3H ELA.,.1_.'.':'-J1 ._A. “whiny. . .u my”; 1; ”Mg; :~;.- 1...»,sz % THE LOWLAND HARDWOOD FORESTS OF INGHAM COUNTY. MICHIGAN: THEIR STRUCTURE AND ECOLOGY Thesi: for the Dogma of Ph. D. MICHIGAN STATE UNIVERSITY Robert Louis Bryant 1963 IH ESlS This is to certify that the ‘ thesis entitled THE WND HARWOOD FORESTS OP INGHAM COUNTY, MICHIGAN: THEIR STRUCTURE AND ECOLmY presented by ROBERT LOUIS BRYANT has been accepted towards fulfillment of the requirements for # degree in M 9 /3 ‘ , L Zfififl‘z (£1 Major professor Date September 6, 1963 0-169 LIBRARY Michigan State University 06L” WM llllll \II ‘ III—.0 ‘1 III II I ll1. \\ III I ‘ I |2 ‘1 I“ III I IN 4 i L E: DATE DUE DATE DUE REMOTE STORAGE P CE IN RETURN BOX to remove this checkout from your record. DATE DUE LIBRARY Michigan Sm. University {tn-:24 l rm t; l 1:325 tg;a-:itx:.‘i,>~+mNIL-vamp r-‘(mssrs oF‘ 13cm»; canny, mummy; lsuzs S RUCTdnE Ann Etntocy by tiubtzrt Lewis ilryrmt A considerable acreage occupied by lowland-hardwood {crests in southern Hichiun'i is considered to inn/e a storage and stabilizing influence -3n ground water supplies. Since little is known about tnc composition, {runnd water hydrology, successional relationships, and general ecology of [uuSe forests the present study is an attempt to provide preliminary ingor- naatlon on these Subjects. Neasurcmeut ot the nowher, size, and distribution of tree species in t? we lowland-hardwood stands was accomplished using (our sampling methodsc .5. 1 plot method was used as a standard for the study ag inst which data were com ared from the other methods. Non—areal methods included random pairs, the point*centered quarter, and the variable plot-radius methods. in order to determine the relative reliability of the various samp- ling methods, simple correlations (r), were calculated between three parameters (basal area, frequency, and density), as determined from each of the methods. The relationships indicated by the use of coefficients oi determination (r2) showed that any of the four sampling methods gave almost as much information about the three parameters as did the best method. The same measures (basal area, frequency, and censity) were used in describing the composition of the stands selected for the study. Con- tinuum. indices were calculated for all lowland~hardwood stands. These huhces indicated a rather complete coverage for the spectrum of lowland surw. Importance values calculated trom the three parameters proved to ‘5in“ the best measure of species dominance. lhe exteris of lire and pathogenic iniluences, windthrow and root— ing systems on community structure of lowland~hardwood stands were dis- cussed. Windtnrow and rooting habits of certain lowland tree species served to influence the eventual stand~age~structure of lacustrine turests. An examination of the soil proiiles showed that the nmjority of soil types were alpha-gleys which were poorly drained. Soils were analysed for total sand, silt, and clay content. Other analyses included moisture equivalent, loss on ignition, and pH. Diff- erences in soil characteristics between the stands were evaluated stat- istically. Simple correlations were calculated between the importance values and basal areAS of the ten most abundant species and soil physical prop- erties. Two speciesrnnwcda tendency to be found on soils with certain physical characteristics--black cherry and slippery elm. Because the presence of non-significant correlations between soil characteristics in different horizons was great, it was suggested that future vegetation studies in the lowland~hardwood land type snould ignore the measurement of physical soil parameters. Keekly measurements of water table levels during the 1961 growing season were recorded for 72 ground water wells. Differences in water table depths between the stands were eValuated statistically. The great- est changes in true water levels occurred in beta-gley and certain alpha~ gley soils. Actual water table variation was least in poorly-drained mocks. ’impie correlations which were calculated between the importance values and basal areas of the ten most abundant species and ground water depths during tne growing season involved two species--Sugar maple and Amvrican basswood. Additional statistical analyses were performed barwven water table depths and the date of measurement. It was suggested that a subsequent series of ground water measurements to give maximum intornmtlun on nater table depth could be substantially reduced. As a means of depicting the behavioral pattern 0t lowland-hardwood {crests the stand to stand relationships based on vegetation similar- ities and environmental measures were included in a series of graphs. using the continuum as an index of similarity was not unlike the figures derived from a formula method. Succession in lowland-hardwood stands implied by the must significant environmental measurements was repre- sented in a dimensional figure. COpyright by ROBFRT LOUIS BRYANT 1961+ 'fllll LOflLAND-HARDNOOD FORESTS OF INGHAM COUNTY, MICHIGAN: THEIR STRUCTURE AND ECOLOGY by Robert Louis Bryant A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree DOCTOR OF PHILOSOPHY Department of Forestry 1963 A 1': 9' ”(W Li W ‘ L. “1’ ‘2' -0— The author wishes to express his appreciation to Dr. Jonn h. "antlon of the Department of Botany and Plant Pathology for his helpful advice in the preparation of the material on stand survey methods and stand similarities: to Dr. Donald 8. white of the Forestry Department and Mr. Ivan F. Schneider of the Department of - Soil Science for their assistance on soils and ground water hydrology, and to Dr. Johnathan w. Wright of the Forestry Department for his suggestions and help in computer technique and statistical analyses. I am also indebted to Drs. R. Keith Hudson and Victor J. Rudolph of the Forestry Department for their patience in reviewing the man- uscript. Appreciation is also extended to Mr. John L. Arend of the Lake States Forest Experiment Station and the P. 3. Forest Service for furn- ishing transportation for the ground water study, and to the personnel of the computer laboratory in the Department of Electrical Engineering, who voluntarily offered their services on some of the computations. I am especially grateful to Dr. Terrill D. Stevens of the Department of Forestry ior his generous counsel and efforts in coordinating the various phases of work concerned with this study. Lastly, I am indebted to my wife, Nadine, for her encouragement and help throughout the study. ii .\“L.l(;\‘j()‘:.:1"1Dfijlv;:\ll-‘:“Yr E.) o o n 9 ...... O 0 0 ooooooo O O o O O O . :\:1 “.~)I):3(:1‘]();\10 I I O O O I O . D O O O U I O 00000 O 0 C O O O O O O .RLY HESTURY 0F fin AREA. DESCRIPTION AND E: CLIMATE.1. IOOIIOODIIOOQUOOI P}{\’ST{)(:lz-I\E)}l\) 8‘)! 145 g lA‘K‘VI‘) (;l.-(}ll()l(:\’o o o o o o o o o o nnnnnnnnnnnnnnnnn Ecology in Cenorai-Pre-Uistory........... V Lowland Hardwood Types in the Distribution. ..... ................... hydroworphic Soils and Cround Hater. Lowland Hardwood Distribution.. Hydromorphic Types in the Previous studies-Hydronorpnic soils and lowland hardwoods and Eastern finited ‘t*1tes O O l O O 0 Lake States 0.- Ceuira &:Oii.:)o.oi00.....IOOCOOIIOOOOOIIOOD .00.... Ground Water........................................... hardwoods.... Lowland Hardwood Types in Southern Fichlgan................ Distribution............................................. Hydronmrphic Soils....................................... Previous studies-Hydronmrphic soils and lowland hardwoods Ground Water............................................. hardwoods. Previous studies-Cround water and lowland Previous studies-Hround water and lowland sampling blethOClSooooooo-ooooooooooooooo PlOtSooOOOOO0000.00.00.00.-QOOOOoo-oo Sampling Efficiency~Time............. Sampling Efficiency-information...... Stand SLMilaritiESoocooo000.000.00-0000 METHODSODOOOOOOOOOOOOOOOOOOOOOOOIO0.0.0COD Stand Survey........................... Plot Selection......................... Sampling Hethods....................... Stand Characterization................. Soil Measorements...................... 