71-18,269 PARKER, George Ralph, 1942THE STRUCTURE OF A SWAMP COMMUNITY IN NORTHERN MICHIGAN AND ITS REACTION TO PARTIAL DRAINAGE. Michigan State University, Ph.D., 1970 Ecology U n iv ersity M icrofilm s, A XEROX C om p an y , A n n A rbor, M ich ig an THE STRUCTURE OF A SWAMP COMMUNITY IN NORTHERN MICHIGAN AND ITS REACTION TO PARTIAL DRAINAGE By George Ralph Parker A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Forestry 1970 ABSTRACT THE STRUCTURE OF A SWAMP COMMUNITY IN NORTHERN MICHIGAN AND ITS REACTION TO PARTIAL DRAINAGE By George Ralph Parker The study was conducted in a swamp community on the eastern edge of Michigan's upper peninsula. It was designed to (1) characterize the present structure of* the primary producers and significant edaphlc factors in a swamp community, and (2) to determine what effects a 6-year-old drainage ditch has had on the site. Quadrats and water wells were located at three distances and on two sites along an existing drainage ditch. Herbaceous and woody shrub species were clipped for biomass, density, and frequency determination. and shrub species were measured for basal area, and density estimates. Tree frequency, Sample stems of Alnus rugosa and Fraxinus nigra were clipped for biomass and production estimates. Foliar samples from dominant species were collected for nutrient analysis. Basal discs of Alnus were used for growth ring analysis. Impermeable soils, physiographic location, and periods of excessive precipitation result in a high water table through much of the growing season. Boils George R. Parker consisted of a shallow layer of black, mucky organic matter underlain by silty clay loam on one site and very fine sand on the other s i t e . The community is enriched by nutrients carried by runoff water from surrounding uplands. This is indicated by nearly neutral pH of the ground water and high X'oliar nitrogen percentages in the five dominant plant species. These edaphic characteristics result in a rich and characteristic flora. on the study area. More than 100 species were identified Alnus rugosa and Fraxinus nigra were dominant overstory species, Cornus stolonifera and Ilex verticillata were dominant shrubs of intermediate height* and the understory was predominated by Glycerla striata and Xmpatiens capensls. Alnus has maintained its dominance on the better drained of the two study sites* while the more poorly drained site is in a transitional stage of succession leading to a swamp-hardwood community. These changes in overstory species have resulted in changes in the under­ story species. The structure of the community has not been Influenced by either fire or logging for the past 50 years. The above-ground dry weight net primary production for the predominant species was estimated to be 5^5 and 572 g/m2/year for Sites 1 and 2* respectively. ground dry weight biomass Above­ (g/m2 ) of these predominant George R. Parker species was 3,235 for Site 1 and 2,451 for Site 2. Predominant species include all herbs, small shrubs,, and the two dominant overstory species. The effect of the drainage ditch on edaphic factors and the structure of the plant community was minor. Differential drawdown of the water table did not vary significantly between 7 and 91 meters from the drainage ditch. However, there was evidence that rate of discharge of surface water is greater near the ditch. decrease in depth of organic matter, ground water, Significant increased pH of increased biomass of herbaceous species, as well as a slight increase in foliar nitrogen for four of five dominant species tested indicate greater rates of decomposition and mineralization are occurring within 40 m of the drainage ditch than farther away. This is partly responsible for a significant increase in number of individuals for the herbaceous species Impatlens capensis. However, there has not been a major change in species composition or a significant shift in biomass among the species. increased, Diversity per individual significantly and redundancy significantly decreased with distance from the drainage ditch based on number of individuals and measured with the factorial expression of the Shannon-Weaver function. However, when biomass per species was used with this function, diversity per individual was not significantly different with distance from the drainage ditch. George R. Parker A growth ring analysis of Alnus indicates that insufficient time has elapsed since construction of the drainage ditch_, and/or its effect has not b ee n great enough to cause a growth response. ACKNOWLEDGMENTS I would like to express my appreciation to Dr. G. Schneider under whose guidance and encouragement this study was conducted, and to the other members of the guidance committee, Dr. W. E. Cooper, Dr. V. J. Rudolph, and Dr. S. N. Stephenson, and reviews of the manuscript. for their suggestions Appreciation also goes to Dr. J. E. Cantlon and Dr. D. P. White for their suggestions in initiating this research, Dr. Edward Voss, Michigan, Professor of Taxonomy, and to University of for his help in identification of the more difficult plant species. Special thanks is extended to Maurice and Vivian Day for making m y stay at the Dunbar Forest Experiment Station a very pleasant and memorable experience. Mr. Day's knowledge of the study area and help in establishment of this research project were invaluable. Finally, m y greatest thanks and appreciation goes to my wife, Mary Lee. Her encouragement and assistance were essential in completing this viork. ii VITA George Ralph Parker Candidate Tor the Degree of DOCTOR O F PHILOSOPHY Final Examination: Guidance Dr. Dr. Dr. Dr. September 4, 1970 Committee: G. Schneider (Chairman), Department of Forestry W. E. Cooper, Department of Zoology V. J. Rudolph, Department of Forestry S. N. Stephenson, Department of Botany Dissertation: The Structure of a Swamp Community in Northern Michigan and Its Reaction to Partial Drainage Biographical Items: Born September 16, 1942, Tulsa, Oklahoma. Married Mary Lee Williams, 1965* Education: Diploma from Charles Page High School, Sand Springs, Oklahoma, i960 B.S. degree in Forestry from Oklahoma State University, 1964 M.S. degree in Botany from Oklahoma State University, 1967 Ph.D. degree in Forestry from Michigan State University, 1970 Professional Experience: 1961-1964 Summer employment with Forest Service, U. S. Department of Agriculture, In Idaho and Washington while earning B.S. degree. 1965-1967 Graduate Teaching Assistant in dendrology and general botany, Oklahoma State University, while earning M.S. degree. 1967-1970 Graduate Research Assistant, Michigan State University, while earning Ph.D. degree. Professional Organizations*. Society of American Foresters Phi Sigma Society Society of Sigma Xi Xi Sigma PI Ecological Society of America American Association for the Advancement of Science American Museum of .Natural History iii TABLE OF CONTENTS Chapter I. II. III. IV. Page I N T R O D U C T I O N ........... 1 DESCRIPTION O F A R E A ................... 3 METHODS 6 ....................... Abiotic Sampling Procedure ............ Biotic Sampling Procedure . 11 Analysis of D a t a ....................... 16- RESULTS AND D I S C U S S I O N ................. 22 Present State of the S y s t e m ......... 22 Abiotic Factors . . . . . . . . . . 22 .................. 27 Foliar Nutrients The Flora ..................... 29 Species Importance ................ 30 Successional Change . . . . . . . . 36 Biomass and Production ........... 43 Effects of D r a i n a g e .................. 33 Abiotic Factors Biotic Factors V. 6 ........... 53 ..................... 6l S U M M A R Y ................................. 72 BIBLIOGRAPHY .................................... 77 A P P E N D I X ......................................... 83 iv LIST O F TABLES ace Analysis of Variance Tables for Factors Measured on Two Sites and Three Distances From Drainage Ditch, 1 9 6 9 ............*... .......................... 26 Foliar Nutrient Values for the Dominant Species Found on Each Study Site ......................... 28 Importance Values for Herbaceous Species at Three Distances From Drainage Ditch for Both Study S i t e s ......... .. ................................... 31 Percent Frequency of Quadrat Occurrence for Low Shrub and Vine Species on Each Site at Three Distances From Drainage Ditch ..................... 34 Percent Frequency of Quadrat Occurrence for Trees and Shrubs on Two Sites and Three Distances From Drainage Ditch, 19^9.. .............................. 35 Number of Clumps and Stems for Trees and Shrubs on Two Sites and Three Distances From Drainage Ditch, 1 9 6 9 ........................ * ............... 37 Number of Seedlings and Basal Sprouts of Tree Species at Three Distances From Drainage Ditch for Both Study Sites .............................. 39 Importance Values for Tree Species at Three Distances From Drainage Ditch for Bo t h Study Sites ................................................. 39 Basal Area for Tree Species on Two Sites at Three Distances From Drainage Ditch .............. 40 Means and Standard Deviations of Biotic Factors for Site 1 and Site 2, 1969 ....................... 42 Regressions of Sample Stem Components of Alnus rugosa and Fraxinus nigra on Sites 1 and 2 ^ . . 46 Dry Weight Biomass, Net Production, and Relative Distribution by Component Part for Alnus rugosa on Two Sites and Three Distances From Drainage ' Ditch, August, 1 9 6 9 ......... * .................... v 48 Page Table 13. Dry Weight Biomass, Net Production, and Relative Distribution by Component Part for Fraxinus nigra on Two Sites and Three Distances From “ D rainage Ditch, August, 1 9 6 9 ...................... 49 14. Dry Weight Biomass, Net Production, and Percent Distribution by Species Category for Each Site . 52 15- Differences in Abiotic Site Factors and Standard Deviations With Distance From Drainage Ditch, June-July, 1 9 6 9 .................................... 57 16. Rates of Recharge and Discharge From Wells on Site 1 Determined From Continuous Water Level Recorder C h a r t s .................... .............. 59 17. Mean Monthly Precipitation and Deviation From 20-Year Norm at the Dunbar Forest Experiment Station . 60 18. Relative Above-Ground Dry Weight and Relative Density for 12 Most Important Herbaceous Species With Distance From D itch on Each S i t e ............ 63 19. Means and Standard Deviations of Biotic Factors for Both Sites With Distance From Drainage Ditch, 1 9 6 9 .................. 64 20. Percent Dry Weight Foliar Nitrogen for Five Dominant Species on Two Sites and Three Distances From Drainage D i t c h ............... .............. 69 21. Mean Growth Ring Index Values of Sites 1 and 2 for Alnus rugosa at Three Distances From Drainage D i t c h .................. 71 22. Relative Elevation of Pipe Located at Each W e 11 and Along Center of D i t c h .........................83 23. Relative Elevation of Soil Surface at Each Well and Along Center of Drainage Ditch ................ 24. Average Depth of Organic Matter on Each Quadrat and Average Depth to Mineral Soil Textural Change at Each Well L o c a t i o n ....................... 84 25. PH in the Upper 15 cm of Ground Water Measured at Each Well and Drainage Ditch, June-July, 1969 85 26. Oxygen (mg/1) in Upper 15 cm of Ground Water in Each-Well and Drainage Ditch, July, 1969 • * 85 vi 83 Page Table 27. 28. Weekly Fluctuation in the Water Table and Precipitation for Each Site During the 1 969 Growing Season ..................................... 86 Current Growth Dry Weight, Number of Stems, and Percent of Quadrat Occurrence of Species on Two Sites and Three Distances From Drainage Ditch . 88 2 9 . I/ist of Species Seen on Study Sites But Not Found on Sample Q u a d r a t s ......................... 30. Dry Weight, Density and Number of Herbaceous Species Found on Each 1 Q u a d r a t ............... 98 99 31. Species Diversity Per Individual and Redundancy for Numbers of Individuals of Herbaceous Species Per 1 Q u a d r a t ..................................... 100 32. Species Diversity Per Individual and Redundancy . for Biomass of Herbaceous Species Found on Each 1 Q u a d r a t ......................... 101 33. Current Dry Weight, Density and Number of Woody Species Found on Each 1 Q u a d r a t ..................102 34. Basal Area of All Tree Species by Quadrat on Two Sites and Three Distances From Drainage Ditch, 1969 ................................................ 35- 103 Stem and Branch Data for Sample Stems of Alnus rugosa by Site and Distance From Drainage Ditch, T $ 6 9 ' ~ ............................................. 104 36- Stem and Branch Data for Sample Stems of Fraxinus nigra by Site and Distance From Drainage Ditch, 106 37. Dry Weight and Percent Distribution of Bark and Wood for Stem Discs of Alnus r u g o s a .............. 108 38. Dry Weight and Percent Distribution of Bark and Wood for Stem Discs of Fraxinus n i g r a ............ 109 39. Macro- and Micro-Nutrient Values for Foliage of Three Dominant Woody Species on Two Sites and Two Distances From Drainage Ditch, July, 1968 . vii 110 LIST 01*' FIGURES Figure 1. 2. 3. 4. Page Aerial View of Study Area Showing Drainage Ditch and Location of Study Sites (May, 1 9 ^ 9 ) ......... k Early Spring View of Drainage Ditch Adjacent to Site 1. Note Species Mixture of Fraxinus nigra and Alnus rugosa on Site to Right of Ditch ... 7 Early Spring View of Drainage Ditch Adjacent to Site 2. Note Predominance of Alnus rugosa on Site to Left of D i t c h ............................. 7 Distribution of Quadrats, Wells, and Precipitation Gauges in Relation to Drainage Ditch for Sites 1 and 2 ............................................... 8 5. Precipitation Gauge Located on Cleared Area Near Site 2 ................................................10 6. A I M 2 Quadrat on Line 2 of Site 1 Before Clipping. TheDominant Grass Species is Glyceria striata; Woody Stems are Alnus rugosa . 12 7. A 1 M Quadrat on Line 3 of Site 1 After Clipping of All Above-Ground Herbaceous and Woody Vegeta­ tion (<1 cm Basal D i a m e t e r ) ......................... 12 8. Field Determination of Stem Green VJeights for Alnus rugosa and Fraxinus n i g r a ..................... 15 9- Early Spring View of Line 2, Site 1 Showing Variable Ground Surface. Larger Stems Near Center are Fraxinus nigra; Stems in Right Foreground are Alnus "rugosa; Log Is Betula papyrifera ... 23 10. Early Spring View of Line 2, Site 2 Showing Ground Surface. Stems are Predominantly Alnus rugosa; Dead Snag in Center Background is Populus balsamif e r a ........................................... 23 11. Mean Relative Elevation of Ground Surface and Water Table and Amount of Precipitation Per Week for Both Sites, 1969 •* ............................. 