.5148 0.0173 0.0449 0.0155 2.4249 0.3727 0.3952 0.0468 0.21 -.05 0.11 0.33 -1.9101 -0.355A -0.3503 -0.0313 -0.9548 -0.4007 -0.4757 -0.2106 -0.5544 0.5016 -0.0102 -0.0015 -0.0612 -0.1042 2.4820 0.6654 0.8082 0.2546 0.7672 0.20 -0.02 -0.00 -0.24 -0.14 -1.9803 -0.6756 -0.8096 -0.3158 -0.8714 DETRITUS TOTAL EPILITHON TOTAL -1.2876 -1.3086 -0.2679 0.5925 1.73778 3.2396 -0.15 0.18 -2.0056 -2.6472 COMBINED -2.5962 0.3246 0.07 -4.6528 PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 nun nun urn urn PH1 -0.0466 -0.1992 -0.2887 -0.1987 -0.5544 4 1 250 75 mm mm urn nm 4 1 250 75 0.45 mm mm urn nm um EPILITHON EPILITHON EPILITHON EPILITHON COMBINED COMBINED COMBINED COMBINED DETRITUS TOTAL NCP g o 2/m ~2d ~ 1 GCP g o 2/m ~2d 1 4.9774 348 CR g o 2/m ~ 2d -1 SEDIMENT TYPE Table H - 2 SITE: (cont.)- U P P E R 43RD DATE: 28 F e bruary 1975 NDM PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm um urn jum -0.0036 0.0015 -0.0145 -0.0344 -0.0344 0.0050 0.0080 0.0026 -0.0068 0.0327 0.0185 0.0140 0.0368 0.0302 0.1444 0.27 0.57 0.07 -.23 0.23 -0.0136 -0.0060 -0.0341 -0.0371 -0.1117 4 1 250 75 mm mm um jum 9.8445 0.2696 0.1474 0.0296 9.9500 0.2905 0.1474 0.0296 0.2271 0.0449 0.0000 0.000 43.82 6.46 0.00 0.00 9.7229 0.2455 0.1474 0.0296 4 1 250 75 0.45 mm mm jum jum jum 9.8409 0.2711 0.1330 0.0088 -0.0344 9.9550 0.2985 0.1501 0.0228 0.0327 0.2456 0.0589 0.0368 0.0302 0.1444 40.53 5.07 4.08 0.75 0.23 9.7094 0.2396 0.1133 -0.0074 -0.1117 DETRITUS TOTAL EPILITHON TOTAL -0.0719 10.2912 0.414 10.4176 0.2439 0.2720 0.17 38.30 -0.2025 10.1455 COMBINED 10.2193 10.4590 0.5159 20.27 9.9431 COMBINED COMBINED COMBINED COMBINED DETRITUS TOTAL GCP G02/M"2D-1 g o 2/m ^D 1 349 EPILITHON EPILITHON EPILITHON EPILITHON NCP g o 2/m - 2d -1 CR G02/M-2D-1 VR SEDIMENT TYPE Table H - 2, ( cont.)* SITE: UPPER 4 3RD DATE: 12 June 1975 VR SEDIMENT TYPE PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm jum jum jum -0.0709 -0.0428 0.0051 -0.0139 0.9609 -0.0016 0.0271 0.0416 0.0584 1.3816 0.1091 0.1101 0.0574 0.1138 0.6628 -0.01 0.25 0.72 0.51 2.08 -0.1107 -0.0830 -0.0158 -0.0554 0.7188 EPILITHON EPILITHON EPILITHON EPILITHON 4 1 250 75 mm mm /am /am 2.8204 0.0687 0.1692 0.0243 4.3018 0.1213 0.2666 0.0325 2.3340 0.0828 0.1535 0.0130 1.84 1.46 1.74 2.51 1.9679 ■ 0.0384 0.1132 0.0196 COMBINED COMBINED COMBINED COMBINED DETRITUS 4 1 250 75 0.45 mm mm jum jum pm 2.7495 0.0259 0.1744 0.0104 0.9606 4.3002 0.1483 0.3082 0.0909 1.3816 2.4430 0.1929 0.2109 0.1268 0.6628 1.76 0.77 1.46 0.72 2.08 1.8572 -0.0446 0.0974 -0.0359 0.7188 DETRITUS TOTAL EPILITHON TOTAL 0.8385 3.0826 1.5070 4.7223 1.0531 2.5832 1.43 1.83 0.4539 2.1390 COMBINED 3.9212 6.2292 3.6363 1.71 2.5929 GCP g o 2/m "2d -1 CR G02/M“ 2D-1 NDM g o 2/m -2 d -1 350 TOTAL NCP G02/M"2D-! Table H - 2 SITE: (c o n t .). U P P E R 43RD DATE: 15 August 1975 VR g o 2/m “ z d ~ 0.366 0.0657 0.1248 0.1908 0.5310 0.30 -0.17 -0.03 0.73 0.11 -0.0254 -0.0771 -0.1282 -0.0519 -0.4726 3.3947 0.1215 0.0502 0.0292 1.5861 0.0824 0.0236 0.0242 2.14 1.48 2.13 1.21 1.8086 0.0392 0.0266 0.0051 2.4649 0.0243 -0.0392 0.0435 -0.2495 3.4059 0.1101 0.0469 0.1681 0.0584 1.6227 0.1481 0.1484 0.2149 0.5310 2.10 0.74 0.32 0.78 0.11 1.7832 -0.0379 -0.1016 -0.0468 -0.4726 -0.3565 2.6005 0.1937 3.5957 0.9488 1.7162 0.20 2.10 -0.7551 1.8795 2.2440 3.7894 2.6651 1.42 1.1243 SEDIMENT TYPE PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 tnm mm jum jum jum -0.0101 -0.0495 -0.0757 0.0283 -0.2495 0.0111 -0.0114 -0.0033 0.1389 0.0584 EPILITHON EPILITHON EPILITHON EPILITHON 4 1 250 75 mm mm /urn jum 2.4750 0.0738 0.0365 0.0152 COMBINED COMBINED COMBINED COMBINED DETRITUS 4 1 250 75 0.45 mm min pm pm pm DETRITUS TOTAL EPILITHON TOTAL COMBINED TOTAL NCP G 0 2/M_2D - 1 GCP G02/M"2D_1 CR g o 2/m -1 d - 1 NDM . Table H - 2, SITE: (cont.). NAGEL DATE: CR G02/M-2 d ~1 VR 0.0032 -0.0079 -0.0083 -0.0114 -0.0228 0.0113 0.0137 0.0130 0.0172 0.0601 0.28 -0.57 -0.64 -0.67 -0.38 -0.0081 — .0216 -0.0213 -0.0286 -0.0829 2.0705 0.0410 0.5737 0.0082 2.1999 0.0417 0.5737 0.0082 0.2740 0.0015 0.0000 0.0001 8.03 27.62 0.00 101.95 1.9259 0.0402 0.5737 0.0081 2.0684 0.0266 0.5592 -0.0114 2.2031 0.0338 0.5654 -0.0032 0.2853 0.0152 0.0130 0.0172 7.72 2.22 43.45 -0.19 1.9178 0.0186 0.5524 -0.0206 -0.1016 2.6933 -0.0472 2.8235 0.1153 0.2756 -0.41 10.24 -0.1625 2.5479 2.5917 2.7763 0.3909 7.10 2.3854 PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm jum jum jum -0.0021 -0.0143 -0.0144 -0.0195 -0.0512 EPILITHON EPILITHON EPILITHON EPILITHON 4 1 250 75 mm mm jum jum COMBINED COMBINED COMBINED COMBINED 4 1 250 75 mm mm mm jum COMBINED TOTAL NCP _2 d - 1 g o 2/m GCP G02/M“2D-1 NDM g o 2/m -2 d _ - 352 SEDIMENT TYPE DETRITUS TOTAL EPILITHON TOTAL 03 M a r c h . 1975 Table H - 2 SITE: (cont.) NAGEL DATE: SEDIMENT TYPE PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 COMBINED COMBINED COMBINED COMBINED DETRITUS 1975 GCP g o 2/m -2 d ~ 1 g o 2/m -2 d - 1 VR CR NDM g o 2/m D_1 mm mm jum pm pm -0.0317 -0.0251 -0.0480 -0.0224 -0.2384 -0.0047 -0.0077 -0.0161 -0.0038 0.0978 0.0429 0.0276 0.0506 0.0294 0.5332 -0.11 -0.28 -0.32 -0.13 0.18 -0.0476 -0.0353 -0.0666 -0.0332 -0.4353 4 mm mm 250 pm 75 um 2.0393 0.4558 Q.4484 0.0387 4.2717 0.7086 0.7308 0.0639 3.5404 0.4010 0.4477 0.0400 1.21 1.77 1.63 1.60 0.7313 0.3076 0.2830 0.0239 2.0076 0.4307 0.4005 0.0163 -0.2384 4.2670 0.7010 0.7147 0.0601 0.0978 3.5833 0.4286 0.4983 0.0694 0.5332 1.19 1.64 1.43 0.87 0.18 0.6837 0.2723 0.2164 -0.0093 -0.4353 -0.3655 2.9822 0.0655 5.7750 0.6836 4.4292 0.10 1.30 -0.6181 1.3459 2.6167 5.8406 5.1128 1.14 0.7278 1 4 1 250 75 0.45 mm mm pm pm pm DETRITUS TOTAL EPILITHON TOTAL COMBINED NCP "2 d _1 g o 2/m June TOTAL 353 EPILITHON EPILITHON EPILITHON EPILITHON 04 Table H - 2. SITE: (cont.) NAGEL DATE: g o 2/m -2 d - 1 V* 0.0050 0.0151 0.0062 -0.0108 0.1975 0.0119 0.0333 0.0413 0.0866 