0 O 0 Ground Water and Precipitation Measurements. iii l States ..... I ' u - M... ii l} l3 l7 17 20 22 22 2a 24 27 27 29~ 29 30 31 32 32 34 34 35 36 39 41 Al 41 A3 45 46 a8 TABLE OF CONVENTS (continued) S3LatisticsooOOOOOOQCOOOOCOOOOOOOOOOCOOOOOOOOOQOOOCOOOOO.IOIOOOOCOO {fliSL1CIQOIOOOOOOOIO....OOOOOOOOOOOOO0.0.0.0....00......OO...O... Analyses 0f \',ariallc-t,o . o 0 o o o O O o o O O. O o o O O O O I I D O I O O o O o 0 I o O o o o O 0 o I O O (torrelé‘tion ;\l‘a]ysebo O I O O O O 0 O O I O O O O I I I O O O I O O O O O O I I O O G O O O O I I I O O 9 O Sttt-nd SimildritiesoOOOOOOOOOOOOOIIOOOOODO0.000.000...no.00000on. it‘ll.” LHJ' idoooooooooooooooooonoascoop-cocooooooo000.000.0000....coco-coo Sampling MethodS.................................................. Stand Characterization3....................o...................... Comparison of Quantitative Measurements and Species Dominance... Determination of Soil Properties.................................. Beta-Gley Soils................................................. Alpha-Gley Soils................................................ Organic SoilS................................................... Soil Properties and Species Development........................... American le.................................................... Red Maple....................................................... Green Ash....................................................... Swamp Hhite Oak................................................. Silver Maple.................................................... Sugar Maple and American Beech.................................. Lesser Species.................................................. Slippery E1m00000000000000000900.00.00DOOIOOOOOOOOOOCOOIIODOOO Northern Red Oak.............................................. Black Cherry.................................................. Ground Water and Precipitation Measurement........................ Ground Water and Species Development.............................. Silver MapleOOO...OD...OOOOOOOOOOOCCOCOI.QOOOOOIOOOOOCOOOOOOOOOO RBd MaPIQOOOOOOOOOOOIOOOO0.00.0000...OOOOOOOOOOOOOOOOOOCOO0.0.0. American Basswood............................................... Understory Vegetation and Ground Flora in Lowland Hardwoods....... Biotic Factors and Their Eiiects on Community Structure........... Fire Influences................................................. Nindthrow, Rooting Systems, and Community Structure............. Pathogenic Influences and Community Structure................... StatistiCSooooococonut....ooooooooooocoo-ootoo-0000.00.00no...coo. Correlations Between Sampling Methods........................... Improvements in Methodology~5ampling Lowland Vegetation......... Soil Measurements............................................... Profiles and Horizons oi Beta-Cley Soils...................... Profiles and Horizons of Alpha-Gley Soils..................... Profiles and Horizons of Organic Soils........................ Underlying Materials of Beta-Gley, Alpha—Gley, and Organic Soils......................................................... Correlations Between Species Distribution and Soil Character- istics........................................................ Distribution of one species not related to that of another.. Species distribution slightly related to soil physical prOperties...................o.............................. Correlations Between Soil Characteristics in Diiferent Hor- izons...O....0.IUOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOCOOOC0...... iv 79 79 81 83 84 855 85 8o 86 87 87' 68 88 88 89 90 103 103 105 105 106 114 114 115 123 125 125 128 130 130 130 131 131 133 133 134 137 Tfihuh OF COMTHNTS (continued) '1 ._r 1 kn t improvements in Methodology—Sampling Lowland Soils.............l3U Ground Water and Precipitation Measurement....................,,.La] Fluctuations of the water Tables Within Soils..................!n} Fluctuations of the water Tables Between Soils.................inn Correlations Between Species Distribution and Cround hater.....)wl Species distribution related to ground water depths.,......u.15i Correlations Retween.fiater Table Depth and Date of Measurement.135 Improvements in Methodology-Sampling Ground hater.....,,,,“.,,,159 stand Similarities....................u............g..o..........101 Stand Shift Related to the Number of Stands Sampled............103 Stand Shiit With Changes in Organic Matter.............,.,..,.,163 Stand Shift with Changes in pH.................................lb? Stand Shift With Changes in Sand, Silt, and Clay Content.......]n7 Stand Shift With Changes in Moisture Equivalent................172 Stand Shift With Changes in Water Tables.......................174 Succession in the Lowland Hardwood Complex................a....181 DiSCt‘SSiOnOIO...OOOCOCCOCOIOOOOCOOOOOOOOIOOOOOO0.0.30.000000090 187 C‘I_7_'1:!:“;&'o0.0.0.00.00.0....I00.0.0000...OOOOIOOOOOOOOOOOOO00.00.00.000! 18’ LIL~ER4“\L"1JI{E:.II'V‘T-IVHJL'V‘JLo-coogg0.00.00.00.00.OOOOOOOOOOOOODOOI0000.00.00.00...19‘ :11)I’r:NI)1XIOOVO.00...O.OOOOOOIOOOOOOOOOOOOOOOOOi0.......OOOOODIOIODO...2f}; ‘1’11"\..OOOOO0O00.00.O0....00.0.00...IOOOOOOOOOOIOOOOOO‘00.9.00.0.0.0.. 3]} 'I‘ah 1 e -‘nt a—oo-m 1V V11 Vlli 1X XI X11 XIII XIV 1,18“? OF TABLES 5' Mean unnthly temperature and precipitation data for a 60-year record and the 1961 growing season for ingham County, Mich- ,- ‘o/ 1gan............. 00000000 0.00.0.0... 0000000 OOOQOOIOOOOOOOIOOI Najor canopy species in the lowland-hpriwood types of the (85:91." [It‘ite‘i Stilt-CS....oooo‘o'... IIIIIII 0.0.0.0000000100090 1‘} Clinmx adaption numbers for lowlandehardwood tree species found in lngham County, Michigan............................. 47 A summary of forest survey tree data tor the lowland-hardwood forests of lngham County, based on the 100 meter2 plot method. 55 A summary of forest survey tree data for the lowland-hardwood forests of Ingham County, based on the random pairs method... 57 A sunnery of iorest survey tree data for the lowland-hardwood forests of lngham County, based on the quarter method......... 53 A summary of forest survey tree data for the lowland~hardwood forests of lngham County, based on the full Bitterlich method. 59 Importance values of tree species found in 19 lowland-hard~ wood stands in Ingham County................................. 67 Basal area per acre for tree species found in 19 lowland-hard- wood stunds in Ingham County................................. 68 Soil series in the area covered by the soil survey arranged to show the parent nuterials and catenal relationships....... 80 Species frequencies in 19 lowland-hardwood stands in Ingham County, based on saplings 1.1 to 4 inches in diameter........ 108 Species density in 19 lowland-hardwood stands in lngham County, based on saplings 1.1 to 4 inches in diameter................. 109 A species list of the ground flora common to the lowland-hard- wood stands of the Ingham County showing the degree of import- ance attained by each speCieSOOOOIOOOOOOOOOOCO000000.00.00.000 112 Species mortality (basal area) in 19 lowland-hardwood stands in Ingham county-009.0000.000.000.0000..coo...cocoooooooooooooooo 122 vi (C Table XV XVYYI XIX XX XXI XXII XXIII XXIV XXV XXVI XXVII XXVUIII XXIX LIST 0F WAHLES (continued) Correlations between species' basal area, frequency, and den- sity as measured by four sampling methods...................... _ , _- , f) Coefficrents ot determination (r‘) for species' basal area, frequency,and density as measured by [our sampling methods..... Relations between some physical characteristics of soils and QDECieS iin¥)ort-R’IC(.OOOOI06.00....COCOOODOOOOOCOOOOOOOOOO0.0.0... Correlations between various soils characteristics in 10 stands 0f lowland-hardw00d$ooooooo00o¢oo0.00so00.000.00.00000000900000 Eumnwaryr of the analyses of variance showing the difference in perched water tables due to week of measurement throughout t!]e1961growingseaso!1-IODOOOOO0....OOOOOOOOOOOOOOOOO....00... Sunmmry of the analyses of variance showing the differences in phreatic water tables due to week of measurement through- out the 1961 grOWing season........o.o......oo..........oo.... Summary of the analyses of variance showing the differences in perched water tables between stands throughout the l9bl gTOWlllg season...000.000.0-...no...0.0000000000000000...