2k viii Figure 12, Page Week l y Relative Water Level W ith Distance From Drainage Ditch and Amount of P r e ci pi ­ tation During 19^9 Growing Season . . . . . . ix 55 CHAPTER I. INTRODUCTION Approximately 100 million acres in bhe eastern United States are classified as poorly drained Wright, 1907). (Schwab, et al., 1966; Much of this area exists in the northern regions where extensive basins possessing poor natural drainage were formed during the last glacial stage. On such lands the high water table is often the dominant factor controlling the structure of the plant community. If these ecosystems are modified through partial drainage,, they should serve as good sites to study the dynamics of the system along the resulting water table gradient. Over the past 100 years, European scientists have gained much insight into the dynamics of poorly drained forest lands when modified by drainage. This was primarily done to increase plant productivity (Holmen, 19*54; Huikari, 1 9 6 5 ; Pyavchenko, 1957)* A review of Forestry Abstracts back to 1955 indicates that while little research was done by American scientists in this area, a great deal of research was conducted during that period by Russian scientists. However, their work, was often not readily available to American Investigaters. This lack of research by American scientists thus far is probably due to the abundance of available productive forest lands. As pressures for more and more forest lands increase* more intensive management of the better forest areas and/or an extension of our management practices into the more marginal areas will be needed. Basic research on lowland areas has been done (Bay* 1 9 6 6 ; Damman* 1964; (bogs and swamps) Heinselman* 1 9 6 3 ; Pierce* 1957; Moss* 1953; V/hite* 1 9 6 5 ; Watt and Heinselman* 1 9 6 5 ; Sjors* 1959; Ritchie* Vianna* 1952; Conway* i9 6 0 ; Dansereau and Segadas- 19^9; and Gates* 1942). More information is now needed in determining the best possible use and management for these lands. Their value for uses such as water recharge areas* wildlife habitat* and genetic pools must not be sacrificed for increased production of wood products to obtain a short-term economic gain (Stoeckeler* 1962; Klawitter* 197^)* To avoid this* research needs to be done to determine their reaction to man's activities. This study is the first part of a long-term research project at Michigan State University* which is investigating the dynamics of a poorly drained ecosystem when subjected to man's management practices. The objectives of this study were to (1) characterize the present state of the system* and (2) to determine what effects an existing drainage ditch has on the structure of the plant community. CHAPTER II. DESCRIPTION OP AREA The study was conducted during 1968 and 1969 on the Dunbar Forest Experiment Station located in Chippewa County on the eastern edge of Michigan's upper peninsula. The area can be characterized by Isolated, low, rounded ridges or hills intersected by broad, swampy valleys and lakes (Veatch, 1927). Most of the present surface features are the result of glacial drift and lacustrine deposits laid down during the last glacial stage. The general climate of the area has a short growing season with moderate summer mean temperatures of 63°P and long, frequently rigorous winters. Annual precipitation averages about 31 inches, with somewhat greater amounts occurring in the summer and fall than in the viinter and spring. Average snowfall is about 96 Inches Summary, 1942-1961). (Climatological The frost-free season averages about 116 days, but light frosts may occur throughout the growing season. The prevailing winds are westerly. One of the many glacial basins located in the area served as the study location. The basin extends north and south with a parallel sandy and gravelly ridge located between it and the St. Mary's River (Figure 1). Soils vary 1. AERIAL VIS'/I OF STUDY AREA SHOWING DRAINAGE DITCH AND LOCATION 1969). from very Tine sand to silty clay overlain with shallow deposits of peat and muck* Impervious soils, poor natural drainage, and drainage from surrounding uplands onto the area combine to create its present edaphic characteristics. Mr. Maurice Day, research forester on the station, relates that logging and fire have been excluded from the area for over 30 years. The vegetative type is classified as a shrub-carr (Curtis, 1959; White, 1 9 6 5 ). Alnus rugosa and Fraxinus nigra are the principal overstory species, and Cornus stolonifera the main low shrub. Principal ground vegetation genera are C a r e x , Gl yceria, I m patiens, A s t e r , and Solidago, similar to that described for a sedge-alder swamp by Damman (1964). CHAPTER III. METHODS A biotic Sampling Procedure During the summers of 1961 to 1 9 6 3 * & shallow drainage ditch (40 cm mean depth) the study basin was plowed through the center of (Figures 2 and 3 ). Drainage of the water in the ditch moves in two directions; to the north and to the southj both sections of the ditch emptying into the St. Mary's River. The study area drains to the south. Two study units were first located on aerial p h o t o ­ graphs, then located in the field using recognizable landmarks found on the aerial photographs. extends 130 meters Each unit (m) adjacent to and 92 m perpendicular to the ditch (Figure 4). Available soil maps indicate a Bergland silty clay loam on one area (Site 1), and a Bruce very fine sandy loam on the other (Site 2 ).1 Four 1 m water wells were randomly located along each of three lines which parallel the ditch at distances of 7.6 m (Line 1), 45*7 m (Line 2), and 91.4 m (Line 3) to determine the effectiveness of the ditch in lowering the water table. Wells were not placed within any sample U. S. Department of Agriculture, Service soil survey, 1964 6 Soil Conservation 7 FIGURE 2. EARLY SPRING VIEW OF DRAINAGE DITCH ADJACENT TO SITE 1. NOTE SPECIES MIXTURE OF FRAXINUS NIGRA AND AT.NUS RUGOSA ON SITE TO RIGHT OF DITCH. FIGURE 3. EARLY SPRING VIEW O F DRAINAGE D I T CH ADJACENT TO SITE 2. NOTE PREDOMINANCE OF ALNUS RUGOSA ON SITE TO LEFT OF DITCH. 8 Site 1 a • = o *— Quadrat Well Drainage Ditch Precipitation Gauge Direction or Hater Plow I'* 7.Sen Site 2 I 7.6m J 38.1a O -- * r O D 03 FIOURE DISTRIBUTION OF QUADRATS, WELLS, AND PRECIPITATION GAUGES IN RELATION TO DRAINAGE DITCH FOR SITES 1 AND 2. quadrat to avoid disturbance of the vegetation. A 3-f°°t length of sbeel pipe was driven into the soil beside each well, and four similar pipes were placed in the center of the ditch along each study site. nating area, An aluminum tag, d e s i g ­ line and well number, was placed on each of the steel pipe markers. The relative elevation of the top of each pipe was then determined with a transit and stadia rod to use as a reference elevation for measuring the fluctuation of the water table. Shelters for Belfort continuous water level recorders were located over one well on each line of Site 1 to obtain an estimate of rates of recharge and discharge of the water from the area. Two functional recorders were available for use; one was placed on l.ine 1 for the entire summer, and the other was rotated between bines 2 and 3* A precipitation gauge was located near each of the study units in a cleared area. All obstructions were removed by a distance at least three times their height from the gauge (Figure 5)• W a t er levels in the wells and precipitation measurements were made weekly throughout the 1969 growing season, except during one period of very intensive precipitation when water levels were the highest recorded on both study sites. The ground water from ditch and well points was analyzed for both oxygen, in milligrams per liter (mg/1), measured in the field with a portable galvonic cell oxygen analyzer, and pH, measured with a Beckman electrode in the FIGURE 5. PRECIPITATION GAUGE LOCATED ON CLEARED AREA NEAR SITE 2. 11 laboratory. All measurements were made in the upper 15 centimeters (cm) of the ground water. Soils were checked in the field down to 100 cm, and depths of organic matter and changes in texture and structure were noted. The study area was flown in May, physiographic features, graphic record. 1 9 6 9 , to observe the the study sites, and for a p h o t o ­ Weather balloons, four feet in diameter, were used to m ark the corners of the study sites. Biotic Sampling Procedure Within each study unit, ten 4 by 4 m quadrats were randomly located along each of three lines which parallel the ditch. Bach quadrat was permanently marked with a 3 - foot length of steel pipe and numbered with an aluminum tag designating the study site, line, and quadrat number. A one square meter quadrat vias located in each of the larger quadrats utilizing the steel pipe marker as one of its corners. All plant species found on the two sites were identified 1 9 5 0 )9 and specimens placed in the (Fernald, Beal-Darlington herbarium at Michigan State University for future reference. Woody species less than 1 cm basal diameter and herbaceous species were sampled on each 1 m^ quadrat (Figures 6 and 7)* Density, frequency, and above-ground dry weight biomass were determined for each species found. All clipping was accomplished in two weeks during the 12 FIGURE 6 . A I M 2 QUADRAT ON LINE 2 OF SITE 1 BEFORE CLIPPING. THE DOMINANT GRASS SPECIES IS GLYCERIA STRIATA; WOODY STEMS ARE ALNUS H U G O S A . FIGURE 7. A I M 2 Q.UADRAT ON LINE 3 OF SITE 1 AFTER CLIPPING OF ALL ABOVE-GROUND HERBACEOUS AND W O ODY VEGETATION («Cl CM BASAL DIAMETER). 13 period oi' maximum above-ground biomass August). Site 2. (late July and early Quadrats on Site 1 were clipped before those on The order of clipping quadrats was randomly assigned to minimize sampling error due to temporal changes in the vegetation. Samples were first air dried and then oven dried at 105°C for 24 hours before weighing. A collapsible 1 m 2 plot frame similar to that described by Blair was used (1 9 6 3 ) to delimit the edges of each quadrat. Diameters at breast height (1.37 ni) were measured with calipers for each stem of all tree species on each of the 4 by 4 m quadrats. The dominant high shrub^ Alnus r u g o s a 3 was measured as a tree. Each stem of each clump for this species was measured separately, and number of clumps per quadrat were tallied. Heights of Alnus and Fraxinus were measured to the nearest 0.5 m. Less abundant shrub species were counted individually or by clumps. Above-ground biomass determinations of Alnus and Fraxinus were made on three of the ten 4 by 4 m quadrats that were randomly selected on each line. One stem from each 2 cm size class found on each quadrat was clipped to obtain the range of size classes for each species. avoid sampling bias, To the stem located farthest to the north on each quadrat for each size class was taken as the sample stem. Stems occurring on quadrats other than the biomass quadrats, but which constituted a larger size class, were also sampled. 14 A chain saw was used to cut each stem at ground level. Branches were clipped from the stem and separated into five size classes. A representative branch was selected from each size class and separated into fruit and flowers, current growth (twigs plus leaves), wood. Total length was measured live wood and dead to the nearest 10 cm for each representative branch, and basal discs were clipped from all remaining branches. Diameters measured to the nearest 1 millimeter (outside bark) were (mm) just above the butt swell for each of the five representative branches and for all basal discs from the remaining branches. The last five growth rings were measured to the nearest 0.1 mm. for each representative branch to determine wood production by weight (Whittaker, 1 9 6 5 )- All growth rings were measured on branches less than 5 years of age. Component parts were then oven dried at 105°C to constant weight. After removal and field measurement of all branches, the stem was measured for total length (nearest 0.5 m) and cut into 1 m sections. Total green weight was determined to the nearest 1.0 gram (g) (Figure 8). taken from each 1 m section and weighed. Sample discs were Laboratory measurements of the stem discs included bark thickness, diameter (inside bark), width of last six growth rings (growth ring for 1969 measured separately), and oven dry weight (105°C to constant weight). A representative sample of the stem discs from each species was separated into wood and bark for oven dry weight ratio determination. Moisture FIGURE 8. FIELD DETERMINATION OF STEM GREEN WEIGHTS FOR ALNUS RUGOSA AND FRAXINUS NIGRA. 16 contents of stem discs were used to determine total dry weight for each sample stem. Methods of woody stem analysis generally follow those of Whittaker and Woodwell (1968, 1969)• Foliar samples were collected from the dominant woody an^ herbaceous species for nutrient analysis 1964; White, 1958) . (Kenworthy, Analysis of samples was made in the Plant Analysis Laboratory at Michigan State University. Nitrogen was determined by the micro-Kjeldahl method, potassium by flame photometer, and all other elements by direct-reading spectrograph. Sixty basal stem discs (one from each quadrat) rugosa were collected in mid-September, ring analysis. 1969, for a growth Discs were air dried, sanded with 300 grid paper, and marked into four quadrats 1968 ). of Alnus (Stokes and Smiley, Ring width along each of these marks was measured to the nearest 0.1 mm. Analysis of Data All data were entered on computer forms and punched on cards for analysis. The Michigan State University Agricultural Experiment Station Computer Stat Series was used for statistical analysis and computation of regression equations. Where necessary, programs were written to perform other computations. Orthogonal comparisons were made for species differences betvieen quadrat lines and for water levels, pH and oxygen 17 in wells at the three distances from the ditch. The statistical design is a randomized complete block utilizing the study sites as blocks and the lines of quadrats and wells as treatments (Steel and Torrie, i9 6 0 ). The following calculations were made on data collected for each species: 1. Herbaceous species p a. Above-ground dry weight b. Density c. Frequency of quadrat occurrence d. Importance value (no/m2) the sum f 3) Total number of individuals duals per species (%) (relative dry weight + relative density + relative frequency, e. (g/m ) (N), number of indivi­ (n^), and number of species (S) found on each m s quadrat were used to calculate community diversity (D), diversity per individual minimum diversity equations (D), maximum diversity (Dm ^n ) and redundancy (VJilhm., 1 9 6 7 ; Patten, s D = log Ni -*51 log n< ] 1=1 D = (i)(log Ni - £ (R) in the following 1 9 6 2 ; Pielou, 1969): log n± !) Dm a x = loS Ni - s log (§) i D min = 1°S Ni - log N - (s - 1 ) J (Om a x)> 1 9 6 6 ; Margalef, l8 cm a x " % i n These same equations were then used to compute equivalent values based on biomass for each species per m (Wiliam* 1 9 6 8 ; Dicltman* 1 9 6 8 ) . 2. Woody shrubs (<1 cm basal diameter) and trees (<1 cm d .b .h . ) n a. Above-ground dry weight (g/m ) separated into current and previous growth 3. 