0.4213 0.42 0.45 0.15 -0.12 0.47 -0.0068 -0.0182 -0.0351 -0.0974 -0.2238 3.2310 -0.0307 0.0098 0.0192 4.6062 0.0831 0.0438 0.0412 2.3975 0.1984 0.0593 0.0382 1.92 0.42 0.74 1.08 2.2087 -0.1153 -0.0155 0.0030 3.2292 -0.0347 -0.0077 -0.0412 -0.0442 4.6112 0.0982 0.0500 0.0304 0.1975 2.4094 0.2317 0.1005 0.1248 0.4213 1.91 0.42 0.50 0.24 0.47 2.2019 -0.1335 -0.0506 -0.0944 -0,2238 0.1279 3.2293 0.2130 4.7743 0.5943 2.6934 0.36 1.77 -0.3813 2.0809 4.9873 3.2877 1.52 1.6996 SEDIMENT TYPE PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm >im jum um -0.0018 -0.0040 -0.0175 -0.0605 -0.0442 4 1 250 75 mm mm jam pm 4 1 250 75 0.45 mm mm Jim pm pm EPILITHON EPILITHON EPILITHON EPILITHON COMBINED COMBINED COMBINED COMBINED DETRITUS DETRITUS TOTAL EPILITHON TOTAL COMBINED TOTAL 18 A u g u s t . 1975 NCP “2d ~ 1 g o 2/m 3.1014 GCP G02/M-2 d ~1 CR NDM g o 2/m _2 d - j Table H - 2. SITE: (cont.). KELLOGG FOREST SEDIMENT TYPE PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 COMBINED COMBINED COMBINED COMBINED DETRITUS NCP "2d -1 g o 2/m GCP g o 2/m _2 d _1 CR g o 2/m ~2 d ~ 1 VR NDM g o 2/m -2 d - 1 -0.0085 -0.0514 -0.0583 -0.0832 -0.1105 -0.0018 -0.0037 -0.0181 -0.0084 0.2752 0.0107 0.0755 0.0635 0.1183 0.6104 -0.16 -0.05 -0.29 -0.07 0.45 -0.0125 -0.0792 -0.0816 -0.1267 -0.3352 4 1 250 75 mm mm pm pm 3.1854 0.0913 0.2128 0.2526 4.5516 0.1907 0.3343 0.3812 2.1618 0.1573 0.1922 0.2035 2.11 1.21 1.74 1.87 2.3898 0.0334 0.1420 0.1776 4 1 250 75 0.45 mm mm pm pm pm 3.1769 0.0399 0.1545 0.1694 -0.1105 4.598 0.1870 0.3161 0.3728 0.2752 2.1725 0.2328 0.2557 0.3218 0.6104 2.09 0.80 1.24 1.16 0.45 2.3773 -0.0458 0.0604 0.0509 -0.3352 -0.3119 3.7421 0.2432 5.4577 0.8784 2.7148 0.28 12.01 -0.6352 2.7429 3.4302 5.7009 3.5932 1.59 DETRITUS TOTAL EPILITHON TOTAL COMBINED 06 M a r c h 1975 TOTAL 2.1077 355 EPILITHON EPILITHON EPILITHON EPILITHON mm mm pm jUffl pm DATE: Table H - 2 SITE: ( c o n t .). KELLOGG FOREST DATE: 05 June 1975 VR SEDIMENT TYPE PARTICLE SIZE DETRITUS DETRITUS DETRITUS DETRITUS DETRITUS 4 1 250 75 0.45 mm mm Aim urn Aim -0.0001 -0.0095 -0.0160 -0.0346 -0.0722 0.0097 0.0197 0.0077 0.0175 0.1461 0.0207 0.0614 0.0498 0.1095 0.4588 0.47 0.32 0.15 0.16 0.32 -0.0110 -0.0417 -0.0421 -0.0920 -0.3128 4 1 250 75 mm mm pm jum 1.6342 0.0795 0.0460 -0.0021 2.2674 0.1336 0.0578 -0.0007 1.3310 0.1136 0.0247 0.0030 1.70 1.18 2.34 -0.22 0.9364 0.0200 0.0331 -0.0036 4 1 250 75 0.45 mm mm jum Aim Aim 1.6341 0.0701 0.0301 -0.0366 -0.0722 2.^771 0.1533 0.0655 0.0169 0.1461 1.3517 0.1750 0.0745 0.1125 0.4588 1.68 0.88 0.88 0.15 0.32 0.9254 -0.0217 -0.0090 -0.0956 -0.3128 -0.1323 1.7577 0.2008 2.4581 0.7002 1.4723 0.29 1.67 -0.4995 0.9858 1.6254 2.6588 2.1725 1.22 0.4863 COMBINED COMBINED COMBINED COMBINED DETRITUS DETRITUS TOTAL EPILITHON TOTAL COMBINED TOTAL GCP . G O ^ M ^ D -1 CR g o 2/m - 2d _1 NDM g o 2/m _2 d ~ 1 356 EPILITHON EPILITHON EPILITHON EPILITHON NCP g o 2/m -2d -1