-coo.0 Summary of the analyses of variance showing the differences in phreatic water tables between stands throughout the 1961 ngWing season.0....OI00.0.0.0...0.00.00.00.0000COOOOOOOOOOOOC Mean depth of water in soils for the 1961 growing season...... Correlations between ground water depth and importance of sugar maple and Anerican basswood................................... Relationship between the range of ground water depth and im- portance of sugar maple and American basswood................. Correlations between water tabbedepth and date of measurements Optimum sampling data in 1961 to give maximum information on water table depth at minimum effort....o......oo....o......... Community coefficients (W: 3C) based on the importance values of tree species in the 19 lowland-hardwood stands of lngham countYOooo0.0.0.0000...00.00.000.00...0.0.00.0..00000000000000 Profile means for alpha-gley and beta-gley soils, showing differences between soil texture, moisture equivalent, loss on ignition, and reaction...o..o.......o..................o... vii l)\1 I»): L? 127 135 1/42 1A4 1&5 14b 152 153 156 160 lb 2 306 XXX ~in t" A-‘ I. .L IVIXI ll AAXlV LIST 0? lABLHS (continued) Horizon means for beta-gley soils developed from glacial drift, showing difierences in soil texture, noisiure equiv— alent, loss on ignition, and reaction........................ Horizon means for alpha-gley soils developed frow glacial drift, showing differences in soil texture, moisture aqui- aleni, loss on ignition and reaation......................... Horizon means for alluvial soils showing difterences in soil texture, noisture equivalent, loss on ignition and reaction.. Profile and horizon means for organic soils, showing difier- ences in loss on ignition and reaction....................... Horizon means of underlying materials of the beta~gley, alpha-3133,! and organic soils showing diifervnces in texture” viii Pi! a: 2b? 203 2U9 210 LlnT (IF FTCI'R‘r.) _E;i_u 111:5: l‘;1,:e- l qu oi lngham County showing the surface geolngy and locaLion of the lowland-hardwood stands selected for the ve£:etdti-On Slu‘-iy°.......5.....IOOOO.II.OOOOO.09..b...9.. {41. ll bampling methods used in the lowland-hardwood forest S‘lrve‘j'000o0Iaoooooo0000000090.0.octoooooouoooooooooooooo Ill Map of Inghdm County showing the drainage system, loca- tion and number of ground-water wells in the stands se- lected for the ground water studY....................... 31 IV A ground water observation well in the shallow Linnwood IUUCk Of Stand [.10Looooeoooacouoosoooooooaoooooo3.0.0.000 5‘ V Interior view of stand was on poorly-drained benewu,loum -~ ' '1 COULinuum IHdGX 1)QQOOOOOOOOOOOOOOOOOOaOOOOOIOOOOI10.0.00 6; VI Interior View of stand HZOH on poorly~drnined Riile peat, continuumindex1408...“,H.“H.....,,,.,,.,,,.....,.°.U) . 9 VII interior view of stand 3225 on poorly—drained bloan 10am, COhtilIUUm in‘ifi’x 1"...“3'3000000000000OuOOOOOQOIOOIOOOOOOOOOOO0 [’3' Vlll Interior view or stand M17C on ponrly~drained Cohoctuh sandy loam, continuum index Dam.......................q “1" Ix Interior view of stand M268 on poorly—drained Hronkston loam, continuum index 1059...,,...,...n.,,,,,,,,_,,,,,,,, no A Interior view of stand VZUC on poorly-drained Carlisle muck, Continuum index 1818......°............o.¢..a..a... «9 AI Interior iiew of stand 5238 on poorly-drained Wrookston soil, continuum index 1827;,.................l............ ? Jill Interior view of stand LlOL on shallow Linnwnod muck, 7§ corltinklun‘ illdcx ’8Zipgggggoablcoioon00.000.090.000030000-l.‘l AIlI Interior view of stand £38 on poorly-drained Pewnmo loam, mti‘1utm il‘uex L92(~).....,IOIOQIOOIOb0.....00090090000000’) “1" Interior view of stand A148 on poorly-drained BrOOkston sandy loam, continuum Pun nflu....a........................ ’“ ix Figure XV XVI XVII XVIII XIX XX XXI XXII XXIII XXV XXVI XXVII XXVIII XXIX XXX XXXI. LIST OF FIGURES (continued) Interior view of stand NSC on imperfectly-drained Conover 5011’ colitj-nuum index 2155....IOOOOOOOOOOOICOOIOOOOOOOOOC... Interior view of stand M193 on imperfectlyvdrained Spinks lomy sand, continuum illdex 2160....0OOIOIOOOOOOOOOOOOOOOOOI Interior view of stand Ml9L on imperfectly-drained Locke sandy loam, continuum index 2429............................ Hydrographs of true and perched water tables in moisture regimA (St-ands rllgs and h11914)000000000.00.000.000.0.09.... Hydrographs of true and perched water tables in moisture regime B (stands AlQC, A148, and AlQL).....o.............s.. Hydrographs of true and perched water tables in moisture regime C (Stands LIOG, LIOL, and VZUC)OOOOOOOOOCOOO9.000.... Hydrographs of true and perched water tables in moisture regime D (St-and was).O...OOOOOOOIOOOOOOOOOOOOOOOOOOOOCOOED... Hydrographs of true and perched water tables in moisture regim D (Stalld WISB)OOOOOCOOOOCOOCOOOOO0.00.0.0...0...0.00.. Hydrographs of true and perched water tables in moisture regime E (stands M268 and M26R)...........................°. Hydrographs of true and perched water tables in moisture regime F (Stands M228 and M17C)oooooooooooooocoooooooooocooo An 18 year record of mean water levels for the only obser- vation well in the Lansing area glacial drift............... A layer of silver maple (Acer saccharinum,Harsh.) seedlings on the Cohoctah sandy loam of stand M17C.................... A fire scar gives mute testimony to the fact that this area was swept by a low intensity fire ten years previously...... A young stand of quaking aspen (Pogulus tremuloides)on the alpha-gley Sebewa loam adjacent to stand wts................ A silver maple (Acer saccharinum) located on the deep Car- 11318 mUCk Of Stand VZOCOCOO.0...’..............OC..OOOCOCOO An American elm (Ulmus americana) growing on the Pewamo loam Of Stand W480...no.coco-cooooooooooooooooooooooooooooooooooo 77 78 91 94 95 96 97 100 104 116 117 119 121 The shift in spatial relationships of lowland-hardwood stands, which occurs when stand M26R is selected in place of stand 026R as a base of reference..................................16a X LLST OF FIGURES (continued) than: Bees: XXKII Relationship of lowland-hardwood stands based on the organic matter content of the soils (loss on ignition) and W values... 166 XXXILI Relationship of lowland-hardwood stands based on the pH of the $01.15 al'Li ‘1‘: valuescoconnect-00000000....ooccoooooooooooooooooo 1‘38 Hikiv Relationship of lowland-hardwood stands based on the percent SK1IId 1:11 the SOils (1“‘(1‘3W‘Vi‘lueSOOOCOOOCOOOOIOOOOOOIOOOOOOOCOCQO 169 IXKV Relationship of lowland-hardwood stands based on the percent Silt 21‘] the SC’ils al‘d b: V.é11‘JeSOCOOOOOOOOI0....OOOOOOOOOOOOOOOO 17(3 EXXVL Relationship of lowland-hardwood stands based on the percent Clay 1“ tile: SC)le 3‘1‘1k‘Vail-(188..OOUOOOCIOOOCO..0......IOOCOOOO 171 XXXVL! Relationship of lowland-hardwood stands based on the moisture eQUivalent of the soils and w values.......................... 173 TKAVIYI Relationship of lowland-hardwood Stands based on mean water table depth and w values showing the toposequence oi water tilbles Witi1ln t‘t‘e soil-s:QIOOCOUODOOOIOOOO’COOCOOCIOCCOOOOUOCOC 175 I u > ,V ‘21 v-fl X Relationship of lowland-hardwood stands based on mean water table depth and w values showing the range of water table fluctuation during the growing season......................... 176 XL Relationship of lowlandshardwood stands based on their con- tinuum index placement and average water table depth during tile 8r0w1ng SEBSOH..........o...........o.....o...o..o.oo....o 178 XL? Relationship of lowland-hardwood stands based on the organic matter content (loss on ignition) and a logarithmic scale of W values X mean water table depth X range of water table fluctuation................................................... 