4. b. Density (no/m^) c. Frequency of quadrat occurrence Shrub species (>1 cm basal diameter) a. Density (clumps and individual stems/ha) b. Frequency of quadrat occurrence ($) Tree species a. Density (clumps and individual stems/ha) b. Frequency of quadrat occurrence c. Basal area d. Importance value (m2/ha) (relative basal area -t- relative density + relative frequency* 5- {%) the sum ~ 3 ) Alnus rugosa and Fraxinus nigra a. Above-ground dry weight (1) (g/m^) Stems - Percent moisture of stem discs and total stem green weight were used to estimate total dry weight; for each sample stem. Linear regressions of e s t i ­ mated total dry weight on parabolic volume X tree height* (stem basal area the product 7 2) were computed separately 19 for the two species on each site. These regressions, wood/bark weight percentages and parabolic volumes for stems on each quadrat were used to determine total dry weight (g/m2 ) for the two species on each line of the two study sites. Total volume of wood per sample stem for each 1 m section was computed using the Smalian formula the areas, (means of inside bark., at the end of each 1 m stem section times the l e n g t h ) . Volumes of sections were then summed to obtain the total wood volume for each sample stem. The volume of wood included in the mean increment for the last five years, not including 19^9 growing season due to early sampling date, was computed for each stem section with the following formulas: V = (a^ + a 2)L a^ = area of mean increment at the end of stem sections A = area of wood at end of each stem section r = mean width of growth rings at end of stem section R = mean radius at end of each stem section V = volume increment L = length of stem section Volume increments for each stem section were then summed to give total growth volume for each sample tree. Regressions 2 of the volume estimates on DBH 4- DBH were used to compute 20 volume of wood ( c m V m 2 ) and volume increment (cm3/m2 / y e a r ) for the two species on each line of both study sites. An estimate was made of stem wood weight produced (g/m2 /year) by: Est. Total Dry VJt. (g/m2 ) X Est. Vol. Increment Total Wood Vol. (cm3/m2 ) (2) weight Branches (cm3/m2/year) - Independent estimates of current (twigs + leaves) and dry wood weight + bark for branches of each sample stem were obtained by regressing the log10 of 10 times current dry weight and the log1Q of lO times dry wood weight + bark on the log1Q of 10 times the basal diameter of each representative bra nc h for each species on each site, and by using these regressions along w it h basal diameters of all branches on each sample stem. Estimated dry weights of current growth and wood + bark for branches of each sample stem were then regressed on D B M + D B H 2 for each stem by species and site. These regressions were then utilized to compute total current p growth and wood weight + bark of branches per m c for both species on the two sites. Annual weight increment of representative branch wood + bark was obtained by regressing the log-^Q of 10 times the mean weight of the last five growth rings including bark (less than five rings in younger branches) l o g10 -*-0 times the basal diameter. on the Total branch wood increment for each stem was then computed utilizing basal diameters of all branches on each stem and the above 21 regressions. This total was then regressed on DBH + DBH2 of each sample stem to obtain an estimate of branch wood + bark, weight increment (g/m2/year) site. for each species on each Branch data were multiplied by 10 to avoid zeros when taking the log of 1 . b. Growth ring analysis of Alnus - A growth ring index was computed for two of the growth ring width series measured on the basal discs of Alnus as outlined by Pritts (1966). Index values correct for variability in width of rings due to factors other than climate, crown size. such as age and It does not eliminate such variability as sampling error. Index values were computed by first regressing each ring width series on year and then dividing the actual ring width into its corresponding value on the regression line. The two index values for each year on each stem were used to obtain an average Index value per quadrat. CHAPTER IV. RESULTS AND DISCUSSION Present State of the System Abiotic Factors The ground surface on the two sites iu typical of poorly drained areas with hummocks and depressions scattered throughout (Figures 9 and 10). The average soil surface slopes about 1 percent on both study sites. Downslope is to the east on Site 1 and to the west on Site 2. Mean depth of the black, mucky organic layer (includes A^ horizon) was 25.6 cm on Site 1 and 27.0 cm on Site 2. This was underlain by a 10 to 15 cm strongly leached, bluegray A^ horizon on both sites. The A^ plus the B horizon on Site 1 was a silty clay loam texture and averaged 62 cm in thickness. This corresponds to a very fine sand horizon on Site 2 which averaged 52 cm. Little variation was found in thickness of the A2 and B horizon over each study site. Total precipitation measured from June 18 to September 17 during the 1969 growing season was 2 5 .^ cm (Figure 11). Amounts measured on the two sites never varied more than 0.3 cm and were well distributed over 22 23 FIGURE 9. EARLY SPRING VIEW OF LINE 2, SITE 1 SHOWING VARIABLE GROUND SURFACE. LARGER STEMS NEAR CENTER ARE FRAXINUS N I G R A : STEMS IN RIGHT FOREGROUND ARE ALNUS R U G O S A : LOG IS BF.TULA PAPYRIFERA. FIGURE 10. EARLY SPRING VIEW OF LINE 2 3 SITE 2 SHOWING GROUND SURFACE. STEMS ARE PREDOMINANTLY ALNUS R U G O S A : DEAD SNAG IN CENTER BACKGROUND IS POPULUS BAISAMIFERA. Relative Elevation (cm) 24- Site 1 300 Ground Surface 200 260 ■ 240 220 Site 2 Relative Elevation (cm) 300 200 ■ Ground Surface 260 240 220 200 100 PPT (cm) 6 4 2 JL n IL 6/10 7/2 IL 7/16 7/30 0/13 Date of Measurement n Q/27 9/10 FIGURE 11. MEAN RELATIVE ELEVATION O F GROUND SURFACE AND WATER TABLE AND AMOUNT OF PRECIPITATION PER WEEK FOR BOTH SITES, 1 9 6 9 . 25 the growing season except for two-week periods in mid-July and late August. Recharge of the water table with precipitation depended on its initial height at the time precipitation started and the duration and intensity of the precipitation (Figure 11). Bay (1 9 6 6 ) found this same relationship in bogs of northern Minnesota. If the water table was near the soil surface at the time precipitation o c c ur r ed 3 height of net water table rise was less, due to more direct runoff, than if the same precipitation occurred when the water table was well below the soil surface. This assumes the same moisture conditions in the overlying soil stratum. Both sites remained wet through the middle of July for the 1969 growing season. During this period the mean water table was 2.8 and 11.5 cm below the soil surface on Sites 1 and 2, respectively. (Table 1) . This difference was significant (e<=.01) During this wet p e r i o d d e p r e s s i o n s in the soil surface remained inundated while most of the hummocks remained exposed; however, contrast between hummocks and depressions was not as great on Site 2 as on Site 1, probably due to better drainage characteristics on Site 2. Both sites were completely Inundated on June 27 after a very intense rainfall. After July 16, water levels began to drop more rapidly due to decreasing amounts of precipitation and increasing evapotranspiration rates. Water levels on Site 2 dropped to below 60 cm of the soil surface by TABLE 1 . ANALYSIS OF VARIANCE TABLES FOR FACTORS MEASURED ON TWO SITES AND THREE DISTANCES FROM DRAINAGE DITCH. 1969 Line Line Hell Remaining Rep Rep Line Rep Error X X X X X X (Site) Line Quad Line Quad Quad Hell Week Hell Week Week (df) Degrees of Freedom (df) Herbaceous Species Dry Height Stems Species Dive r si ty/Ind ivid ua 1 Stems Redundancy (Stems) Diversity/Ind ividual Biomass Redundancy (Biomass) Tree Species Basal Area Relative Elevation of Hater Table Depth of Hater Table Oxygen in Ground Hater pH of Ground Hater Organic Matter Depth * ** *** T 10 15 1 2 9 2 9 18 T * * *** T #** T ■jp# T T T * T T T * T T T T T T 18 18 18 18 *** T *** T T T T T T T T T 18 18 ** * T T T T 18 * *** T ***- T T T T T *** T T T * * T Significant at .10 Significant at .05 Significant at .01 Tested for factor but not significant T T ** T 3 5 6 T *** T T T 18 101 T T T *** T T 101 f T T 9 9 27 27 July 24 and remained there for the rest of the growing season except for a wet period in early August. Water levels decreased more slowly on Site 1 due to its less permeable soils and physiographic location. Site 1 drains an extensive basin and its surrounding uplands, while Site 2 drains a much smaller area (Figure 1). Other studies have shown similar sites to be fairly eutrophic due to the high nutrient content of incoming ground water from surrounding uplands 1 9 6 5 ; Bay, 1 9 6 7 ). Fierce (Heinselman, 1963j (1 9 5 7 ) believes the relatively high productivity of tree species and stand composition on these sites to result from high levels of dissolved salts and oxygen carried in drainage waters. He found pH to vary from 6.0 to 7.5 and oxygen levels from 0.9 to 4.6 ppm in the upper 15 cm of the ground water. Oxygen levels in this study were found to vary from 1.9 +• 0.8 mg / 1 on Site 1 to 2.0 + 0.8 mg/1 on Site 2. Mean pH was 6.4 4- 0.2 on Site 1 and 6.0 + 0.2 on Site 2. Foliar Nutrients Comparison of foliar nutrient values with those found in other studies gives further evidence that this system is eutrophic (Table 2). Watt and Heinselman (19&5) have shown foliar nutrient percentages to be correlated with site quality. They show a decrease in foliar percentages of nitrogen and phosphorus for black spruce w i th distance from a dense speckled alder stand to a black spruce 28 muskeg. Moizuk and Livingstone nitrogen values (1 9 6 6 ) present foliar for red maple seedlings taken from a bog mat and for those grown in a complete nutrient culture. Percent nitrogen for those from the mat averaged less than 1 ,0 * while those grown in nutrient culture averaged greater than 3 *0 * TABLE 2. FOLIAR NUTRIENT VALUES FOR THE DOMINANT SPECIES FOUND O N EACH STUDY SITE Species Element (Percent Oven Dry Weight) Ca K P N Site Site Site Site Site Site site Site 2 2 1 2 1 1 2 1 Alnus rugosa July 1968*(2 )D Aug. 1969 (6) 2.68 2.99 _c 2.37 0.17 0.20 0.79 0.75 0.92 0.84 Fraxinus nigra July 1968 (2) Aug. 1969 (6 ) 2.26 2.26 2.60 0.14 0.20 1.18 1.24 1.89 1.80 0.26 1.12 Cornus stolonifera July 1968 (2) Glyceria striata July 1969 (6) 2.51 Impatiens capensis July 1969 (6) 2.26 2. 31 - 2.25 - - - - - - - - - - - - - - a Samples collected in July and August 6 Number of samples in each nutrient value c No analysis made 29 Foliar analyses of tree species are not always an indication of the fertility level in soil however, (Wilde, 1 9 5 8 ). foliar nitrogen percentages for all species sampled were in the upper part of the range given for red maple seedlings by Moizuk and Livingstone discussed above. These high levels of nutrients found in the five dominant species - tree, high shrub, medi um shrub, grass and annual of this community indicate this site to be richer in available nutrients than similar stagnant water systems. The Flora The combination of such edaphic factors as nearly neutral pH, high nutrient content, high fluctuating water table, and good aeration result in a rich and distinct flora (SJors, 1959» Conway, 1949)• A search of the flora on both sites revealed 109 species, 6l genera, and 31 families. Gramineae was the most represented family with 10 genera and 15 species, and Carex with 17 species was the most represented genus. Of the 109 species found, 29 did not occur on any sample quadrats. These species were either rare in occurrence or were located only along the ditch banks. D a m m a n 1s (1964) description of a sedge-alder swamp in central Newfoundland presents edaphic factors and a species composition very similar to those found in this study. Glyceria s t r i a t a , which was a dominant grass species on both study sites, was 30 only of’ mi nor importance; and Fraxinus n i g r a , a dominant tree species, was absent from D a m m a n 's species lists. Species Importance Importance value is believed to be a better indication of a species' structural importance since it combines distribution, McIntosh, size and number or individuals 1951; Rice, 1 9 6 5 )- (Curtis and However, it is not necessarily an indication of a species' functional importance in the community. Importance values for herbaceous species Involve relative dry weights, relative densities, and relative frequencies; those for tree species include relative basal area, relative densities, and relative frequency. was determined from quadrat occurrence. Frequency All stems which originated below 1.37 m were sampled as Individuals for woody species, and clumps were used as individuals in grass and sedge species. VJoody species which had a main stem greater than 1 cm at 1-37 ni were considered a tree. Of the 57 herbaceous species sampled, 16 were well distributed over both study sites (Table 3). Glyceria striata and Impatiens capensis vie re the most important species sampled. High importance values for Glyceria are primarily due to its large relative dry weight, vihile those for Impatiens are due to large relative densities. Glyceria is a grass which occurs as clumps, vihile Impatiens is a very succulent single-stemmed annual. Impatiens vias more important on the better drained soils of Site 2 than 31 TABLE 3. IMPORTANCE VALUES® FOR HERBACEOUS SPECIES AT THREE DISTANCES FROM DRAINAGE D I TC H FOR BO T H STUDY SITES Species Distance from Ditch 7.6m 9 1 .4m 45. 7m Site 1 Site 2 Site 1 S'I'te 2 Site 1 S'ite '2 _ . 0.33 1.62 2.54 1.25 0.50 6.59 1.19 0.42 0.26 — 0.32 3.90 2.14 2.87 1.29 1.74 1.43 1.65 — 3-59 0.95 4.78 I .98 2.11 0.31 0.50 — — — - — 2.51 — - — o. 38 0.26 O .38 1.13 1.08 0.39 O .69 2.20 5.18 — 1.61 0.32 1.85 — — — — — 0.76 O .83 1.99 1.54 — — 0.47 — — 5.00 3-17 2.22 4.13 — 2.95 1.14 - 4.82 0.31 __ 1.47 — O .38 0.45 1.93 0 .31 1.89 1.59 — 0.85 1.61 - 5.18 1.83 3.95 1. 38 1.38 . 0.43 2.24 2.26 4.83 l.4o 1.57 0 .34 8.17 0.29 2.71 0.64 2.90 2.53 - - 4784 0.29 0.32 — — — 0.90 — — 0.34 — 1.02 3.61 1.39 0.37 — — 0.36 — — 3.36 0. 68 — 3.46 0.65 0.74 0.29 1.82 4.51 1.50 0.48 — 0.34 — — 1.29 2.84 0 .89 — — — 1.70 0.68 0.58 3.37 — 0.60 3.04 0.67 0.54 0.78 — — — o.4i — — 1.12 _ 2.19 0.31 — - 00 0 * H Agrostis perennans Arisaema atrorubens Aster lateriflorus A. puniceus A. umbellatus Bromus c H i a t u s Calamagrostis canadensis Caltha palustris Cardamine pensylvanica Carex canescens C. crinita C. disperma C. gracillima C. hystricina C. interior C. intumescens C. leptalea C. leptonervia C. projecta C. stipata C . tribuloidcs C . Tuckermani C. vesicaria Chelone glabra Cicuta bulbifera Cinna arundlnacea C. latifolia Dryopteris spinulosa Epilobium glandulosum Equisetum arvense Eupatorium maculatum E. purpureum Fragaria virginiana Galium palustre G. tinctorium G. tririorum - - 2.53 2.57 7.69 4.87 2.09 0.49 2.27 0.33 7.57 2.60 0.39 — 2.75 2.13 32 TABLE 3 (cont'd) Species Distance t'rom Ditch 45 -Tin 7. bin ~ 91.4m Site T- S"it'e"'2 Site" 1' Site "2 'Site l"SiTe 9-60 0.26 23.35 — 18.73 0.29 21.13 _ ;4.63 1.57 0.89 1.77 • 0 N O Glyceria striata llabenaria psycodes Ifieracium aurantiacum Impatiens capensis Iris versicolor Lactuca biennis Lycopus uniflorus Maianthemum canadense Mitella nuda Molinia caerulea Onoclea sensibilis Ranunculus abortivus R. recurvatus Rumex orbiculatus Scutellaria lateriflora Senecio aureus Smilacina trifolia Solidago rugosa Stellaria graminea Viola cucullata V. p a l l e n s ______ 39.35 — 0.42 - 0.43 : - 0.55 1.93 5.36 — 0.58 0.92 — 0.76 3.33 25.01 O .29 4.72 0.32 1.40 5.63 - — 18.21 — 24.24 0.50 1.07 2.99 0.28 2.50 0.52 0.31 0.34 - — 3.06 1.55 — 4.96 8.21 24.30 0.95 3.24 — 1.89 0.52 0.32 1.30 7.63 1.59 6. 31 0.33 0.35 0.90 — 0.64 0.32 2 .31 4.22 0.32 8.37 1.76 6.51 2.27 3.41 1.48 1.66 I .50 4.67 2.61 5-31 2.56 a Relative dry weight + relative density + relative frequency, the sum ^ 3 33 on Site 1, while Glyceria was fairly constant in importance over both sites. The next most important species were Senecio aureus and Solidago r u g o s a . Arisaema atrorubens and Habenaria psycodes were found on Site 1 only. distribution is limited It appears their to the more poorly drained Site 1. The distribution of other rarer species ma y be actual, but may also be due to sampling error. It should be pointed out that importance values are valid only during that part of the season (late July) when data were c o l l e c t e d . An earlier or later sampling date would probably result In a shift of importance values among species due to vernal changes. Curtis (1959) discusses three aspects of such communities occurring in early spring, mid-summer, and late fall. Species such as Caltha palustris dominate the view In early spring. Mid­ summer brings the flower of orchids, grasses and sedges. Solidagos and Asters dominate the late fall aspect. Low shrub and vine species were minor in importance compared to herbaceous species in this community except for localized areas (Table 4). Clematis virginiana was the only vine found which was common on both sites. It usually dies back: to a low woody base each winter, but grows rapidly during the summer, overtopping crowns of overstory species in a few instances. Ribes americanum was the most common of the four Ribes species found, but still m inor In occurrence. Rubus idaeus apparently does 34 best on the better drained soils of Site 2, while Rubus pubescens occurs commonly over both sites. This latter species is one that is found almost exclusively on hummocks, but sends out runners to colonise depressions during the drier summer seufc.cn. TABLE 4. PERCENT FREQUENCY OF QUADRAT OCCURRENCE FOR LOW SHRUB AND VINE SPECIES ON EACH SITE AT THREE DISTANCES FROM DRAINAGE DITCH Low Shrub & Vine Species Distance from Ditch 45': 7m 91 .4m 7 . 6m Slt'e 1 Site 2 Site 1 Site 2 Site l Site 2 Clematis virginiana 20 — 60 60 10 40 Rhamnus alnifolia - - 10 - 10 - Ribes americanum 10 - 10 10 20 10 R. glandulosum 10 - 10 - 10 - R. hirtellum - - - - - 20 R. triste 10 - - 10 - 10 Rubus idaeus 10 4o 20 60 - 70 R. pubescens 6o 70 100 100 8o 80 Cornus stolonifera was the most prevalent species in the second story of the community (Table 5)• Alnus is considered to be a third-story species since it dominates the crown surface of the community. Ilex verticillata, the only other second-story shrub sampled, was not found on Site 2. 35 TABLE 5PERCENT FREQUENCY OF QUADRAT OCCURRENCE FOR TREES AND SHRUBS ON TWO SITES AND THREE DISTANCES FROM DRAINAGE DITCH, 19^9 Species Distance from Ditch . bm 91 .4m 7 45 7m Site 1 Site 2 Site 1 Site 2 Site 1 Site 2 . Tree Species8 Abies balsamea Alnus rugosa Betula papyrlfera 10 100 - 100 - Fraxinus nigra 70 Larix laricina 10 - 30 Populus balsamifera 20 20 10 1O0 100 1O0 - 90 - 90 - - 20 4o 30 4o 10 20 - Salix gracilis - - - - S. pyrifolia - - - - - - - - Viburnum trilobum - 60 - - Ulmus americana - - - - 20 70 100 - Prunus virginiana S. serissima 10 - 10 - - - 4o - - 10 - - 10 - 10 - - 10 - Shrub Species13 Cornus stolonifera 4o Ilex verticillata 30 20 f* Steins > 1 cm d.b.h. b Stems > 1 cm basal diameter - 20 50 4o 20 - 10 70 - 36 Table 6 emphasizes the basal sprouting characteristic of the dominant woody species in this community. The tight canopy, heavy herbaceous cover, and poor moisture conditions found in these communities are not conducive to high seedling survival. Table 7 gives some indication of the number of new seedlings and sprouts occurring on the sites. They seem to be in sufficient number for Alnus to maintain its dominant position in the community. sprouts Distribution of seedlings and for Fraxinus and Fopulus correspond with the present distribution of old established trees. The presence of Populus on the 7.6 m transect of Site 1 was due to nevj seedlings. The same is also true for Ulmus americana on the 45.7 m transect of Site 2. This species occurred on Site 1 also, but was rare in occurrence. Of the 12 tree species sampled, distributed over both study sites only two were well (Table 8). Alnus r u g o s a , a high shrub, was the most important, and Fraxinus nigra was next most important. Site 2 was more heavily dominated Alnus than was Site 1. All other tree species were sporadically scattered over the sites. Fopulus balsamifera and Larix larlcina were mostl y large, over-mature individuals which resulted in large basal areas and importance values (Table 9)* Successional Change It appears that m uc h of Site 1 is In a transitional stage of succession leading from an A l n u s -dominated TABLE 6. NUMBER OF CLUMPS AND STEMS FOR TREES AND SHRUBS ON TWO SITES AND THREE DISTANCES FROM DRAINAGE DITCH, 1969 _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ Distance from Ditch_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 7.6m AE.7m Q1.4m Species Clumps Steins Clumps Stems Clumps Stems _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ ■ - — ■— - — - - - - - - - - - (no/ha) — - - - Tree Species8 Abies balsamea Site 1 125 125 125 125 125 63 Site 2 125 125 Alnus rugosa Site 1 10,812 9,688 3,687 3,687 15,562 4,125 ll,2p0 Site 2 3,625 3,562 10,125 3,750 12,437 Betula papyrifera Site 1 63 63 Site 2 Fraxinus nigra 2,000 Site 1 3,687 2,375 3,375 3,875 4,063 625 3,000 625 Site 2 3,125 313 313 Larix laricina Site 1 63 63 Site 2 Populus balsamifera Site 1 1,312 1,125 Site 2 687 625 187 187 Prunus virginiana Site 1 125 125 Site 2 625 313 313 313 Salix gracilis Site 1 Site 2 63 313 - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - TABLE 6 (cont'd) Distance from Ditch 46.7m Stems Clumps 7.6m Species Clumps Tree Species (cont'd) Salix pyrifolia Site 1 Site 2 Salix serissima 2p0 Site 1 Site 2 Ulmus americana Site 1 Site 2 125 Viburnum trilobum Site 1 Site 2 Total Tree Species Site 1 6,125 Site 2 4,063 5,094 Site Average Shrub Speciesto Cornus stolonifera 562 Site 1 Site 2 125 Ilex verticillata 500 Site 1 Site 2 Total Shrub Species Site 1 1,062 Site 2 125 Site Average 594 a Stems >1 cm d.b~.hu to Steins >1 cm basal diameter Stems 91.4m Clumps Stems o3 125 - - - - - - - - - - 43? - - - - - - - - - - - - - - - 125 187 250 - - - - - - - 63 63 - 187 313 - - - 13,812 11,688 12,750 7,499 7,812 7,656 13,876 14,937 14,406 9,126 4,938 7,032 21,500 13,938 17,719 1,312 812 375 1,750 500 4,187 250 1,312 437 4,125 1,000 125 250 63 313 - - - - 750 4,187 2,468 313 1,312 812 750 4,125 2,338 - 2,312 812 1,562 . 500 1,750 1_,125_ 39 TABLE 7• NUMBER OF SEEDLINGS AND BASAL SPROUTS OF TREE SPECIES AT THREE DISTANCES FROM DRAINAGE D I T C H FOR BOTH STUDY SITES Species Distance from Ditch 4B.7m 01.4m 7. 6m Site 1 Site 2 Site 1 Site 2 Site 1 Site 2 ^no/ *_L iiu )l Alnus rugosa 431 Praxinus nigra 19 Populus balsamifera 12 Ulmus americana 412 500 56 594 85 762 69 588 12 12 IOO 56 TABLE 8. IMPORTANCE VALUES3 FOR TREE SPECIES AT THREE DISTANCES FROM DRAINAGE D I TC H FOR BOTH STUDY SITES Tree Species13 Distance from Ditch 7. 6m 45 . 7m 91. 4m Site 1 site 2 Site 1 'Site 2 Site 1 Site 2 Abies balsamea 4.8 Alnus rugosa 47.6 Betula papyrifera — Fraxinus nigra 33.3 4.8 Larix laricina Populus — balsamifera Prunus virginiana — Salix gracilis S. pyrifolia S. serissima 9.5 Ulmus americana Viburnum — trilobum 83.7 — 13-4 3.8 51.0 — 45.2 2.9 50.4 — 28.3 —. 2.0 46.8 66.4 1.4 21.3 16.8 - — — - - 7.8 23*8 - — 5.2 1.5 — — - 8.2 - - - - - 4.2 - - - 1.5 - — — — - - 2.9 — 5.4 — 4.4 — *“ — 1.7 a Relative basal area + relative density + relative frequency., the sum f- 3 k Stems > 1 cm d.b.h. 4o TABLE 9. B A SA L AREA FOR TREE SPECIES ON TWO SITES AT THREE DISTANCES FROM DRAINAGE DIT CH Species Distance from Ditch 4B. 7m 7. 6m 91. 4m Site 1 Site 2 Site 1 Site 2 Site 1 Site 2 ( /ha^® ,1 , Abies balsamea 0.23 - 0.19 0.12 0.50 Alnus rugosa 4-39 10.28 6.08 7.39 6.58 Betula papyrifera - - - - 0.03 Fraxinus nigra 6.90 1.98 Larix laricina 7-78 - - - — — 1-93 12.71 Prunus virginiana - - - 0.01 0.01 Salix gracilis - - - - S. pyrifolia - - - - - - - Populus balsamifera S. serissima 0.16 Ulmus americana - Viburnum trilobum - - 15.50 - 2.46 1.28 - 1.07 - 0.10 - 0.77 8.37 - - - - 1.12 - 0.05 CO 18.26 - • a 1 m 2/ha = 4-36 ft2 /acre 12.30 - 5.20 H 19-46 - - 5.29 in cu Totals 0.04 11.79 - - 14.30 41 community to a mixed hardwood-swamp c o m m u n i t y , while Site 2 is, except for small areas, still dominated by A l n u s . This change in Site 1 is indicated b y large importance values for Fraxinus and Populus and high numbers of sprouts and seedlings for these two species. Also, the significantly greater and larger standard deviation for basal area of trees on Site 1 is an indication of the presence of more larger-stemmed species such as P o p u l u s , Fraxinus and La rix (Table 10). Brickman (1950) drained clay soils. found that Alnus grows best on well This may indicate that once Alnus colonizes these sites, it will maintain control of the better drained areas for a longer period of time due to its more vigorous growth. Present distribution of species could also be the result of past history of the area, including such factors as fire, logging, and i n s e c t s ) . and pests (disease However, evidence indicates such activities have not occurred on either site since the early 1 9 0 0 's. The oldest stem of Fraxinus clipped for biomass d e t e r m i n a ­ tion had 69 growth rings with no evidence of fire scars. Several others were greater than 50 years of age. Signs of logging were not present on either site, and the only major insect pest is the larch sawfly. This pest m a y be partially responsible for the present distribution of Larix laricina since there are a few dead trees standing on Site 1. From this evidence it appears the sites have had at least 50 years to change in composition without 42 TABLE 10. MEANS AND STANDARD DEVIATIONS O F BIOTIC FACTORS FOR SITE 1 A N D SITE 2, 1969 Basal Area of Trees (m2/ha) ro 0 Site 1 Biotic Factors + 19.6 Site 2 * 14.0 dfc* 4.9 * 162 + 103 Density Herbs Q (no/m ) Vines, Shrubs & Trees8 (no/m2 ) Shrubs ft; T reesb (no/.l ha) Quadrat Di versity 2 Herbs (no/m ) Vines, Shrubs ft: T rees8 (no/m2 ) hi 2 Trees & Shrubs (no/l6 m ) 134 + 75 IT 14 I 10.2 13-9 1,640 -f* 884 14 + 3 1,353 + 511 *** 11 + 2 3 + 2 3 + 2 3 + 2 3 + 2 f* Vines and shrubs < 1 cm basal diameter; trees 1 cm basal diameter; trees > 1 cm d.b.h. *** Significantly different at ,01 level * Significantly different at .10 level ^3 major dis tiirbances. Over that period change in relative importance or woody species has occurred more slowly on Site 2 than on Site 1. The present flora in the un de rs to ry is in part due to these successional changes and in part to the variation in drainage characteristics of the two sites. The significant decrease in number of herbaceous species sampled per from 14 on Site 1 to 11 on Site 2 may also be the result of these two factors. Increase in number of plant species In terrestrial ecosystems w it h advancing succession has been well established 1966; Whittaker, (Weaver and Clements, 1965; Monk, 1938; Schneider, 1967) • Standing water in depressions on these sites is believed to create a mosaic of micro-gradients from depression to hummock. Since Site 2 is better drained than Site 1, standing water remains for m uc h shorter periods of time. Visual observation also indicates the relief change from depression to hummock Is less on Site 2. This may result in a more variable environment for the shallow-rooted herbaceous species on Site 1, resulting in a greater number occupying a given surface area. (1 9 6 7 ) and Clausen Whittaker (1957) discuss changes In species composition along environmental gradients. Biomass and Production Determining biomass and production for woodland communities proves to be a very time-consuming, difficult 44 and laborious operation due to the size and complexity of the plant species. Early studies on this subject were m o s t l y restricted to single species estimates 1957; Ovington and Madgwick, 1959)- (Ovington, Determination of biomass and production for woodlands with mixed composition was started during the past decade et al. * 19^3; Whittaker* species (Ovington* 1 9 6 2 * 1 9 6 3 ). O In this study* above-ground dry biomass (g/m ) and net production (g/m2 /year) were determined for herbaceous species* woody shrubs seedlings 1 9 6 1 * 1 9 6 2 ; Bray* (less than 1 cm basal diameter)* tree (less than 1 cm d.b.h.)* and the two dominant overstory species. Herbs* shrubs* and tree seedlings were clipped from 10 randomly located m 2 quadrats on each of three transects for each site. Clipping started at the time of m a t ur it y for Glyceria striata and Impatiens capensis (latter July)* the two dominant herb species. Their time of maturity was considered to be time of m a x i mu m biomass accumulation for the sites. In temperate communities such as these where all grovith begins anew each spring* standing crop biomass is considered to be a crude estimate of pr od u c ­ tion per year for herbaceous species (Newbould* 1 9 6 7 )- Above-ground dry weight at the time of clipping was, therefore* considered to be an estimate of production for herbaceous species. Current growth of shrubs 1 cm basal diameter) and trees (less than (less than 1 cm d.b.h.) was separated from old growth as a partial estimate of their production. 