180 XLII A construction for 15 lowland—hardwood stands based on their present continuum position, organic matter and moisture equiv- alent units, and units of mean water table depth and fluctu- atLOnOOOCCUOOOOIOODO0.0IIOOCCOOCOOOOOQOOQOOIOOIOOOOOOCOIOCO... 182 xi introduction in recent years the use of quantitative measurements in phytcsocioiugi- cal studies has been devored largely to upland forest communities (130, 23, Fewer oh Losoc10107ical studies have been carried out i .V b 1 7,1'52,60,28,S3). 'c": exclusively in lowland hardwood iorests probably because of the numerous discomforts which are always present, including baggy terrain, irritating insects and ohnoyious plants, which usually impede progress in the col- lection of quantitative information. Those studies carried out in spite of these difficulties, include the efforts of Christensen, Clausen and Curtis (40), Ware (150), Lindsey (102), and Wistendahl (lb7). The lowland hardwood forests of southern Michigan are of considerable importance. Although they occupy only six to ten percent of the com- mercial forest land throughout the region, this amounts to about 750,000 acres in the lower peninsula of the state.1 The considerable acerage occupied by lowland hardwoods mnkes it one of the major forest types in Michigan, whose future economic potential as a source Of forest FIOductS Indy be great. Such a large_£orest0d area. occupying sites which are so poorly drained, serves as a catchment basin for precipitation and runoff (hid acts as a storage and stabilizing influence on the ground water sup- ;ulies in this section of the state. the present condition of the majority of stands in this type consists orT‘overmature, defective and cull Specimens left aiter cutting and utilize- ticn1 practices which removed only the most valuable trees. Lowland hard- umxads Lmually have been bypassed as an area of research in southern lUnpublished information, Lake States Forest Experiment Station. -1- -2- Michigan and most or the research effort requiring detailed investibato ions has been devoted to the more valuable oak, oak-hickory, and sugar maple-beech iorests on upland soils. fhe lowlands in hardwood forests are primarily of glacio~lacustrine or fluvial origin, occupying such depressional features as elongated glacial drainageways, swales and basins of till plains, ground moraines, and shallow muck swamps. in addition, these forests are often found along the borders of many lakes and streams, small tributaries, and enclosed upland depressions and bogs. The major tree species of these iorests in the northern part of the lower peninsula consists of pure conifers to mixed conifers and hardwoods, while the southern part consists of nearly pure hardwoods. The primary objective of the present study was to describe and compare the structure and several environmental characteristics of a series of stands in this poorly known lowland forest type. The environmental parameters were mechanical analyses of soils and the changes occurring in water table levels. Basic information on the ground water hydloiogy for this section of the state consists only of information on represent- ative wells influenced by pumping. A second objective was to construct estimates 0: the successional relationships among the lowland forest types. Because the collection of hydrolobic data requires a deiinite schedule oi obserVations in wells, the area within which the stands were <1hoson was kept snwll. lngham County, in which the study was made, is CCNISidered reasonably representative of the 13 other counties ir scutwgrn [agwen'fiichignn. it contains 13,200 acres (112) of lowland hardwoods, it rather large portion of the type occurring in the l3 connLy area. 3.19:9$.IZLPES;S?13-_LJQ.§. -!:».«..tt_l_r_~‘-.7.ziter1_0.t- steered lngham County, the area in which this study was made, is located in the south-central part of the lower peninsula of Michigan. The county is roughly Square in outline, composed of four tiers of townships, and includes an area of approximately 550 square miles within its limits. Several hundred lndians were the first known human inhabitants of Ingham County (03). Since they relied heavily on water as their principal means of transportation, it is only natural that the first dwellings were established along the few waterways that exist in the county. The largest Indian village in the county was established at Okemos, beside a clear shal- low river called the Red Cedar. Early surveyor's notes also indicated the presence of two smaller villages. One was located beSide a small lake in the northern part of Stockbridge township and the other along the Grand River in Onondaga township (639, Overland trails used by the Indians usually followed established water routes.’ Away from the rivers, trails were established on higher ground, usually along the t0ps of winding eskers and ridges of the larger moraines. Two large eskers, running from the northeastern to the south- western part of the country. were used rather extensively as travel routes. According to Fuller (6)), the tribes depended heavily upon upland beech-maple and oak-hickory forests, as well as floodplain forests for food, shelter, and clothing, No mention was made, however, of the extensive low— land hardwood forests, in old glacial drainageways and lowland basins, being usenlior any purpose. It is possible that these areas were avoided because of fear of sickness, iever and pestilence, bred by the hordes of mosquitoes unlich inhabit such places. Small open areas oi wet marshes were mentioned {us being extensively used tor trapping purposes. «L I»\. u. it the time that permanent settlement by white men began, during the 1830's, the entire county was covered by dense hardwood forests. Little pkg was present, except for the area around Lake Lansing. Tamarack, how- ever, was prevalent in the swamps and drains of the watersheds. The early settlers immigrating into the southeastern part of the county, in Stock- bridge township, testified to the fact that dense forests ranged to the west and north. Initial purchases of land for timber and minerals were made in the northern part of the county. It was here, and in the western sections of the county, that the greatest variety of timber species was found (63). The principal species found on the well drained mesic sites were beech and sugar maple, which are considered climax for this region. Originally, the heavi- est timbered sugar maple stands were found in Vevay township. Maple trees were tapped in these early days for the sap, which was made into syrup and together with venison served as items of trade between the Indians and pion- eer settlers. It was reported by Fuller (03), that the most heavily stocked lowland hardwood stands were found in Meridian, Leslie, Wheatfield, Alaie- don, Onondaga, Stockbridge and Vevny townships. Toward the south and east, homesteads were established on the well drained and intermediate textured soils. Unk stands were. found on the (keper sands and somewhat drier upland soils. it appears from the early records that oak timber was very abundant on these soils, and was the most Valuable forest asset in the country. According to the early lumbermen, lflgham County produced the greatest quantity and some ot the finest quality oax timber found in the state (6}). Before the l-.:3£:i}’s, over 21 small saw— mil is had been established throwfimut the county. The largest mills were ‘ V v n . .