45 Biomass and production for Alnus rugosa are based on data from 18 and 23 sample stems from Sites 1 and 2, respectively. Values for Fraxinus nigra are based on 21 and 17 sample stems from Sites 1 and 2 5 respectively. Methods of woody stem analysis are similar to those of Whittaker and Woodwell (19 6 8 ^ 1 9 6 9 )• Data for individual sample stems are listed in ap pendix Tables 35 and 3 6 . Regression values based on sample stem data were used to obtain estimates of biomass and production for component parts of Alnus and Fraxinus for each site separately (Table 11). Estimates of relative error for component parts are listed w ith regression values. Stem bark is based on dry weight ratios for wood and ba r k of 32 and 19 sample stem discs for Fraxinus and A l n u s 3 respectively (appendix Tables 37 and 3 8 ). Stem bark was found to constitute 12.6 percent of total stem dry weight for Alnus and 13-5 percent for F r a x i n u s . Biomass and production values give further evidence that Site 1 is further along in its transition from a shrub-carr to a swamp-hardwood community. Average biomass and production for Fraxinus were 642 g/m2 and 23 g/m2 /year greater than for Alnus on Site 1 (Tables 12 and 13). On Site 2 biomass and production were 868 g/m2 and l4l p g/m /year greater for Alnus than for F r a x i n u s . These differences indicate that Alnus is much more dominant on Site 2 than Site 1 where it is a co-dominant with Fraxinus. 46 TABLE 11. REGRESSIONS3 OP SAMPLE STEM COMPONENTS O F ALNUS RUGOSA A N D FRAXINUS NIGRA ON SITES 1 "AND 2 Alnus r u g o s a F r a x i n u s _________Site 1 Site 2 _________Site 1 Regressions on Parabolic Volume Linear, stem dry weight (g)(y) a 292.4962 72.5465 b 4 9 4 3091.4463 482,449.7525 r2 0.901 0.987 e 0.247 0.124 o 17 23 nigra Site 2 (m3)(x) 427.4615 451,546.3499 0.995 0.070 21 185.5513 433,684.0605 0.988 0.148 16 Regressions on DBH -f D B H 2 (cm) Quadratic, stem volume (cm3) (x + x2 ) a 397.8500 -783.4718 1,203.0956 b -338.8211 400.5638 -850.5234 b2 275.1847 146.3068 322.0462 r2 0.859 0.991 O.98I e 0.350 0.095 0.139 o 18 23 21 -31.9751 - 363.5160 291.0194 0.989 O.l4l 16 Quadratic, stem growth volume ( c m V y e a r ) a -8.4658 - 162.2632 -97.4949 b 18.5201 97.4121 57.0271 b2 16.1107 6.7643 7.7283 r2 0.857 0.880 0.713 e 0.310 0.339 0.525 o 18 23 21 -517.6971 252.5635 -7.0491 0.735 0.530 16 Quadratic, branch wood weight (g) a -92.1850 102.9558 b 44.3166 -I6 l.4 l65 b2 27.6811 70.2989 r2 0.816 0.989 e 0.467 0.144 o 17 23 36.7907 -177.7236 67.6788 0.958 0.302 16 396.9873 -216.7656 55.4556 0.835 0.452 21 Quadratic, branch current twig and leaf weight a -37.4690 -84.2034 -75.3251 b0 43.4742 49.0586 100.3852 b 5.0968 12.2450 0.7528 r2 0.847 0.983 0.844 e 0.311 0.130 0.264 o 17 23 21 (g) ^ - 1 3 1 -2566 74.3974 9.9921 0.959 0.226 16 47 TABLE 11 (cont'd) Alnus rugosa Site 2 Site 1 Fraxinus nigra site" £ Site 1 tr1 0 Oq H 0 Regressions on Log;LQ Branch Basal Diameter A B r2 E o branch wood weight (g) X 10 -6.8492 -0.7707 3.2146 3.0573 0.924 0.951 1.56 1.48 77 109 (cm) -0.7626 3.1583 0.943 1.59 80 -0.9904 3.3583 0.935 1.65 100 Log-, 0 branch current twig and leaf weight X 10 O .1838 0 .1824 0.4224 A B 1.8447 1.9617 2.0299 r2 0.742 0.704 0.777 E 1.96 1.65 1.97 o 100 77 109 X 10 (s) L o g 10 branch wood weight production (g/year) X 10 0.7526 A 0.1191 0.3109 1.2286 B 1.3031 0.8951 r2 0.541 0.775 0.539 2.52 E 1.63 2.31 o 108 100 77 0.2409 2.0421 O .769 1.98 80 0.6822 1.0435 0.650 2.02 79 Regressions on Logio Stem D B H X 10 (cm) LogiQ branch wood production X 10 (g) A 0.5794 1.3784 0.1593 B 1.6308 0.9266 1.2675 r2 O .892 0.920 0.718 E 1.18 1.31 1.45 o 21 17 23 0.7957 1.3063 0.829 1.54 17 a Regressions are in the forms: (linear) y = a 4- bx, (quadratic) y = a + b x + b 2 x, and (logarithmic) log10 (y*1 0 )= A + B logjLQ (x*10) . The independent variable (x) is listed in the heading for each section, and the dependent variable (y) is listed with each individual regression. Breastheight diameter (d.b.h.) was measured at 1.37 m above ground. Coefficients of determination are listed as r2 ; estimates of relative error as e (standard error of the e st imate/mean(y)) for linear and quadratic, and E (antilog of standard error of estimate) for logarithmic regressions. Number of observations are listed as o. TABLE 12. DRY WEIGHT BIOMASS. NET PRODUCTION, AND RELATIVE DISTRIBUTION BY COMPONENT PART FOR ALNUS RUGOSA ON TWO SITES AND THREE DISTANCES FROM DRAINAGE DITCH, AUGUST, 1969 _ _ _ _ _ _ _ Site 1_ _ _ _ _ _ _ _ _ Site 2_ _ _ _ _ _ _ - - - - - - - - - - - - - Distance from D i t c h - - - - - - - - - - 7.6m 4-5.7m 91.4m Average 7.6m 45.7m 91.4m Average Percent Distribution of Components Site 1 Site 2 Biomass (dry g/m2 ) Stem Wood 642 841 930 804 1,070 759 866 898 64.9 56.2 Stem Bark 93 121 134 116 154 109 125 129 9.4 8.1 155 236 237 209 476 304 338 372 16.9 23.3 86 112 130 109 239 170 188 199 8.8 12.4 1,939 1,342 1,517 1,598 100.0 100.0 Branch Live Wood & Bark Current Twigs & Leaves Total 976 1,310 1,431 1,238 'oduction (dry g/m2/year) Stem Wood 65 82 95 81 108 78 88 91 37.9 25.8 Branch Wood & Bark 20 m CM 30 24 49 37 42 43 11.2 12.2 Current Twigs & Leaves 86 112 130 109 254 186 216 219 50.9 62.0 171 217 255 214 411 301 346 353 100.0 100.0 Total TABLE 13. DRY WEIGHT BIOMASS, NET PRODUCTION, AND RELATIVE DISTRIBUTION BY COMPONENT PART FOR FRAXINUS NIGRA ON TOO SITES AND THREE DISTANCES FROM DRAINAGE DITCH, AUGUST, i960 Site 1 Site 2 7.6m ^5*7m 91• ton Average 7.6m 45.7m 91.4m Average Percent Distribution of Components Site 1 Site 2 Biomass (dry g/m2 ) Stem Wood 1,019 1,770 801 1,197 303 648 330 427 63.7 58.5 Stem Bark 159 276 125 187 47 101 52 67 9.9 9.2 Branch Live Wood & Bark 299 535 235 356 125 234 140 166 18.9 22.7 Current Twigs & Leaves 120 193 108 l4o 40 124 47 70 7.5 9.6 515 1,107 569 730 100.0 100.0 28 . 29.1 25.0 Total 1,597 2,77^ 1,269 1,880 Production fdry g/m2/year) Stem Wood 60 98 48 69 15 54 16 Branch Wood & Bark 23 37 24 28 7 27 8 14 11.8 12.5 Current Twigs & Leaves 120 193 108 140 4o 124 47 70 59.1 62.5 Total 203 328 180 237 62 205 71 112 100.0 100.0 50 Relative distribution of* component parts are approximately equal for both species on both sites. Biomass was composed of approximately 6 l percent stem w o o d , 9 percent stem bark, 21 percent branch wood plus bark, and 9 percent current twigs plus leaves. Production by component averaged approximately 29 percent stem wood, 11 percent branch wood plus bark, and 60 percent current twigs plus leaves. Biomass accumulation ratios, the relationship of total biomass to production, are an indication of stature and structure development in terrestrial communities. Values range from 1.0 for herbs through 2.0 to 10.0 for shrubs of increasing size, and above 10.0 for trees (Whittaker, 1962). Values of 5.8 on Site 1 and 4.5 on Site 2 for Alnus characterize it as a shrub of intermediate size. Corresponding values for Fraxinus of 7*9 and 6.5 indicate its very small stature as a tree. Diameter, height and age are further evidence of the small stature of these two species. Mean diameter (1.37 m above ground) and height were 2.3 cm and 3*5 m on Site 1 and 2.9 cm and 4.0 m on Site 2 for A l n u s . Maximum diameter and height for the species were 5*1 cm and 5*5 m respec­ tively on Site 1, and 5*4 cm and 5*5 m on Site 2. Age ranged from 10 to 25 years for canopy individuals on both sites. Mean diameter and height for Fraxinus viere 4.5 cm and 5*0 m on Site 1 and 4.9 cm and 5.0 m on Site 2. Maximum diameter and height were 12.0 cm and 9*5 m on 51 Sito 1, and 15-3 cm and g.O m on Site 2 for this species. Canopy trees ranged in age from 40 to 69 years. Biomass and production values presented in Table 14- are a conservative estimate of the total net above-ground primary biomass and production for the community. Biomass and production of all subterranean organs and of the rare woody species are not included. bark: production, not included. Also, estimates for stem losses to animals and dead branches are Whittaker (1 9 6 8 ) found about 9 percent of annual production is consumed b y animals, and 52 percent of the total biomass is concentrated in the subterranean organs. Pie believes this large concentration in the roots to be due to the basal sprouting characteristic of the oak-pine forest involved. Percent biomass in the roots is not considered to be that h igh in this study due to the high water table. matter layer Roots were not found belovi the organic (25 to 27 c m ) . However, they are believed to be higher than the 20 percent given by Ovington (1 9 6 2 ) for mature forests due to the basal sprouting characteristic of the community and its y o un g age. Herbaceous root biomass has been shown to be approximately three times greater than above-ground biomass (Bray, 1 9 6 2 ). Above-ground biomass and net production range from 2,000 to 20,000 dry g/m^ and from 200 to 1,200 dry g/m^/year for woodland and shrubland communities (Whittaker, community, 1970). Values for production in this 545 g/m^/year for Site 1 and 572 g/m2/year 52 TABLE 14. DRY WEIGHT BIOMASS, NET PRODUCTION, AND PERCENT DISTRIBUTION BY SPECIES CATEGORY FOR E AC H SITE Site 1 Biomass Percent Dis tribution Site 2 Percent Distribution (dry g/m! !l 83 2.6 94 3.8 Vines, Shrubs & Trees8 34 H 1.2 0m Herbs 29 Alnus rugosa 1,238 38.3 1,598 65.2 Fraxinus nigra 1,880 58.1 730 29.8 Total 3,235 100.0 2,451 100.0 Production (dry g/m2 /year) Herbs 83 15.2 94 16.4 Vines, Shrubs & Trees8 11 2.0 13 2.3 Alnus rugosa 214 39.3 353 61.7 Fraxinus nigra 237 43.5 112 19.6 Total 545 100.0 572 100.0 Biomass Accumulation Ratio*5 5-9 4.3 ® Vines and shrubs < 1 cm basal diameter; trees < 1 cm d.b.h. ° Biomass/Production 53 for Site 2, fall in the upper part of this range if allowance is made for that production not sampled. plant production Other (vascular plant species not sampled) make up a small part of the structure of the plant community, and they probably would not add more than lOO g/m2 /year to the production. Biomass values, 3*235 g /m2 for Site 1 and 2,451 g/m2 for Site 2, fall in the lower part of the above range, Biomass accumulation ratio for the community was 5.9 on Site 1 and 4.3 for Site 2. This ratio would probably increase on Site 1 if L arix and Populus were included in the biomass and production estimates since they were mostly large, over-mature individuals. Whi tt a ke r (1 9 6 3 ) gives biomass accumulation ratios of 2.5 to 11.0 for heath balds and forest heaths of the Great Smoky Mountains. I Effects of Drainage 1■ ' — ■' ... - Abiotic Factors Drainage of swamps and bogs may affect such edaphic factors as hydrology, nutrient availability. oxygen content, pH, temperature and These changes may result in a change in productivity and structure of the vegetation (Arkhipova, 1955; Pyavchenko, 1942; Khainla, 1957; I.eBarron and Heetzel, 1 9 6 3 ). Abiotic and biotic factors were 2 measured along transects of water wells and 1 m quadrats to determine the effect of the drainage ditch. 54 The drainage ditch averaged about 40 cm in depths extending into the B horizon of the soil profile on both sites. This depth is shallow compared to other studies reported in which ditch depths usually averaged 1 m or greater. Water movement in the ditch adjacent to Site 1 was variable. The northern third of the ditch on this site was nearly level and, therefore,, drained more slowly than the southern two-thirds. This northern portion was approaching the point at which water drained to the north instead of the south in the basin. Water movement in the ditch adjacent to Site 2 was uniform. Water table level exceeded the depth of the wells on Site 2 after July 16 and on Site 1 after August 20 for varying periods of time (Figure 12). Therefore, only those measurements made from June 18 through July 16 were used tc determine the effect of the drainage ditch on the water table. This was the wettest part of the growing season and is considered to be the most critical period affecting the structure and function of the plant community. Also, sampling of herbaceous species was accomplished during the last two weeks in Julyj and once the water level had exceeded the depth of the wells, it was well below effecti\ ditch depth anyway. During this wet period, June 18 to July 16, the mean relative elevation of the water table in wells J ,& m (Line 1) from the ditch (average of both sites) was 8 cm 0 CJ Site X 3 20 H 300 S 0) 2 8 0 *-1 F -0M) 260 h (U P* 240 4> > W +> 220 ■a (—< 41 D5 200 Ditch Level s Cl 280 Site 2 r-t 41 2 60 > a 240 4) 4) P* 220 41 > •H 200 t., Ditch Level 4 5 .7 m (0 0) 180 91.4m 6 0 tJ 4 2 (U 6/18 JX JL 7/2 n ' 7?!^ 7/30 ' 8/13 Date of Measurement 8 / 27 " 9$/ 1 0 FIGURE 12. WEEKLY RELATIVE WATER LEVEL WITH DISTANCE FROM DRAINAGE DITCH AND AMOUNT OF PRECIPITATION DURING 19^9 GROWING SEASON. 56 lower than in wells 45*7 m (Line 2) from the ditch and lO cm lower than in wells 91.4 m (Line 3) from the ditch (Table 15 These differences were significant (^<=.01). However, if relative water table elevations are transformed to actual depths below the soil surface, differences between mean depths on lines were not significant (^^.lO). Therefore, differential drawdown of the water table between 7*6 m and 91.4 m from the drainage ditch is less than can accurately be measured b y the methods used. The significant increase in relative elevation from the ditch results in a natural water table gradient corresponding to the sloping soil surface. Other studies have reported that unless a second ditch is used to intercept runoff water from surrounding areas, the effect of the first ditch will be greatly reduced (LeBarron and Neetzel, 1942; Averell and McGrew, 1929)* This appears to be the situation here since this single ditch drains an extensive area of lowland and upland. Huikari (Black, 1 9 6 7 ) found that ditch spacing is more important than ditch depth in significantly lowering the water table in peat soils. He believes 5 to 10 m to be the m a x i m u m effective distance between ditches and 70 cm to be an effective depth. Burke (1 9 6 7 ) reports maximum spacing of 12 to 15 feet for effective drawdown. Optimum depth of the water table to achieve maximum wood production varies from 20 to 50 cm, depending on the tree species concerned (Huikari, 1 9 6 5 ; Black, 1 9 6 7 )- Based 57 on th } s information and the rooting depth of species found on the two sites, the present ditch is probably sufficient, but lateral ditches will be needed to effectively control the water table. Natural discharge and evapotranspiration rates were sufficient to mai nt ai n a lowered water table except during periods of intensive precipitation. Therefore, drainage ditches will be useful only in removing excess water during these wet periods. TABLE 15. DIFFERENCES IN ABIOTIC SITE FACTORS AND STANDARD DEVIATIONS W I TH D I ST AN CE FROM DRAINAGE DITCH, JUNE-JULY, 1969 Abiotic Factors Relative Water Levelsa (m) Depth of Water Table3 (cm) distance from Ditch 91. 4m 7. 6m 46.7m Mean Mean Mean Sites Sites Sites 1 & 2 1 & 2 1 & 2 2 .6lq +0.23 7.9 +15.8 Mean Site 1 Mean Site 2 2 .7199 2 .6 gqq +0.21 ±0.25 6.2 + 11.2 7.3 +14.4 2 .8q ±7.5 11.5qq +17-1 Organic Matter Depthb (cm) 24. 8 s ±3.7 25.9 +4.5 2 8 .i SG +4.8 27.0 ±5-0 25*6 ±3 .9 pH of Ground Waterc 6.3S +0.3 6 .lss +0.2 6 .l ss ±0. 3 6 ,4q + 0.2 6 ,Oqq + 0.2 Oxygen In Ground W a t e r c (mg/1) 2.0 +0.8 1.7 +0.7 2.2 + 0.8 1.9 +0.8 2.0 +0.8 a Average of wells from June 18 to July 1 6 , 1969 b Includes Aj soil horizon c Measured In upper 15 cm of ground water Significantly different at .01 level s *ss Significantly different at .10 level 58 Hates of recharge and discharge varied with distance from the ditch (Table 16) . The greatest difference in rates of discharge between Lines 1 and 3 on Site 1, based on continuous water level recorder informationj when the water level was above the soil surface. occurred This difference can also be seen in Figure 12 from June 27 to July 2. Hates of discharge tended to equalize as the water level dropped below the soil surface. This differ­ ential drainage of surface water may be quite important to the shallow-rooted species vof this community. The quantity of precipitation occurring in June* 1 9 69* was 2.94 inches greater than the 20-year norm (Table 17)* July and September were drier, and August was approximately the same as the norm. From April through September of 19^9? there was a 0 .7 6 -inch deficit from the norm. This probably indicates water levels on the sites were somewhat lower than average during July, August and September. Mean depth of organic matter (includes A^ horizon) Line 1 was 24.8 cm (Table 15)* on Depths on Lines 2 and 3 averaged 25.9 and 28.1 cm, respectively. were significantly different (°C=.10). Lines 1 and 3 However, there was also a significant line-site interaction (°< =.0 5 ) due to an increase in depth with distance from the ditch on Site 1 compared to a slight decrease on Site 2. The slight decrease in depth on Site 2 may be the result of its better drainage properties. Also, Line 3 of this site drains as well as Line 1, while Line 2 drains more slowly. TABLE 16. RATES OF RECHARGE A!® DISCHARGE FROM HELLS CL SITE 1 DETERMINED FROM CONTINUOUS WATER LEI'LL RECORDER CHARTS 7. 6m Date (1969) 45.7m 91.4m Precipitation --- Recharge (cm/hr) (cm) _a Distance from Ditch 7.6m 45.7m 91.4m Initial Discharge — (cm/hrT T T 0.7 T T _ T T - - T T 0.3 0.2 0.2 0.2 - 0.4 3.4 0.0 - - - July 9 - 16 0.0 - - August 6 - 1 3 1.0 5-0 - 1.2 5.6 June 25 - July 2 5*8 0.5 July 2 - 9 a No data b <. 05 cm/hr 4.6 _ 45.7m 91.4m Extended Discharge — (cm/hr) q*Q 1.6 June 18 - 24 7.6m 0.1 6o TABLE 17MEAN MONTHLY PRECIPITATION AND DEVIATION FROM 20-YEAR NORM AT T H E DUNBAR FOREST EXPERIMENT STATION Precipitation (I n c h e s ) 20-Year Average61 1969k 1942-1961 Deviation From Norm (I n c he s) April 2.33 2.87 +0.54 May 2.96 1 .64 -1. 32 June 3-40 6.34 +2.94 July H 00 • CM 1.06 -1-75 August 2.86 3.38 +0.52 September 3.88 2 .19 -1.69 18.24 17-48 - O .76 Total a From U. S. Climatological Summary b Measurements made at Dunbar Headquarters Mean pH of the upper 15 cm of the ground -water was 6.3 on Line 1 and 6.1 on Lines 2 and 3 (Table 15). was significantly different Oxygen content (mg/l) (^=.10) from Lines 2 and 3 . in the upper 15 cm ol* the ground water was not significantly different distance Line 1 (** = . 1 0 ) with from the ditch. Differences in organic matter depth and pH with distance from the ditch indicate rates of decomposition and mineralization are greater on Line 1. Gardiner (1 9 6 6 ) found that drainage creates a more aerobic environment* thereby stimulating the growth of decomposing bacteria 61 which results in increased rates or d e c o m p o s i t i o n . 11' this is true, in the present study the increased rate of drawdown of surface water on Line 1 is important to the functioning of the system. There may also be increased light intensities on Line 1 during the morning hours on Site 1 and the evening hours on Site 2 due to construction of the ditch. This difference was partially reduced by locating the first line 7*6 m from the ditch. Greater light intensity would increase surface temperatures and cause increased activity of decomposing organisms. These factors were not measured, but do present questions to be answered in a later study. Although differences great, in abiotic factors were not it must be remembered that the ditch has been fully effective for only 6 years (ditch was completed in late summer of 1 9 6 3 ), and its effect on the water table has bee] small. Biotic Factors Changes in edaphic factors should result in changes in the plant community. These changes were not believed to be great enough or of long enough duration to have caused changes in the structure of the woody species. Therefore, only herbaceous species were analyzed for changes in structure with distance from the ditch. Other studies have shown changes in the composition of herbaceous species after drainage. Holmen (1964) 62 found a decrease in total number of species on bogs and fens drained for 3 3 y e a r s . He believed this resulted from increased eutrophy of the environment., giving those species with greater nutrient demands a competitive advantage. Satterlund and Graham (1957) found little change in species composition except along the ditch banks. Results from the present study indicate there has been very little change in species composition due to drainage, but there has been a change in distribution of individuals among the species (Table 18). Impatiens capensis was much more abundant on Line 1 than on Lines 2 and 3* Decrease of this species with distance from the ditch was m uch greater on the more poorly drained Site 1^ indicating it grows best on better drained sites, or is responding to some other condition associated w i t h drainage. Increased importance of Impatiens near the ditch results in a significant decrease (<=<=.0 1 ) of total individuals from Line 1 to Lines 2 and 3 mean number of individuals Line 1 was 109 per greater than on The number of (Table 19). (all herbaceous species) The on m 2 greater than on Line 2, and 112 per Line 3 * species per m 2 (quadrat diversity) not significantly different between lines. was Quadrat diversity averaged 12 per m 2 for both sites. Species diversity per individual based on the factorial expression of the Shannon-Weaver function may provide more insight about the species relationships than 63 TABLE 18. RELATIVE ABOVE-GROUND DRY WEIGHT AND RELATIVE DENSITY FOR 12 MOST IMPORTANT HERBACEOUS SPECIES WIT H DISTANCE FROM D I T C H ON EA C H SITE _________Distance from Ditch_________ 7.6m 46.7m 0 1 .4m 7 . 6m 4 6 . 7m 01.4m Relative Relative Herbaceous Species Current Growth Density ____________________________ Dry Weight _____________ (%)_______ Aster laterlflorus 2.0 Site 1 — Site 2 A. puniceus Site 1 1-7 Site 2 1.8 A. umbellatus Site 1 1-9 Site 2 Calamagrostis canadensis Site 1 12.0 — Site 2 Caltha palustris Site 1 0.9 Site 2 2.4 Dryopteris spinulosa Site 1 3.0 Site 2 1.6 Equisetum arvense Site 1 2.7 Site 2 0.1 Eupatorium purpureum Site 1 2.5 Site 2 0.9 Glyceria striata Site 1 22.0 Site 2 54.4 Impatiens capensis Site 1 7.9 Site 2 24.2 Senecio aureus Site 1 8.1 Site 2 1.0 Solidago rugosa Site 1 7.5 Site 2 3-1 Other herbaceous species Site 1 27.8 Site 2 10.5 cl rpy'o n o 3*9 1.4 0.6 4.8 1.3 2.6 0.1 1.5 1.6 5-5 0.5 1.0 0.3 1.1 0.3 2 -2 0.8 4.1 2.2 0.2 0.8 0.4 1.4 0.4 1.2 0.5 0.6 2.3 5.5 1.1 3-6 0.8 1.2 5.3 0.6 10.8 2.0 T 1.4 0.6 0.8 1.2 1.2 o.e 5.0 5*0 0.9 3.6 0.8 1 ’I 3*4 3*5 2,1 7.0 0.2 6.1 0.4 4.1 0.3 11.4 0.7 10.1 1.3 2.1 0.4 1.3 3*6 1.3 0.1 2.2 0.4 l.£ 0 .j 40.3 48.9 53.0 38.0 2.0 4.8 9.0 6.6 1 2 .f 8 .f 1.5 10.9 1.0 10.1 59*0 83.0 15*5 56.2 7*: 54. * 6.0 4.6 3-5 0.6 3*8 1.2 11.5 4.1 1 0 .: o.< 7*7 12.1 3*6 5*4 3.2 3*0 5*9 5.1 2.1 3*' 9*5 8.4 18.8 34.5 19*5 5*7 31.8 19.0 (jiS 0.8 — — — 0.1 i.t 42. 15* 64 TABLE 19 MEANS AND STANDARD DEVIATIONS OF BIOTIC FACTORS FOR BOTH SITES WITH DISTANCE FROM DRAINAGE DITCH, 1969 Biotic Factors Distance from Ditch 91. 4m 7 .6m 46.7m Mean Mean Mean Sites Sites Sites 1 & 2 1 & 2 1 & 2 Density of Herbs (no/m2 ) 221r ±90 11 ±5 Quadrat Diversity of Herbs (no/m2 ) 112rr ±75 13 ±3 109rr +55 13 ±3 Species Diversity Per Individual of Herbsa Individual Stems 1 .4 2 q +0.81 2.22qq +0 . 5 5 2. I3qq +O.63 Biomass Units 2 .0 5 j-0.82 2 .1 4 +0 . 7 7 2 .0 5 +O.58 Individual Stems o.8oq +0.13 0.66qq +0.10 0.66qq po.16 Biomass Units 0.66r ±0.19 0 .7 7 rr ±0.13 0.72rr +0 . 1 5 113s +61 7 0 s3 +37 Redundancy of Herbsa Biomass of Herbs (dry g/m2 ) a Based on factorial function. 9^99 Significantly **,rr Significantly s,ss Significantly 82 +72 expression of Shannon-VJeaver different at .01 level different at .05 level different at .10 level 65 quadrat diversity since it also depends on the distribution of individuals among species. Values vary viith number of species and distribution of individuals among species and Ghelardi, 1964-), (Lloyc Species diversity indices have been used successfully in aquatic environments to measure species reaction to environmental change along environmental gradients (Wilhm, 1 9 6 6 * 1 9 6 7 ) and (Sheldon, 1968; Gibson, 1966) Monk (1 967 ) has used diversity indices to characterize terrestrial vegetation. There are at least tvjo problems in using these indices in terrestrial plant communities: (1 ) determining what constitutes an individual of a given species, and (2 ) size of individuals for different species is not accounted for. For example, one individual of Impatiens, a single-stemmed annual, is not equivalent functionally to an individual of Glyceria, a clumping grass. The first problem Is partially solved vJhen these indices are used only to compare changes in the same community, assuming a given species is measured the same each time it is encountered. The second problem is more difficult, but it seems that biomass units for each species could be used to partially correct for inequality of size. Dicfcman (1968) discusses the use of relative biomass units In aquatic systems. It should be noted that the use of the Shannon-VJeaver function with biomass units may be mathematically incorrect. This question arises because each biomass unit for a given 66 individual m ay not have an equal probability of being selected. This problem is still subject to debate, however. Size or the surface area included per sample may also result In differences in the value of the indices. This results because increased number of rare species are sampled on larger sample areas. Wilhm (1 9 6 8 ) indicates these differences to be small, however. In the present study, both the number of individuals and dry biomass units were used to calculate diversity per individual and redundancy for herbaceous species on each m 2 quadrat. tically test and sites These values were then used to statis­ for differences in structure between lines (Table 19)* Diversity per individual based on d ensity was lower on Line 1 (*=<=.0 1 ) than on Lines 2 and 3* However, when diversity per Individual based on dry biomass units is used, differences betv;een lines are non-significant. Redundancy based on density significantly from Line 1 to Lines 2 and units it significantly (°C=.0 1 ) decreases 3# hut when based on dry biomass (tX = . 0 5 ) Increases from Line 1 to Lines 2 and 3These differences can be explained from the relative dry weight and relative density of the two dominant species (Table 18). Impatiens capensis constituted a much higher percentage of the total individuals sampled on Line 1 than on Lines 2 and 3* while more of the relative biomass belongs to Glyceria striata on Lines 2 and 3 than 67 on i.ine 1. However* the distribution of biomass among spucaos is not. so different that it results in a signifi­ cant change in species diversity per individual. capensis is a very succulent* Impatiens single-stemmed annual* while Glyceria striata is a comparatively low-raoisture perennial grass which forms clumps. This results in a large dry weight per individual for Glyceria and small dry weight per individual for I m p a t i e n s . The question now becomes w h i ch index best describes the relationship of species. Small dry weight of Impatiens indicates it is utilizing a comparatively small amount of the available nutrient source* while Glyceria is utilizing a much greater quantity. However* when these species are observed in the field* Impatiens towers over Glyceria * possibly cutting off some of the available light. This competitive relationship may be partially responsible for the Increased dry biomass per m^ for Glyceria with distance from the ditch on Site 1. On Site 2* Impatiens remains relatively important on all three lines* and dry biomass for Glyceria remains approximately the same. It appears that the effect of the drainage ditch on the structure of the understory lies somewhere between these two indices of diversity. The significantly lower diversity per individual and higher redundancy based on numbers of Individuals on Line 1* and the non-significant difference in number of species between lines indicate there has been a shift in number of Individuals among 68 species. However* little change in species composition and the non-signifieant change in diversity per individual based on biomass units indicate there has not been a significant change in biomass among species. Mean dry weight for herbaceous species on Line 1 was ^3 g/m2 greater than on Line 2 and 31 g/m2 greater than on Line 3 (Table 19). was significant The difference between Lines 1 and 2 (°C = .10). The lack: of significance between Lines 1 and 3 may be due to the better drainage c h a r ac te r ­ istics of Line 3 on Site 2* resulting in an increased dry weight. It should be noted that the high dry weight on the line is primarily the result of one quadrat located in a natural opening within the community (appendix Table 30). The drainage ditch may* therefore* be at least partially responsible for more dry weight being produced per m 2 per year. However* this effect may be indirect; i.e., it ma y have affected the overstory and in turn resulted in change in the understory. The effect extends to somewhere between 7 and 40 m from the ditch. Again* it should be stressed that differences in light intensity may also be responsible for this increased dry weight. Foliar analysis for percent nitrogen of the five dominant species with distance from the ditch lends further evidence to the possibility of increased nutrient availability on Line 1 (Table 20) . Values were greater on Line 1 for all analyses made except for those of 69 Fraxinus in 1968 and Glyceria in 1 9 6 9 . However, these differences were not as great as has been reported in other studies. TABLE 20. PERCENT DRY WEIGHT FOLIAR NITROGEN FOR FIVE DOMINANT SPECIES ON TWO SITES AND THREE DISTANCES FROM DRAINAGE D ITCH Speeiesa 7 •6m Site 2 Site 1 Distance from Ditch 91.4m 91.4m 7 •6m 45.7m Alnus rugosa 1968 1969 2.70 2.47 2.65 2.35 3.08 2.90 2. 30 Fraxinus nigra 1968 1969 2.26 2.49 2.26 2.26 2.34 2.18 2.45 — — - 2.34 2.28 Glyceria striata 1969 2.29 2.73 2.51 - - Impatiens capensis 1969 2.38 2.24 2.16 - - Cornus stolonifera 1968 a Samples collected in July , 1 968- 1969 Stanek (1 9 6 8 ) presents data showing a doubling of foliar nitrogen in black spruce after drainage. He attributes this to increased nutrient availability resulting from greater rates of decomposition. Increased decomposition was believed to result from increased oxygen supply due to increase in water movement by drainage. 