- q I o intated JD wheatfield, OUOHOHdd, and hiOCkbflUflL townships. . A. . r- Fm. -~., 'I -5)- During this time, millions oi feet of oak timber were cut to supply lo- cal and regional markets to the west. Oak was used for practically every- thing, including shingles and flooring. staves for barrels and casks, wheel- barrows and hand trucks, bridges, corduroy roads, and railroad ties. After the railroads were built, oak lumber was sent to the shipyards of the east- ern markets. Waste slabs and cull lumber were made into charcoal, shipped to the northern part of the state and used for smelting iron. In these early days the timber on upland areas was also cleared for agriculture. Plowing and planting these lands to various crops reduced the natural fertility of the soil, and more land was cleared as yields became progressively lower. Little was don: by the landowners to protect the soil from erosion or to increase its fertility. As a result of this treatment, a great amount of soil material washed into and filled the natural drains. In a very few years competition for finer textured soils increased as mar- ginal droughty uplands were abandoned and eventually some of the poorly drained mineral soils and mocks were placed under cultivation. In earlier times, these poorly drained areas were considered unfavor- able tor the production of crOps and continued to support lowland hardwood forests. Since the cost of drainage is high many areas have remained rela- tively undisturbed until the present time. it appears from the early records, that lowland hardwood forests were not generally exploited for their timbers, at least to the extent which occurred in the oak forests of the county. One exception to this situation occurred in lowland floodplain forests, whicn favored the deveIOpment of sycamore, cottonwood, yellowpoplar, black cherry, butternut and black walnut. It appears that the walnuts, prized for their use in furniture and Cabinet making, and black cherry, a fine vuaod used in the manufacture of coffins, were subjected to such des— LrUCCiAee cutting that they were practically obliterated on floodplain soils. In the early ddys of pioneer settlement, tamarack was cut rather extensively on the wet organic soils and hewn into ship timbers (53). The agricultural potential of the lowland soils was found to be high under conditions of proper drainage and in recent years some of the better lowland hardwood stands have been cut. Original forest growth on fine tex- tured soils of imperfect and poor drainage, including the wet organic peats and mocks, iavored a devel0pment of large individual trees. Elms, ashes, soft maples, basswood, and sWamp white oak were the principal Species ac- cording to Veatch, £5 21- (l47). Tamarack, quaking and bigtooth aspen, white and paper birch, willows and an occasional black spruce were also present. The neutral to basic organic soils also supported a lush growth of shrubs, sedges, and grasses, in addition to the previously mentioned species of treeso On highly acid bogs, the vegetation was shrublike consisting mostly of leatherleaf, dogwoods, and wintcrberriCS. At the present time about 80 percent of the total land area in Ingham County has been cleared for agricultural purposes, leaving about 15 percent in forests. Although the greater portion of the forested terrain is characterized by rather good natural drainage, about 30 percent of the {crested area has soils which are imperfectly or poorly drained, Alluvial and organic soils comprise about 17 percent of the lands classified as wet or in a permanently swampy condition. "he mineral soils are rather dark in colxn'and have developed on glacial till from calcareous gray and yellow 103m to clay loam. Where drainage is feasible, most of the organic soils at the present time are very valuable for agricultural purposes, L;nerw:ly the climate oi [Johan County nay we characterized as (OHILH‘ tfiifil in naturt, with extremes in temperature, humidity and precipitation “mt: Imcmrmun. (Ming to t'm mttrior location of the county in the south- 9:2”! peninsula, considerable wuiation many be found in meteorological condi- tions. At times, moisture~ladon winds carried inland from the surrounding ('t‘eat Lakes produc e a senvi~azurine influence, with accoxzipanying underately lailgll l‘tUl‘nldillt 5. Low wind velocities and evaporation rates are the norm for this section of Michigan. Prevailing wind movement in the county is from the southwest, averaging 8.2 miles per hour, except for the month of March, when wind di- rection from the northwest has an average velocity of 10.2 miles per hour. The highest wind velocities have been recorded from the northwest compared to other directions (353. he average annual precipitation in the county amounts to about 31.1 inches. The driest year for 60 years of weather records occurred in 1938 with an annual rainfall of 21.3 inches, while the wettest year, with an an— nual rainfall of 39.7 inches occurred in 1947. Most of the precipitation is from storms which are cyclonic in origin, producing less than .25 inches of rainfall per 510111.033). & fairly recent classification of rainfall patterns for the East Lansing area shows that Host of th'se storms are of low intensity, and that most of them occur during the nmnth of November every year (137). The major Pdl‘t of the annual precipitation falls during the mid-part of the growing 5938011, from May until August, a product of high intensity conVectional s to rms . h;rtet' l<)S:n'S thie to rulhrlt FIN)n1 stt)rnus 8Y1? u«nxal_ly lour, t>xcrqat when the soil surface is frozen or not protectec by a cover of Vegetation. Nevertheless, through perColation, infiltration, and subterranean transfer «at unconfined ground water, :fixnxt 3U percent of the annual precipitation ap- pears as stream [low in the creeks and rivers oi the local watersheds (H7). in addition, melting of accumulated snow cover contributes to runoff during the late Spring months. The heaviest blanket of snow cover on record oc. curred during the winter of 1931-1952, with a mean depth of 88.8 inches. The average length of the growing season for the county varies from 184 to 158 days (163). Although the state of Michigan is noted for some of the greatest extremes in temperature within the continental limits of the coun~ try, such is not the case for Ingham County. The 60-year reCord indicates a mean annual average temperature of 46.70F. with a mean deviation of 19.80F. between maximum and minimum means. The highest temperatures usually occur during the month of July and the lowest during the month of February. Aver- age dates of the last killing frost in Spring and first killing frost in autumn occur in early May and late October, respectively. A brief resume of normal climatological data by unnths of the year is COmparcd to the values for the lle growing season in Table I. It may be observed trom the tabular values that mean Monthly temperatures for the pre~ sent year were approxinmtcly nornwl. The rainfall pattern, h-.:z..'ew:t‘, was definitely abnormal. As shown, the tx)Cal annual accumulation was far below the normal trend,.making 1961 one of the dI’ivst years on record. Although not indicated in the table, the accumulated dcParture from normal for each succeeding month was negative tor the entire year, primarily because the initial deficit during the early winter months -9- TABLE.- 1. Mean monthly temperature precipitation data for a 60-year record and the 1961 growing season for Ingham County, Michigan 7— Month Temperature __ Precigitation 60—year 60-year Departure mean 1961 _Jg§g1_ 1961 from normal “F. jig. inches inches inches January 23.8 19.9 1.87 0.45 ~1.42 February 24.2 27.8 1.81 1.60 -0.21 March 33.2 37.0 2.57 2.87 0.30 April 45.3 41.8 2.83 3.45 0.62 May 56.5 52.9 3.75 1.00 -2.73 June 67.4 64.8 3.37 2.97 ~U.MO July 71.1 69.8 2.28 2.28 0.00 August 69.0 68.7 2.68 3.33 0.65 September 61.8 66.6 3.05 4.61 1.36 October 50.5 52.2 2.45 1.58 ~0.87 November 37.9 39.0 2.30 1.77 -0.53 December 27.1 26.3 2.12 1.44 -0.68 Average 47.3 47.2 31.08 27.35 -J.73 or Total O . ' . . ., , ‘- ; 1 . . ‘ ' . math? d ‘h‘: 0‘ .5 .1910 unflilpxu r W' -' r~ -m~-.w-‘~MWM-w lhc wa}or portion of LNG coonty is part of a broad glaciated plain '3- known as the jhumh upland.