70 Watt and Heinselman (1 9 6 5 ) indicated two possible reasons Tor increased foliar nitrogen content on better sites: (1) nitrogen fixation by A l n u s , and decomposition of organic matter. (2) more rapid They found active root nodules primarily in hummocks above the water table where the oxygen supply is above the minimum of 10 percent needed for nitrogen fixation. This could also be a factor in the present study since surface water does drain more quickly from bine 1 , thereby creating a better aerated environment in the h u m m o c k s . A growth ring analysis was made on Alnus to determine whether growth rates had increased since drainage (Table 21). Growth ring index values were computed to adjust ring widths for variability other than that due to climate (Fritts, 196 6 ; Clock, 1955). The larger the index value, the greater the growth for that year due to climate. Index values were not significantly different between lines. They were significantly different (°< =.05) between the years i960 and 1 9 6 2 , both before completion of the drainage ditch. other studies have found response of tree species to drainage varies from 1 to 8 years (Gatterlund and Graham, 1957; Averell and MeGrew, 1929)- Poor growth response m ay be due to this lag effect or may be the result of an ineffective drainage ditch. 71 TABLE 21. MI-1 ,AN ClHOW TII HI MO INDEX VALUESa OF SITES 1 AMD 2 FOR ALNUS RUGOSA AT THREE DISTANCES FROM D RAINAGE D I TC H Year Distance from Ditch 91.4m 7.6m 45*7 hi 1969 1.36b 1.32 1.26 1.31 1968 1.30 1.30 1.25 1.28 1967 1.26 1.22 1.28 1.25 1966 1.35 1.20 1.28 1.28 1965 1. 31 1.24 1.33 1.30 1964 1.22 1.31 1.28 1.27 1963 1.19 1.25 1.2 6 1.23 1962 1.17 1.21 1.14 1.17rr 1961 1.18 1.20 1.18 1.19 i960 1.34 1.23 1.69 1.42r Year Average 1.27 1.25 1.29 Average a Methods from Fritz* 19685 the larger the index value* the greater the ring width. b Mean of 20 stem discs wi t h 2 growth ring series er d i s c . r Significantly different at .05 level. ? CHAPTER V. SUMMARY The study area was located In a glacial basin with poor natural drainage. Hummocks and depressions charac­ terized the soil surface, but change in elevation over the study sites vjas very small. Soils consisted of a shallow layer of black, mucky organic matter underlain by a silty clay loam horizon on Site 1 and very fine sand horizon on Site 2. The A^ horizon was strongly leached and blue-gray in color. Thickness of the A2 plus B horizon was uniform, indicating little difference in permeability over the study sites. Nutrients are probably carried onto the sites by runoff water from the surrounding uplands. This results in a nearly neutral pH (slightly acid) of the ground water and enriched nutrient regime. Foliar nutrient percentages of the five dominant plant species In the community supply evidence to Its eutrophic character. Impermeable soils, physiographic location, and periods of excessive precipitation result in a high water table through the middle of the growing season. Greater permeability of the coarser-textured soils and smaller amount of incoming runoff water on Site 2 result 72 73 in better drainage than on Site 1. VJater table levels during the latter part of the growing season dropped below the plant rooting zone due to natural discharge and evapot ra n s p i r a t i o n . However, comparison of amounts of prec i pi ­ tation with the 2 0 -year norm indicate the first half of the growing season was wetter than normal, and the last half was drier than normal. These edaphic characteristics - high but flowing ground viater and enriched nutrient regime - result in a rich and characteristic flora. were identified on both sites. More than lOO species Gramineae was the most important family in number of genera, and Carex was the most important genus in number of species. The plant community (vascular species only) had three distinct strata. Alnus rugosa and Fraxinus nigra were dominants in the overstory; Cornus stolonifera and Ilex vertici11ata dominate intermediate shrubs; and the u n d e r ­ story was dominated by Glyceria striata and Impatiens capensis. Better drainage characteristics of Site 2 have allowed Alnus to maintain more dominant control than on the more poorly drained Site 1. This more uniform overstory plus the more rapid drainage of water from surface d e p r e s ­ sions result in a more redundant environment for the understory species on Site 2. This is indicated by the greater dominance of Glyceria and Impatiens and herbaceous species per unit area on Site 2. fewer 74 The mixed species overstory, greater importance of ITaxinus n i g r a , and greater numbers of sprouts and seedlings for Fraxinus and Populus balsamifera indicate Site 1 is further along into a transitional stage of succession from a shrub-carr to a swamp-hardwood community than is Site 2. These differences are probably due to the greater vigor of Alnus on the better drained Site 2 and to past history of such factors as fire, logging, and animal pests. However, evidence of fire, logging and pests indicates they have not been a strong factor in changing the structure of the community during the past 50 years. Biomass and production estimates for the dominant overstory species indicate Alnus is a strong dominant on Bite 2, while it is a co-dominant with Fraxinus on Bite 1. Small biomass accumulation ratios for these two species indicate their small stature. Estimates of community net production place it in the upper half of the range, (g/m^/year) greater than 600 g/m^/year, given for woodland and shrub communities; while biomass estimates fall in the lower half of its range, less O than 9,000 g/m , given for such communities. Herbaceous species are more important in this community than has been given for other shrub communities. Biomass accumulation ratios are considered lower than actual for the community on Site 1 since large, over-mature individuals of Populus and Larix laricina were not included in biomass and production estimates. 75 The efi'ect of the drainage ditch on edaphic factors and the structure and function of the plant community vjas minor. Differential drawdown of the water table did not vary significantly between 7 and 91 m from the drainage ditch. [fowoverj there is evidence that rate of discharge of surface water is greater near the ditch. Lateral ditches to stop the inflow of runoff water from surrounding areas are probably necessary to increase the effectiveness of the present ditch. Significant decreased depth of organic matter, increased pH of ground water, increased biomass of herbaceous species, and a slight increase in foliar nitrogen content for four of five dominant species tested indicate greater rates of decomposition and mineralization are occurring within 40 m of the ditch. This may be the result of changed hydrological characteristics, but it may also be partially due to increased light intensities resulting from overstory destruction during ditch c ons true t i o n . This has resulted in a slight change in structure or the herbaceous species. The drainage ditch has created a more favorable environment for Impatiens c a p e n s i s , allowing it to greatly expand its numbers. Diversity per individual and redundancy based on density Indicate this proliferation has resulted in significant change in equitability of the species. However, diversity per 76 individual and redundancy baued on the dry biomass value for each species, along with little difference in species composition between lines, indicate the change in structure has not been severe enough to cause a significant shift in biomass among the s p e c i e s . No significant difference was found in growth of Alnus with distance from the ditch based on a growth ring analysis of stem discs. Insufficient time has elapsed since construction of the drainage ditch, and/or its effect has not been great enough to cause a growth response in this species. BIBLIOGRAPHY BIBLIOGRAPHY Arkhipova, K . P. 1955* Thermal regime of soil on a reclaimed bog. Translated from Russian by Israel Program for Scientific Translations,, 1 9 6 8 . Available from U. S. Department of Commerce, Clearinghouse for Federal Scientific and Technical Information, Springfield, Virginia. Averell, James L. and Paul C. McGrew. 1 9 2 9 . The reaction of swamp forests to drainage in northern Minnesota. Department of Drainage and Vlaters, State of Minnesota. Bay, Roger R. 1 9 6 6 . Factors influencing soil-moisture relationships in undrained forested bogs. Inter­ national Symposium on Forest Hydrology. Pergamon Press, Oxford and New York.. 335-343* Bay, Roger R. 1 9 6 7 * Ground water and vegetation in two peat bogs in northern Minnesota. Ecology 48(2): 3 0 8 -3 1 0 . Black, T. 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Agricultural Experiment Station, Michigan State University, East Lansing, Michigan, pp. 138-140. Wilhm, Jerry L. and Troy C. Doris. 1 9 6 6 . Species diversity of benthic macroinvertebrates in a stream receiving domestic and oil refinery effluents. The American Midland Naturalist 7 6 (2 ): 427-449* Wilhm, Jerry L. 1 9 6 7 * Comparison of some diversity indices applied to populations of benthic m a c r o ­ invertebrates in a stream receiving organic wastes. Journal of Water Pollution Control Federation, Washington, D. C. 39(10): 1 6 7 3 -1 6 8 3 . Wilhm, Jerry L. 1 9 6 8 . Use of biomass units in Shannon's formula. Ecology 49: 153-156. Wright, J. O. 1907Swamp and overflowed lands in the United States. U. S. Department of Agriculture Circular Mo. 7 6 . APPENDIX S3 TABLE 22. RELATIVE ELEVATION OF PIPE LOCATED AT EACH W EL L AND ALONG CENTER OF DITCH W el l No. D i tch Relative Elevation (cm) 45.7m 7 ‘.6m 91.4m Cite 1 I 2 3 4 296.6 2 9 1 .1 285.6 298.4 296.9 292.6 290.8 291.1 303*9 299*9 3 0 4 .2 301.1 305.1 305.4 308.5 3 0 6 .6 281.9 275*8 262.7 2 6 0 .3 279*5 275*5 273-1 272.8 2 7 8 .9 279.5 273-4 272.2 280.1 276.8 2 7 2 .8 270.1 Cite 2 1 2 3 A TABLE 23. RELATIVE ELEVATION OF SOIL Sim FACE AT EACH WELL AMD ALONG CENTER OF DRAINAGE DITCH Well No. Ditch Relative Elevation (cm) /. 6m 45 *7m 91-4m Cite 1 1 2 3 4 239*3 249*3 245*7 246.0 283.5 286.2 276.8 282.9 295.1 2 8 9 .3 292.0 284.7 295*1 294.7 290.8 292.6 221.7 217*4 221.9 214.8 260.6 254.5 254.5 256.0 268.2 262.4 253.9 254.2 269.1 264.6 260.6 261.2 Cite 2 1 2 3 4 84 TABLE 24. AVERAGE DEPTH OP ORGANIC MATTER ON EACH QUADRAT AND AVERAGE DEPTH TO MINERAL DOII, TEXTIJRAL CHANGE AT EACH W EL L LOCATION (. bm Lite 1 Site 2 4 b .7m Site 1 Site 2 Organic Matter Depth 91 •4m Site 1 Site 2 (cm) Quadrat 1 25 .4 25*4 26.7 19.1 38-1 30.5 2 19-1 27*9 22.9 22.9 38.1 19.1 3 25 .4 31.8 22.9 33.0 33.0 27.9 4 27.9 26.7 25.4 27-9 31-8 22.9 9 19.1 2 6 .7 27.9 22. 9 30.5 27-9 6 20. 3 27-9 21.6 27.9 30.5 24.5 7 2 2 .9 27.9 31.8 24 .'5 30.5 27.9 Q 25.4 19.1 31.8 33-0 26.7 24.5 9 2 0 .3 24.1 27.9 22.9 25.4 24.5 29.2 24.1 26.7 17.8 24.1 24.5 23-5 26.2 26.6 25.2 30.9 25.4 10 Quadrat Average Site Average 24. 8 2 8 .2 25. 9 Mineral Horizon Thicknes □ (cm) Well 1 6l 69 58 51 71 51 2 71 51 71 51 51 51 3 56 51 6l 51 71 51 56 61 56 71 71 76 61 58 61 56 66 57 4 Vie11 Average Site Average 59 58 61 85 TABLE 25I'll IN THE UPPER 15 CM f)K GROUND WATER MEASURED AT EACH WEI,I, AND DRAINAGE DITCH, JUNE-JULY, 1969 Well Ho. Ditch ” Site Site 2 1 '7. bm Site Site 2 • 1 ■ 46. 7m Site Site 1 2 9 1 .4m Site Site 2 1 1 71 *"3 J 6.3 6.8 6.2 6.4 6.0 6.5 6.0 2 6.0 6.3 6 .2 6.3 6.4 5-7 6 .6 6.1 3 6.8 6.4 6.6 6.0 6.2 6.0 6.0 5-7 4 7-2 6.5 6.6 6.0 6.2 b.O 6.1 6,0 6.8 6.4 6.6 6.1 6 .3 6.0 6.3 6.0 Well Average ■Site Average 6>. 6 6 .2 6 .2 6 .4 TABLE 26 • OXYGEN (MG/L) IN UPPER 15 1CM OK GROUND WATER IN E A C H W E L L AND DRAINAGE DITCH, JULY, 1969 Dit ch Site Site 1 2 7- bm Site Site 1 2 2 7.8 7.0 3 8.2 4 2.1 1.2 1.4 1.4 1-5 3-7 1.2 1.6 1.7 2.4 8.2 1.6 1-5 1-3 3-5 1.8 8.2 8.2 1.2 1.3 2.0 3-3 3-3 1-5 8.0 7.9 1.7 2.2 1.6 1-9 £-5 1.9 7- 9 2 .0 1 -7 * Well Average Site Average • 7.1 91- 4m Site Site 2 1 CO H 7-8 45- 7m Site Site 2 1 CO • H 1 ro Well No. 2 .2 TABLE 27. WEEKLY FLUCTUATION IN THE WATER TABLE AND PRECIPITATION FOR EACH SITE DURING THE 1969 GROWING SEASON Date c? Measurement (19o9) June 18 Site Site June 25 Site Site June 27 Site Site July 2 Site Site July 9 Site Site July 16 Site Site July 23 Site Site July 30 Site Site Relative Elevation of Water Table ------ (it)- - - - - - - - 7.6m -5 •7m 91 ■4m Depth of Water Table Below Ground Surface Precipitation — - - --- (cm)- - - - - - - - - - - - (cm)7.6" 45 •7m 91*4n Ditch_ _ _ _ _ _ _ _ _ _ _ _ _ 1 2 2.77 2.46 2.86 2.49 2.92 2.51 5*7 9-9 4.8 10.6 1-5 12.6 -13.3 -15.1 2.1 2.1 1 2 2.76 2.43 2.85 2.46 2.90 2.49 6 ^ 13.0 4.8 13.0 3.7 14.5 -13*0 -13.8 1.2 1.2 1 2 2.90 2.71 2.97 2.70 3.03 2.69 -8.0 -14.3 -7.2 -10.4 -12.1 -4.7 -43.1 -45.8 5.8 5*7 1 2 2.31 2.56 2.91 2.60 2.95 2.62 1.1 0.9 -Co 0.1 -2.1 1.6 -19.9 -22.3 1.2 1.1 1 2 2.76 2.45 2.85 2.49 2.90 2.51 5.9 n.o 5.2 11.0 3.0 13.0 -14.6 -14.5 0.0 0.0 1 2 2.69 2.06 2.77 2.30 2.8? 2.19 13.7 30.1 13.2 29.9 11.6 45.1 -11.9 -11.9 0.0 0.0 1 2 2.41 1-93 2.53 1.93 2.72 1.93 in .■? 63.0 22.6 21.1 70.6 -9.2 -9.8 0.2 0.2 1 2 2.62 1.9c 2.70 2.06 2.74 1*93 20.5 60.2 IQ -L_/*0 18.9 70.4 -10.1 -10.9 3*1 3.2 000 5^.° TABLE 27 (cont'a) Date of Measurement (1969) August 6 Site 1 Site 2 Ausust 13 Site 1 Site 2 August 20 Site 1 Site 2 August 27 Site 1 Site 2 September 3 Site i Site 2 September 10 Site 1 Site 2 September 17 Site 1 Site 2 'Seek Average V . t _ r w Site 2 Sire Average Relative Elevat1cn of Eater 2'able „ \(m)j 91.4m 7 .6m 45.7m Depth c:‘ Eater '.able Below Ground Surfa .. . . . . . . . f f m V _ _ . * 1?r»o 7 .6m 45.7m 91.4m Lrwii 2.69 2.34 2.70 2.31 2.SO 2.12 13.6 22.3 14.4 28.2 1^ ^ 51.5 -11.0 -12.4 3.1 3.4 2.58 1.94 2.68 1.93 2.72 1.93 24.0 62.8 22.6 00.2r-j 21.2 70.4 -9.7 -9.8 0.9 1.2 2.21 1.93 2.30 1-93 2.35 1.93 74.5 63.2 54.6 66.4 58.8 70.4 0.0 r* — 0.0 0.0 2.20 1-93 2.20 1.93 2.20 1.93 75.0 63.2 74.7 66.4 74.3 70.4 0.0 0.0 0.0 0.0 2.29 1.93 2.59 1.93 2.64 1-93 54.6 63.2 31.6 66.4 29.1 70.4 -8.1 -8.5 8.9 5.8 2.20 1.93 2.26 1.93 2.42 1.93 73.4 63.2 65.2 66.4 51.0 70.4 -5.2 -4.8 0.6 0.7 2.20 1.93 2.26 1*93 2.36 1.93 75.0 63.2 66.3 66.4 57.9 70.4 -8.9 -8.5 1.3 1.4 2.54 2.17 2.65 2.20 2.70 2.17 28.3 39.6 25o 40.1 2” .^ 46.5 -11.9 -12.7 1.7 1*7 ,"1 2.42 0 liii 34.0 32.8 34.9 -12.3 1.7 — -.02 Wl a * —\ • * Frecipite /pm 'rl V'-m TABLE 23. CURREHT GROWTH DRY WEIGHT,, NUMBER OF STEMS, AMD PERCENT OF QUADRAT OCCURRENCE OF SPECIES ON TWO SITES AMD THREE DISTAMCES FROM DRAINAGE DITCH Species Herbaceous Species Agrostis perennans Site 1 Site 2 Arisaema atrorubens Site 1 Site 2 Aster lateriflorus Site 1 Site 2 A. nuniceus Site 1 Site 2 A. uir.be11atus Site 1 Site 2 Bromus ciliatus Site 1 Site 2 Calamagrcstis canadensis Site 1 Site 2 Caltha ualustris Site 1 Site 2 Current Growth Dry Height / ~/ ±U.*;- ) 91.4m 7,6m 45.7m - - 1.5 0.1 - - Humber cf Stems 7. 5m 45 •7m 91 illT 0.8 - - - 0 - - - tpa 3 3 1 10 20 - - - 26.8 10.0 4.3 T 14 18.6 21,4 10.7 11.0 39.9 4.7 20.5 - 28.6 15.4 ii.•li. - 4.3 - 128.5 - t*3 lo.o 40.3 9-o 74.3 0.4 13-9 12.9 21.5 - 28.2. Frequency of Oc currenee fcf. { vj . _ . 