“ Fhis elevated plain is typical of an area hdv~ "lactatcd, wqose topographic features have bten affected ’.. (J ing been recently hy advances And subsequent retreats of the ice fronts, during Fleisiotcue time. fn inhgam county the plain rises from 200 to 600 ieet above the level 8 o; the Great Lakes and is bordered on the north by a flat lowland plain, which crosses the state from Saginaw Bay to Lake Michigan. lhe lowland plain lies 200 to QOO icet below the ”Thumb Upland" (99). The hnfiesc elevations in Inhgam county are found in the four southeast‘ ern townships, where the first permanent settlement of the county took place. The average elevation in this area is about 990 feet above sea level. The surrounding countryside gradually slopes away from this area in the four cardinal directions. Differences in elevation between the high and low points in the County are not great, averaging about 300 feet. Between low- land swamps and adjacent higher ground, differences average about 100 feet (167}. During the late Wisconsin Age, the ice sheet of the Cary substnge cov- ered earlier Pleistocene tedtures, by depositing an unstratified drift mantle of varying thickness over the county. The soils of lngnam county have de- veloped from this drift and belong, to the (irayfiirown l’odzolic and humic—Cley great soil groups. aneatheled glacial materials oi these soil groups mic usually strong calcareous mixtures of sorted and unassorted deposits. This history of glacial nativity in Ingham county has been described in detail by LCVCrett and Taylor (99, 100). The physiography of the county may best be described as one of smoothly or gently undulating ground moraines and nearly level clayey and sandy outwnsh plains, which are interrupted by old glacial -10.. -11- drainageways and closed basin depressions. A map illustrating the basic geo~ logic formations of the county is shown in Figure 1. In the northern part of the county, 8 low upland called the ”Lansing Moraine , travels eastward from Lansing township into Wheatfield township. The topography along this moraine varies from flat to undulating, and tends to become less pronounced in the northeastern part of the county. South of this area, the Kalamazoo and Charlotte till plains are found, which gradu- ally slope southward at the rate of six to ten feet to the mile. Along the southern part of the Charlotte moraine, the topography again becomes rather hilly. These two moraines, averaging A0 to 75 feet in depth, form a rather thin drift layer in Ingham County, compared to adjoining counties, where the drift mantle averages from 300 to 400 feet in depth (99, 100). Although a system of slender sub-moraines oriented in an east-west di- rection are apparent on the surface f the larger till plains, the nnst striking surface features are the numerous eskers. The majority of these glacial features are oriented in a north-south direction and appear to be strongly associated with elongated glacio-fluvial basins. An examination of the differentiation of materials in these formations indicated that they were probably produced by streams under the glacier, sweeping from side to side in a serpentine manner as the stratified deposits were laid down. Thus, confined and elongated lowlands would have eventually resulted from this type of activity under a rapidly retreating ice trout. The mason esler s stem, one of the largest in the state, and referred to previously as an ancient Indian trail, does not show as strong a rela~ ‘ionship to the elongated basin drptessions, as the Dansville and Williams- tfln csker systems in the eastern part of the county. At the present time, these esker systems are diScontinuous, with several small creeks winding Unwind] Openings created by stream; dissection. -13- Accompanying natural land divisions, produced by the retreating ice masses of the Cary substage, include shallow potholes and swales, shallow and deep muck swamps and bogs widely distributed throughout the county. Additional depressional features include irregularly shaped shallow basins, variable in size and created by sheet flow from glacial melt-waters. lt ap- pears that lowland hardwood forests were predominant in the reneins of these glacial drainageways at the time of settlement of the county by the white nan (63). All of the features previously described are strongly related to con- formations of glacial origin rather than erosion or stream dissection. Ero— sion since the last glacial retreat has produced a modified dentritic drain~ age pattern. Since stream dissection is not prevalent, water tables rennin close to the ground surface in lowlands for the major part of the year, and flooding occurs rather frequently during the spring season. Literature Review igualogy in General -- Pre-history In the field of ecology concerned with the organization and sociologi- CHI relationships of plants, probably no question has been more argued than “What constitutes climax?" Although early workers in the field recognized and described patterns of easily discernable changes in vegetation, Clements (44) contributed most to the concepts of climax and succession. The prin- Clinil concept in his writings was expressed in the unity of the climax and climate. The two were considered inseparable and all successional stages in the development of vegetation pointed toward a single climax. Thus, the nmnclinmx theory evolved, a concept long accepted by many workers in the field. According to this theory, all vegetation, regardless of its starting point in development, would eventually reach theoretical equilibrium under the general control of climate. The visible unity of the ” a unit having similarities between climax was expressed in the ”formation, the life forms of the dominant species. In this sense, it would be expected that all vegetation within a given region, given a sufficient period of time, would be uniform regardless of habitat. if the effects of climate were eventually made manifest in producing stabilization of the vegetation, then a true climax would prevail. in this fPSPvct, climate, climax, and geologic time would be inseparable. The prob- ability for either of the foregoing siuations taking place, however, would be quite low in the natural sequence of events. The specialized often com- rivx terminology used by Clements (Ad) in his descriptions of vegetation obscured the time element, which was largely ignored as an associational lLentity with UN”? Clllf'ldXo -15- -iq- i-i‘ii lo. mst ecologists have, at one that: or anot'wr, stressed the spa- tizil tx:n 1s>w lUDJlJ!FJ ‘Virsuit tV;HFS tit ihzf cerxlrsal axni :n3rtiiexww ix) ‘a?r\‘) I"! (‘ .Ylocust (Kenoyer 86). 9 -m- .»~ 51. .D. J...I-—-. ~50- l:yi.ls occupy an intermediate position between well—drained and poorly drain- eci rnineral soils. High water tables, fluctuating throughout the profiles, have caused strongly mottled subsoils. Wilde (160), in a discussion of 11y‘dromorphic soils refers to this condition, typical of soils which are l>eliiodically wet or insufficiently drained, as the mid-gley or beta-gley Cy?)e of development. The gley layers in these soils are superimposed on tile! profiles at depths of two to three feet. The most common soil series ‘Jitfli imperfect drainage in Ingham County are the Blount, Conover, Locke, VI- ‘ ‘ v n ‘mltlierton and brady series. Poorly drained mineral soils were formed in low areas of glaciofluvial ‘DL'tivash and lacustrine plains which were swampy or permanently wet with the ‘déltier tables remaining continuously at or near the surface. Wilde (160) 0E3].]_s these soils alpha-gleys, owing to the presence of high water tables I’rsbciucing intense gleization of the profiles, except for the inmwdiate sur- f a C e. hot 1220118 . in lngham County the most common soil series with poor drainage atTC? the Brookston, Sebewa, and Cilford series. Alluvial soils, or those found on the floodplains of streams, consist o ... .. f r‘ecently deposited sediments and may be classiiied as either beta or a 13>?)d- gleys depending on the intensity of gleization in the profiles. Since 81 c - , , - —. o u ' a l\J\Ilum is composed mainly oi stratiiied deposits, which continue to be di ‘ . ‘ I v . o l l .