7.6m 91.4n 45.7m p r - - 54 15 28 1 30 19 8 12 3 31 0J 1.8 6.9 8 - 16 -iir - 7 - /■i* - 11 20 9.6 - 00 - - 4o 20 10 - 30 30 10 70 50 50 30 70 50 13 0 20 - 40 30 40 40 10 - - 10 - 30 - - 9 55 6o 13 7 - 20 30 50 10 14 1R 1 30 70 30 10 40 30 9 - TA3LE 28 (cont'a) Current Grcxth Dry Height Species (g/10m2)----_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 7 .6m 45 *7m 91 ■4m Herbaceous Species Cardamine censylvanica Site 1 50 Site 2 Carex canescens Site 1 0.1 0.6 7.5 Site 2 3.6 C. crinita Site 1 Site 2 4.6 C. disperma Site 1 0.7 Site 2 C. gracillima Site 1 13.1 1.4 Site 2 C. hystricina Site 1 Site 2 0,7 C. interior Site 1 0.1 7-7 Site 2 C. inclines cens Site 1 4.2 6.0 Site 2 8.1 0. leptalea Site 1 1.6 3.7 7.3 Site 2 0.3 7.6 1.7 Number o£ Stems (10m-)- - - - - - 7 ■6m 45* 7m 9 1 .4m Frequency of Occurrence (Or)7 •5m 45. 7m 91.4m 1 - - 1 - 1 1 12 - 10 - 10 10 30 - - 1 - - 10 - - 3 - 2 - 4 10 - - 1 1 - 10 - - 10 - 10 10 10 - 1 - 2 3 - 10 - 10 20 21 2 37 47 65 12 30 20 60 70 10 50 100 30 co ^ TABLE 28 (cent1a) Snecies Herbaceous Species Carex leptonervia Site 1 Site 2 C. projecta Site 1 Site 2 C. stipata Site 1 Site 2 C. tribuloiaes Site 1 Site 2 C. Tucfcermani Site 1 Site 2 C. vesicaria Site 1 Site 2 C. spn. Site 1 Site 2 Chelone glabra Site 1 Site 2 Cicuta bulbifera Site 1 Site 2 Current Growth Dry height ■(g/lOni-) 7.6m 45-7n 91 •4m Frequency 03 Occurrence Number of Stems -(10m2)- 7 *oni 45.7m 91 • 4m - 1.4 0.1 15.3 3.1 5.1 9.2 0.8 8 1 1 32.2 2 5 17 3.9 30.6 9o 0.8 2 2 4 1 14.0 5.0 14.1 3 7 .6m 10 20 10 10 20 30 40 10 10 30 10 10 40 50 10 10 10 10 20 20 47.7 14 20 145.1 - 11 20 0.7 2.3 3*3 2.0 0.9 1.2 - 11 30 14 20 8 5.8 15.7 2 1 10 n o 91'. 4m 20 4 5 0.3 - (?0 - 45. fir. 1 10 30 60 20 \Q O TABLE 28 (cent’d) Current Growth Dry Weight Species (g/lOm*)----_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 7.6m 45»7m 91.4m Herbaceous Species Cinna arunainacea Site 1 1.9 Site 2 5-7 C. latifolia Site 1 12.3 3.4 0.3 Site 2 48.1 9.6 12.0 Dryonteris spinulosa Site 1 31*3 34.6 6.8 Site 2 I8.9 35.2 33.O Epilobium glandulosum Site 1 4.7 0.3 Site 2 2.9 4.0 2.2 Eauisetum arvense *Site 1 29.2 48.7 44.3 Site 2 1.1 1.5 3.8 Eupatorium maculaturn Site 1 0.1 75-3 Site 2 196.7 E. purpureum Site 1 26.6 14.5 9.2 Site 2 10.7 3.2 32.7 Eragaria vir^iniana Site 1 0.8 4.1 Site 2 1.0 Galium palustre Site 1 - 1 , 6 Site 2 - Ereauenoy c:‘ Number of Stems Occurrence (10m2)- - - - - - - - - - (-1)7 .6m 45 *7m 9 1 -4m 7.6m 45.7m 51.4m 2 - - - 1 0 2 - - 10 17 31 15 18 2 40 50 90 20 90 10 60 31 22 38 39 17 24 70 40 80 90 40 70 25 18 2 14 2 70 30 10 30 10 75 7 128 8 110 12 80 40 80 40 90 50 - 1 - 14 2 24 2 25 4 13 8 3 - 6 - 2 10 - 10 - - - 5 - - 10 40 10 40 10 50 30 50' 40 10 1 0 M3 M TABLE 28 (cont'd) Current Grcv.'th Dry Height Species Herbaceous Snecies Galium tinctcrium Site 1 Site 2 G. triflorum Site 1 Site 2 Glyceria striata Site 1 Site 2 Habenaria psycoaes Site 1 Site 2 Hieracium aurantiacum Site 1 Site 2 Inmatiens cauensis Site 1 Site 2 Iris versicolor Site 1 Site 2 Laetuca biennis Site 1 Site 2 Iye opus uni flcrus Site 1 Site 2 Humber cf S terns t1r\rr? \ 7 .6m 91.4m 45. 7m Frequency 0: Occurrence let\. 7 .6m 45.7m 91.4n _________ .... 7 .6m 45.7m 91.4m 2.0 5.8 0.9 3.4 0.6 7.7 44 31 45 35 31 44 90 80 60 50 50 40 3.3 3.4 4.0 3.8 0.1 1.6 19 16 59 18 2 21 30 10 50 70 10 50 235.6 6/18.9 278.3 347.2 386.6 345.2 36 125 102 74 133 99 70 100 90 100 100 90 2.6 - 1 1 3 - - - 10 - 10 — 30 — - - - - * - 10 - 1 - 1.076 2,163 175 631 79 616 100 100 100 100 80 90 1 30 — - 10 30 10 20 10 30 40 4o 40 70 40 0.3 T - - - - - - 85.I 288.8 11.3 - 10.2 77.6 - T 7.3 91.9 - 4.9 1.1 1.8 0.8 T 3.1 3.5 3.1 0.8 4.9 3.9 - - 14 - - - 9 1 7 1 40 73 10 - - 3 117 57 - - - - TABLB 28 (cont'd) Current Growth Dry weight Species (g/lOm*)----_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 7.6m 45.7m 91«4m Herbaceous Species Maianthemum canaaense Site 1 Site 2 Mite11a nuaa 4.8 Site 1 3.9 Site 2 1.2 Molinia caerulea Site 1 2.3 Site 2 Onoclea sensibilis Site 1 Site 2 3.9 1-9 Ranunculus abortivus Site 1 Site 2 0.2 0.1 R. recurvatus Site 1 0.7 Site 2 Riunex orbiculatus Site 1 0.6 0.7 1.3 Site 2 Scutellaria lateriflora Site 1 0.2 0.7 0.5 Site 2 2.0 1.2 S.5 Senecic aureus 87.2 Site 1 4lo 25.7 Site 2 32.4 12.0 5.4 - 1— 1 - - - 1 - - - - - - - - - Number of Stems (lOir.2)- - - - - — 7*om 45 ■ 7m 91* 4m Frequency of Occurrence - - - - - - - [%)- - - - - — 7-6m 45.7m 91.4m - 4 3 35 - - - 20 10 10 - 50 - 10 5 - - - - — - - — — - - - - - - - - 2 3 - - - - - - 2 2 - 10 - - 8 - - 10 - - - - 2 1 1 - - - - - - - - - - - - - - - - - - 54 79 17 20 69 130 46 10 20 - — * -* - - - 10 — - 10 - 10 — 20 — - 10 - 19 40 50 30 60 60 30 107' 10 60 50 70 50 80 40 0 TABLE 28 (cont'd) 4 Species Herbaceous Species Smilacina trifolia Site 1 Site 2 Solidago rugosa Site 1 Site 2 Stellaria graminea Site 1 Site 2 Viola cucullata Site 1 Site 2 Viola nailers Site 1 Site 2 Miscellaneous Site 1 Site 2 lotal Herbs Site 1 Site 2 Current C-rov:th Dry Height (g/10m2)----7 .6m 45.7m 91*4m - 1.3 - T 80.1 37.3 - 1.0 53.3 86.1 0.4 - 0.1 2.5 1.9 26.2 49.2 : Number of Stems (10m2 )-----7 .6m 45.7m 91.4m - - 59 30 17 x 00 57 37 19 30 38 - 60 50 - - 2 - - - - - - - 4 3 - - - - - - 0.4 2.3 0.2 2.2 11 1.4 1.6 27 0.2 T 0.2 T - 0*1 7 2 716.3 910.1 1,841 2,609 693.0 711.6 - - - 1,072.7 1,193.4 Frequency of Occurrence (£)■ 7 -6m 45.7m 91*4m 20 - 4o 10 40 30 70 100 50 60 10 — 10 - - — — 13 31 Ill 26 30 40 20 60 70 60 14 3 3 60 20 30 20 30 - 1,169 1,125 1,059 1,134 - TABLE 28 (ccnt’d) _______ 7«8m '45.7rc 14.2 20.4 1.5 34.1 8 - 10.3 - 0.1 rp 7.4 - 1 - 2 8 - - - 0.4 1.0 1 - 1 3 1.2 - 1 - 3 - - 9.4 - - 41.4 9o ii. - 2.4 2 - - o~6 T 21.5 - 91.4m 20 17 - - - 1.7 - - 0.1 49.? 0* 'O0' 1 Low Shrub L Vine Species (rcv;t;n Dry Be Lent number or Stems (g/lGm.2)----(10m2)-----7.6m 45.7m 91* 4m 7*6m 45.7m 91.4m 0.6 - 17.0 - 11.0 - rrequency c: Occurrence (;"*)• 7.6m 45.7m 91*4m 1 - I - 1 - 10 - 10 - 10 - - ry* - - - 1 - - - 10 - - - 4.5 - 0.1 - 4 - 2 - 10 - 10 - 0.1 - - - 1 - - - 10 - - - rr - - 1 - - 10 - 11.0 0.1 1 - 7 1 1 2 325.3 367*9 344.1 1,110.5 1,937 2,o5o 1*358 1,299 1,221 1,320 0.6 1*159*3 l s25o.5 21.6 11 * 'J8 t a u t .r ay. ,i s t o k s p e c i e s s e e n o n s t u d y s i t e s BUT NOT FOUND ON SAMPLE QUADRATS Herbaceous Species *Alopecurus aequalis Carex paupercula C. rostrata C . tenuiflora *Cirsiura discolor Ooptis groenlandica Cornus canadensis Erigeron philadelphicus Nestuca obtusa Galeopsis Tehrahrt Galium Aparine xGeum macrophyllum G. virginianum Glyceria canadensis G. grandis G. obtusa Lysimachia thyrisflora ~*Phragmites communis Schizachne purpurascens Taraxacum officinale ^Veronica americana High .Shrub Species Fslix discolor Sambuous pubens Spiraea alba Tree Species Acer rubrum A. saccharum Betula lutea Picea glauca P. mariana * Species seen only along the ditch 99 TABLE 30. DRY WEIGHT, DENSITY AND NUMBER OF HERBACEOUS SPECIES FOUND ON E ACH 1 M 2 QUADRAT Q.uadrat Number 1 2 3 4 5 6 7 8 9 lo Quad . Avg. Site Avg. 1 2 3 4 9 t; 7 O 9 io Quad . A v g . Si te Avg. 1 2 3 4 5 6 7 8 9 10 Q.uad . Avg. Si te Avg. 7. bm Site 3T Site 2 91.4m Site” 1 Site ? 243 201 96 82 123 80 102 148 91 66 119 94 47 23 40 23 37 79 81 59 249 73 113 Dry Weight (g/m2 ) 13 103 58 83 121 125 97 63 33 149 38 73 87 65 63 37 81 7 42 36 69 71 70 221 368 339 179 210 199 293 244 299 289 249 261 Density (no/m2 ) 96 239 260 163 98 66 127 31 130 227 41 49 211 171 101 4l 42 34 61 46 112 113 112 86 99 99 114 83 10s 170 87 112 108 106 99 188 106 39 224 91 76 191 69 HO 107 96 103 200 32 321 321 276 174 11/ 183 182 ^ 45. 7m Site IT Site ^ 12 9 13 6 13 11 19 22 15 23 14 11 9 7 10 10 11 9 7 10 7 13 9 Species 11 19 9 13 17 10 18 13 9 10 13 (no/in2 ) 19 14 15 11 16 12 13 12 9 11 13 13 82 33 153 17 98 289 78 86 61 29 14 91 133 118 43 32 96 95 280 115 132 31 113 109 8 12 8 8 17 17 11 11 13 10 12 19 16 14 14 13 19 9 9 14 19 14 13 100 TABLE 31• SPECIES DIVERSITY PER INDIVIDUAL AND REDUNDANCY FOR NUMBERS OF INDIVIDUALS OF HERBACEOUS SPECIES PER 1 m 2 QUADRAT Quadrat Number 7. bm Site r Site 2 A5. 7m' Site 1 Site 2” 91.4m Si'be I Sit 4.5 4.5 4.0 3.0 192 I .183 1,938 863 2.712 '464 17 152 231 73 225 62 315 1,178 516 1,530 316 308 168 126 590 93 31 26 26 37 15 151 109 97 217 64 ■iyOT Seem No. Stems Total One -Year3 Total Dry Neight Volume Yc lume Ncod _ N ood Food *r Bar’s t ■■ { } 1 ■■ IsJ TABLB 35 (cont'd) Stem He. Height Cm) 1.8 4.0 5.1 5.0 1.7 5.1 8.8 4.0 2.0 4.5 5.0 5.0 2.5 5-0 5-5 4.5 354 2,983 4.666 4;05S 445 5,241 14,011 3,047 Id 300 584 595 57 774 1,206 318 251 1,575 2,274 2,381 2.5 4.0 2.4 3.1 4.0 5.0 4.0 4.0 2.5 4.5 5.0 968 3,467 982 1,652 648 3,418 5,730 106 371 107 74 77 228 509 5H 1,599 562 823 385 1,747 2,931 Sice 2 7.6m 1 2 3 4 5 6 7 8 45. 7m 1 2 3 4 5 0 7 91. 4m 2.2 4.2 5.4 Total Dry Height Hood 4 Bark (b ) 25 0 2,806 8,332 1,493 Total Dry Height Heed - 3ark Branches One-Year0 Dry Height Dry Height Twigs 4 Heed 4 Bark Leaves. (5 ) 536 1,381 1,098 49 1,060 4,115 710 110 5o2 35 328 112 511 1,297 345 230 33 36 3*5 264 2 914 5*0 2,595 439 3-8 p 5.2 5,79o 963 0.5 593 3,179 389 2.2 114 820 3*5 87 n 4.0 228 822 282 3*0 4,783 ■ 0 5*813 240 1,216 0 *te 2,353 5o PQ 356 26 1.0 7 3*5 175 —> 5*0 2,324 0 376 367 3*7 1,697 a Mean of 5 years ’ grcv;tha 1904-■I906, based on Smalian formula (mean of of stem sec tier. X height) 1 1.0 17 o9 96 106 16 103 176 63 51 289 544 540 50 536 1,293 330 29 68 99 320 37 154 89 294 499 12 30 26 54 82 0s 30 62 271 483 93 106 32 h~ 183 *0 420 64 36 15 270 68 areas at each end 105 DBH (cm) Stems Total Cne-Y=ara Volume Volume Heed "cod I Clil-') io6 -tJ -Cl I IJ> c/1 mxt voco o m a c o roo\hO CM CM CO L— VO O i-l vo o'lm o me w r-i -i ovooo co ro in o o v o i o moo vo ai o OJ OlW H H O O CM CM xt O H O l H inmvo r— vo xi- un cm roi-ico orxj- o v o or Ln CO CM r-l CM VO m mvo vo —f m CM OT r—ICO C- IPcCO COOC\lfC)C 1-1C O • r lC / J C l ) d) hfi *t-l cl! vo r i L-(t :w 1) X . CO >j; hi Cl C) :c pt O X I.:: ■=£ •a; (r , ro rq l-i i —i R Ui „ EQ W Lc/-ij i{ i— i i— i i— i i— i i—i i—ir-i i —i Cl OT C / J o :n OT H x :t : o l-i co i —I0.1i—( ■U Jd .c; p, bl)ctJ ttl ■H (i) In cl) O :-: + L-i ■a X o Pi o Ci X vo o otcm in cr\i-i C- h- CM VO 1-1 OTxJVQ rHVO AJ CM xt*CO •1 *1 •! »l i —I i—IVO CM -n /d .Cl Sh r-l U ) aj CO *rl [A P r, n i— I at o o r— r~ Ii—Ii— I CM h"-r—ivo i— iin -=f COxrJ-co C O CM •» n •» n n n *i CO i-W H VQ OT i — IL— r— 1 (O r—I CO h- O i-l (O OT— 1‘ OTOO r-l O O VO CM ATxJ- i—I •4 i-l *4 I-l ■ i.r n n •> *s »\ n xj o m vo vo cm co 11too in co cr VO H O O W H S •4 *4 ‘ CM VO CM xW CM m i n i n r-i r-i atvo at CM rIN-Hh- m C M or xj cm in.T|- m m L— xj r-l or rovo re i-i~ t r-i m o m XT’ i-HCO CM rH r-ivo o m o r— r— o L— xovo^tvo m o w n h— in o c o rHio vo •4 xt* W L - H LflHVO ,0 Pi CO ai r— i * *H m rH CM Alxi e Nm v o h- * rH cm m.-j- m v o UT --J- in or l- co rH O ( M S I-l CT mATxtco ino Ol*A-* O «x f• O•C --W 1-1 CCiVO xj c\ e xt * rH AI ATx-t IOC i —I OT TA3LE 36 (cont’d) Stem Me. 4 5 local Dry Height Hood + Bark Ia 1 local Dry Height Hood * Bark Branches, One-Yearu Dry Height Twigs r Dry Height Leaves Hood 4-Bark A =) D3H (cm) Height (n) 10.3 4.7 11.5 11.2 4.1 3.0 5.5 9.0 8,0 4.5 31,360 4,301 34,890 31,881 2,904 2,332 397 1,724 972 290 14,125 2,327 17,573 19,903 1,500 4,139 1,175 !>17l 8,950 340 316 146 239 341 80 1,668 629 1,770 2,547 264 4.7 6.6 2.5 4.8 5.4 1.0 3-5 7.0 7.0 4.0 6.0 5.5 3.0 4.5 5,402 12,825 1,195 6,349 4,662 450 2,009 416 1,080 46 447 291 37 210 2,484 6,236 646 2,864 3,057 257 733 3o3 1,484 13 382 747 44 374 77 164 n 33 1 39 28 89 272 763 19 279 480 58 293 1.2 S.o 4,3 7.4 15*3 2.5 8.0 4.5 5-5 3 ~ 236 35 1,202 3.7,027 312 3,713 545 9,163 1,450 61,575 ■vrrv:— . 1■ 143 9, £74 1,656 4,958 34,729 -9 22 of stem section X height1) 'c Mean weight of last 5 years iih *^ 3,074 — > 1,153 130 572 427 150 917 2,237 12,564 431 3,255 formula (mean of areas at each end 107 Site 2 7.6m 1 2 3 4 5 45.7m 1 2 w' 4 5 6 7 91.4m 1 o Stems local Cne-Year* Volume Volume Hood Hood ((”71^^ 108 TABLE 37. DRY WEIGHT AND PERCENT DISTRIBUTION OF BARK AND WOOD FOR STEM DISCSa OF ALNUS RUGOSA Diameter fern) 17.0 16.6 l 4 .0 13-2 1 3 .0 12. 3 1 2 .0 1 1 .8 1 0 .A 9-1 8.7 7.9 7.8 7-5 ‘ 3.8 5.4 4.4 4.3 3.4 3-3 2.8 2.6 2.6 2.3 2.2 2.0 1.7 1.4 1-3 0.8 0.8 0.7 Total Weight ffi) Wood Weight (6 ) 173.0 213.8 1 3 4 .1 109.3 80.3 129.8 62.5 81.5 83.2 87.4 51.2 40.8 52.2 27.6 15-4 19.5 19-5 7.9 3*9 10.5 5.6 4.8 5.8 3.0 3.5 3.2 3.6 1.4 2.5 0.6 0.5 0.2 153.8 195-2 116.0 95-1 70.2 112.2 53-5 71.0 68.9 72.9 42.8 35-5 46.4 24.0 13.7 16.4 16.6 6.4 3.2 8.9 4.9 3.8 4.2 2.3 2.6 2.5 2 .4 1.0 1.8 0.3 0.3 0.1 1 to 3 cm in length Bark. Weight (S) Wood % of Total Bark ‘/o of Total 19.2 23.7 18.1 14.2 1 0 .1 17.6 9.0 10.5 14. 3 14.5 8.4 5-3 5.8 3 •6 1.7 3.1 2.9 1.5 0.7 1.6 0.7 1.0 1.6 0.7 0.9 0.7 1.2 0.4 0.7 0.3 0.2 0.1 88.9 91.3 8 6 .5 8 7 .O 87.4 8 6 .4 85.6 87.1 82.8 83.4 83 ■6 87.0 88.9 8 7 .O 89.0 84.1 8 5 .I 81.0 82.5 84.8 87.5 79.2 72.4 76.7 74.3 78.1 66.7 71.4 72.0 50.0 60.0 50.0 11.1 8.7 13.5 13.0 1 2 .6 13.6 14.4 12.9 17.2 16.6 16.4 13.0 11.1 13.0 11.0 15.9 14.9 19.0 17 •5 15.2 12.5 20.8 27.6 23.3 25.7 21.9 33 28. 28.0 50.0 4o.o 50.0 109 TABLE 38* DRY WEIGHT AND PERCENT DISTRIBUTION OF BARK AND WOOD FOR STEM DISCSa OF FRAXINUS NIGRA Diameter (ern) 6.3 5-8 5-4 5.2 4.7 4, 5 4.5 4.0 3.8 3-2 2.8 2.5 1.8 1-7 1.2 1 .1 1.0 0.7 0.6 Total Weight Bark. Weight Wood Weight U) (s) 16.1 14.7 16.6 14.8 1 2 .8 7-4 4.5 4.8 14.9 12 .5 14.4 13.0 10.9 6.5 4.1 4.2 6.8 6.0 4.7 3-2 3-1 4.0 2.7 2.6 1.6 2.1 _ 1.2 2.2 1.2 1.8 1-9 0.9 0.4 0.6 0.8 0.7 0.5 0.5 0.5 0.4 0.3 0.1 0.2 1.1 0.5 0.4 0.2 0.5 0.4 0.2 0.1 0.1 0.2 0.1 0.1 a 1 to 3 cm in length Bark of Total 92.5 8 5 .O 86.7 87-8 85.2 87.8 91 •1 87-5 7-5 15.0 13-3 12.2 14.8 1 2 .2 8.9 12.5 8 5 .I 84.4 83-9 76.2 73-3 62.5 80.0 50.0 50.0 50.0 14.9 15-6 16.1 23-8 26.7 37-5 (e) 1.5 0.8 Wood % of Total 88.2 11.8 20.0 50.0 50.0 50.0 no TABLE 39* MACRO- AND MICRO-NUTRIENT VALUES FOR FOLIAGE OF THREE DOMINANT W O O D Y SPECIES ON TWO SITES AND TWO D ISTANCES FROM DRAINAGE DITCH, JULY, I 968 p K ( 7 a M g — Species 7.6 _________________________________________________ — Alnus rugosa Site 1 Site 2 Fraxinus nigra Site 1 Site 2 Cornus stolonifera Site 2 91*4 7*6 91-4 - 91*4 — 0.76 0.82 O .96 0.84 0.87 0.20 0.84 0.l8 0.16 0.17 0.15 0.13 0.20 0.19 1.12 1*24 1.24 1.24 1*93 1.75 1*75 0.52 1*84 0.56 O .50 0.60 0.28 0.24 1.00 1.24 2.26 2.24 0.34 0 .36 Mn Fe 7.6 91-4 7*6 PPM Alnus rugosa Site 1 188 Site 2 106 Fraxinus nigra Site 1 152 Site 2 121 Cornus stolonifera Site 2 188 m Cu ---------------------------- 7.6 91*4 ---------------------------- 91*4 7*6 91*4 ------------------------------ 121 144 100 106 84 89 94 118 94 97 1 8 .3 15.6 1 2 .8 15 *6 92 198 62 92 62 106 88 118 91 130 14.9 11.4 13*5 14.9 144 59 45 112 88 ' Alnus rugosa Site 1 Site 2 Fraxinus nigra Site 1 Site 2 Cornus stolonifera Site 2 7*6 — 0.82 0.68 Na Species 91*4 — 0.18 0.16 0.21 0.20 --------------------------- Species 7*6 m W" Zn JrJrM 7.6 91.4 in 8.6 10.7 "AT" ' 7*6 91*4 7*6 25*1 27*7 24.4 29.0 24 21 23 21 63 63 63 82 26.4 26.4 25 *1 30.3 21 19 16 24 76 76 76 88 21.1 19*8 19 19 88 82 91*4