- E‘turhed oy seasonal flooding and deposxtion, clear differentiation 01 the prxcv- . . ~ k'1 1195 is difficult. OTitanic soils consisting of peats and mucks were developed from accumu~ lat - 1 - . . , , . {)T]S oi Vegetation in shallow laKes, marsnes, swamps, or old Blacxal dra ' l . . . . . . . r1algeways. Fne thickness oi the organic dep051ts overlying tne mineral b]? "VIQ 3. . ' I ' _ . o . p ‘ciries according to the duration of suitable aCCuwulatIOn conditions, -_ ~3l~ O {$():Sl,‘fl(fCLHUUlnlille, diwiiiuipi azid lldtjif? iii tlna {liar t iiiLCl‘la l thgpnasi text. i.il:nwood and Carlisle mocks are [he nust convwn organic soils in [ngham Ct)untv. -. _... i‘r42vious Studic‘--Hydr0uuerlC soils and lowinnd éardwoodu-— Khile attenpt~ ill}; to establish area correlations btiween closely related groups of soil t;§i)cs and pressettlcwent Vegetativn, Teatch (135,l46) described several Wii>u v;:~=v= my :nfijsfiQLufi : \id ice (“licuiat1ts). Poore (1-21.) Sitlgjlg¢~stcd that with ldl'5(.? Quantities oi data, a statistical inrm oi multiwvarisnce analyses might be used in tne study of phytosociolobical methods. Rec0gnizing the multitude of variables pre- sent in the enviromnent, Cain (36) suggests that interaction would produce some problems that are unanswerable. Ashby (S) contended that the value of statistical procedures lies in analysing the distribution of individual Species within the community. Using abundancerrequency ratios Whitford (156) studied the relative dispersion of species found in quadrats, whicn were randomly located in 26 stands. To determine pattern in plant communities of several types, Kershaw (89) used cover and frequency and Creig-Smith (72,73) used differences in abundance. Tue occurrence of a particular species on a particular site, whether it be upland or lowland, is not necessarily due to any single cause but rather is a complex interaction of many factors, including all the environ- mental changes that have acted on the species during its life history. That each species in various plant conmmnities possesses specific amplitudes of tolerance, which are reflected in its abundance along an environmental grad- ient WSSfpflu d nut by Curtis (5a) and whittaker (158). The species wnich "er up these communities form a continuous series of‘mosaics mith an array 0f ecotopic overlap taking place on the slope of changing enVironment. There is definite organization to communities expressed as departures ftngm randomness which at least in the initial stages become more and care evfiideni with increasing amounts of time for community development (Kersnaw (54) l. -39- -49.... Moreover, interactions between species further increase the complexities of community patterns (Mnrgalei (109), hairston (7F), Keisnaw (9U)). Discontinuities in the environment. generally result in chzmging rates of vegetational development, as well as diiterences in composithn, and the series of nosaics along the gradient of environment appears to be continous rather than abrupt (5a). Recently, various methods have been used by ecologists to show the var- iability which occurs in the distribution of both woody and h:rbaceous ve- getation along environmental gradients (21,22,53,54,67,ll,158). The syn- ecological requirements of a numuer of trees, shrubs, and herbs in the forests of Minnesota was outlined by Bakuzis and Hansen (6). in a later study a irequency-countour scheme was used to show the edaphic and climatic complexes affecting the distribution of forest communities in the state (7). A dimensional approach can also be used in conjunction with community coefficients to show not only stand to stand relationships, but the species arrangement and reaction to vicissitudes of environment within the community complex (23,43,110). A gradient analysis of the conifer—swamps of Wisconsin based on the similarities of ground flora was constructed by Clausen (43). Bray and Curtis (23) described a method of multi-dimensional treatment in the ordination of forests in southern Wisconsin. Dimensional relationships were also used to pictorialize the ecotopic sphere of influence for a number of tree Species in southern Ontario, Minnesota, Wisconsin, and northern Michigan (Maycock and Curtis (llU) ). The use of this type of representation s*3“st the over-all influence of interaction-complexity within the environ- ."KN1t upon the species. If the ecotOpic sphere encompassing certain important tinIber species c0uld be delimited, the management problems of foresters “Knlld become less complicated. in addition, eCOIOgists are interested in defi_ning the ecotopic amplitudes or tolerances that each species possesses. N!" "‘ ‘ "JDS ftand Furvey A general reconnaissance of the lowland hardwood stands in Ingham County was started during the summer of 1960. The vegetation in representa- tive portions of many of these stands was sampled between the summer of 1900 and the fall of 1901. The sole Lion of stands for the study was made with the criteria that the stands were of natural origin, relatively undisturbed and representative of lowland-hardwoods. The location of the stands sampled is shown in Figure l. Stands which had been recently subject to disturbance from fire, grtz~ inn or extensive cutting were rejected, and only 19 of 37 stands visited met the criteria set forth for selection. Eight Siuhds varied in size from 129 to 302 acres. The remaining ll stands included in the survey were smaller, ranging from 22 to 98 acres, Jhile the total area of all stands chosen for Blimpl ing was I ,300 acres . Plot Selection A starting point for sompling each stand was randomly chosen from u =1fi31 of coordinates. Depending upon the starting point Chosen, a cor- responding random plot pattern was then selected from a group of patterns. which had been previously constructed. In the field the location of the plots was predetermined by the selected grid pattern, however, distances netween plots were adjusted according to the size of euro stand. the pattern was Pfinxnded for large stands and con racted for smnli :Ldnfii, providing for no vnual number of randomly located plots regardless oi stand size. In addition. the sum of the sample areas was the same for each stand, Since one of the major interests of the study concerned quantitative descriptions of lowlandohurkhaxnhg, it was decided that {or such pUTPOScS, ten plots (5 x 20 meters) per stand would be sufficient. Owing to the nXLTl' -;1- uni-r fifiw‘ _.- —. W Cl. lflq 5"Or (J IIDI1|11II j l J an run if: 1*; on (man mun-mun ‘ ‘ * " . - . I 2:52:35- ago—é? --- . * ; —- “k 9‘ , . ’ g s.ot‘.- n . ' T ‘ , '7?) (‘0 ‘ ' . ' V “I" "p! —~. '0 o . O ‘ . . e . K‘I j ”001...:0 330;] 1 O O.“ .3 00‘ “E E1 ré"’g':3 “'6 "i ' "~1111a°3fi$ , - L 1.. - '__ i- Tutu-3.. r w J "|.-.; ”*‘w ‘9. 3 cyan. COIOLtX Fig. l a Map of lngham county showing the surface geology after Leverett (1915) and location of the lowland hardwood stands selected for the vegetation study. A o.’ 3.. -. varig‘soility in the sizes 0t selected stands, any type of partial cruising would have resulted in varying numbers of plots per stand, and the resultant use of unwieldy conversion faetgré for the treatment of quantitative data. All stands were assigned a letteronumber designation referring to its location and predominant soil series, For example: HAB refers to Wheatfield township, Section 4, Brookston series. Sampling Methods Measurement of the number, size, and distribution of each tree species to determine the present composition of lowland-hardwood stands was accom- plished using four sampling methods. The first of these was a standard 5 x 20 meter plot for recording tree data for the calculation of frequency, den- sity, and dominance. For sampling shrub and herb data, nested 2x8 meter and “5 x 2 meter plots were placed in the southeast corner of the larger plot. In the field, plots were oriented alternately in two directions, The three other methods used in the present study (Figure 2) were non- areal point samples and included the random pairs and point—centered quarter methods (47, A8, 49) and the variable plot-radius method (13, 7Q). The point methods are brieily described as follows: 1. Random pair method. after choosing the first tree a second tree is chosen which lies outside of a 180” sector. N o kuarter method. The nearest tree in each of four quadrits is ChQSen. 3. Full Bitterlich method. All trees are selected through the use of a 3 diopter prism. The latter method was modified by actual measurement of the tree diam- eters within the sample area (103). The plot method was used as a standard for the study, and point samples were obtained concentrically trom points lficated in the geometric center of each plot. All four methods were used In only 15 of the 39 stands investigated. In the remaining stands, only the .2 f. 5.8ng no @-Bitterlich lethal @— "uarter l'iethsi @- Egandom rairs I-ethoa 23 Int" O O o o 00 @ , e of ’I a. , ' fl. I V“-“ I. III d 6;? "2“ C) o o a o O l i 2J ~~---a . Sampling methods used in lowland hardwood forest shrvey. Random pairs method. 180° sector. Quarter method. is chosen. Full Bitterlich method. Second tree is chosen outside of Nearest tree in each of four quadrants. Trees are selected with a prism. slot wethod was used. Stand Characterization - The field data for all tree species were calculated in terms of rel- Live: frequency, density, and dominance. All three oi these indiees ex- pressed on a relative basis have been described in the literature (3a, 52). For the plot nethod, the lower diameter limit for trees was one inch d.b.h. ’n a comparison of methods, four inches d.b.h. was selected as the lower diameter limit. Density per 1,9 hectares and nvrcent frequency determinations were made for seedlings and saplings less than an inch in diameter. Percent frequency on .b by 2 meter nested puxs was determined for herbs. The sum of the relative values, having a single total of 300 is referred to as the importance value. The "importance value” denotes a species' inportance in a particular stand, high Values being assigned to the dominant species (53). importance value is defined mathmatically as the sum of relative basal area, relative density, and relative frequency. Or, stated in algebraic terms. I : B {-D + F who. r e I importance value relative basal area C II relative density. Derre'tije of individuals/stand in ‘ which Species occurs. F : relative frequency: percentage of stand; in wnich Species occurs Since a few species having high importance values usually characterize stand composition, and since differences in the importance valuix of two or (:ven three dominant species may be slight, it ap)ears appropriate to charac- terize a stand on the basis of more than a single species. In describing Inv ~zands selucted {or study, ct least three Species with tne highest im— portance values were usvd to indicate dominmme within each stand. lnportance values of the tree Species in each stand were multiplied by climax adaptation numoers approoriate tor southern Michigan hardwoods. A climax-adaptation number is a subjectively determined numher which indicates the relatch position of a species in succession. The coefficients are usually delined on a scale of 0 to 10, the lowest numbers being assibned to Species wnich occur earliest in succession and often on the were extreme Sites. in this paper, the Michigan modification of the climax~adaptation coefficients recommended by Curtis and Mclntosh for Wisconsin are used (52). They are given in Taule 3. A continuum index is a number which shows relative position of a plot or stand to the average species composition of an old forest on a mesic site in the same region. It is computed from the data for all tree species by summing the products of climax-adaptation numbers X importance values. ‘he relation can be expressed mathematically as follows: (‘2 I Z (CA) (l? continuum index for a stand M wnere CI CA and I 3 clinax-adentntion number and importance values, respective‘y. for the species in the stand. Continuum indices may vary .1‘3 0 for stands on severe sites in very early successional Stabwfl to less than 3000 for old stands on mesic sites. Tue continuum indices were calCulatcd for all lowland hardwood stands in the survey (Table 10 and ll). 5:011 Nee sturements An intensive survey oi thu soils found in 15 oi the 19 SLAnds revealed the presence of ll mineral and inrte organic soil series. lasiho‘3 Climax adaptation numbers used for lowland hardwood tree I ‘ v ‘ u U C a species tound in lognam County, Michiganu -a-ww.‘ *- .—.-.—-.-- a... ~--—.~--— Sgeciv§__ Climax adaptation nunfigfs Acer sacchnrum 10 Fagus grandifolia 10 Tilia americana Hamamelis virginiana Ostrya virginiana Carpinus caroliniana Carya cordiformia Ulmus americana Acer rubrum Quercus rubra hyssa sylvatica Fraxinus americana Quercus alba Ulmus rubra Carya ovata Carya ovalis Fraxinus pennsylvanica Quercus bicolor Prunus serotina Acer saccharinum Celtis occidentalis Crataegus spp. Platanus occidentalis Sassafras alhidum Fraxinus nigra Populus deltoides Larix laricina Salix nigra Betula X purposii Populus tremloides "'\, V ‘ --———-—u-u~c~-¢.—-.—.——---~— . r afiodifled after Curtis (52). bull nrultles were ex ored my digging shallow pits with a spade, and samples from each horizon were extracted with a bucket auder. A soil series may contain tore than one type depending on the texture of the suriace soil, and among the 14 series examined, l5 soil types vere identified. Samples were taken from horizon midpoints of each soil type within a series, rebard— less ot the depth at which the horizon was located. a total of 320 samples, consisting oi tour sample replicates from five horizons in each soil type were collected. All physical determinations followed the prOceedures out- lined in the standard methods oi analysis tor torest soils by the Forest Soils Committee oi the Douglas-Fir region (b2). Only mineral soils in the stands where water tables were to be observed were completely analyzed for physical characteristics. One soil, an alpha-gley Brookston series, was ex— cluded frOm the analysis because of its small areal extent and marginalposi- tion in one of the stands. Horizon samples of the mineral soils and the mineral materials under- lying the organic soils were analyzed for total sand, silt, and clay content by'tle Bouyoucos hydrometer method. Samples were screened through a 2 mm. sieve prior to analysis. The determination of moisture equivalent was ac- complished with an International centrifuge, equipped with an autonntic con- trolling rheostat. Soil organic matter content was approximated with a py- rameter controlled Temco muffle furnace and is reported as loss on ignition. Finally, soil pH was determined by the glass electrode method, using a Beck- man (Zeronatic), pH meter. Groundwater and Precipitation Measurement A preliminary traverse of a prOposed route showed that 12 hours was the minimum time needed to nuke a complete circuit of several tentatively se- lected stands. it was suspected that some difficulty might be encountered in excavating to the true water table, due to perching ot tables in the “,4- liner textured soils. In addition, it was desirable to record water table depths within a reasonable time period to minimize observational errors in well drawdown. Owing to these factors, and because of excessive distance from Lansing, Michigan, five stands were eliminated from the ground water study. During the month of March, 1961, beiore the start of the growing season, 72 ground water wells were installed in 14 of the 19 stands selected for the vegetation study. Four wells, located at random, were drilled on each of 14 soil series using a five-foot bucket auger equipped with a 3% inch barrel. A resultant drop in water tables, as the growing seeson progressed, required the addition of five-foot extensions on the auger. This was necessary in order to reach the true water tables on beta-gley soils during the summer and fall of 1961. If a perched water table was suspected in a particular well, additional excavations were made with the auger until no further drop could be detected and the water level stabilized. It is obvious that confounding of the records on true well levels would have occurred, if the possibility of perching had not been taken into account. Many times sloughing and caving of the excavations between sucessive reading occurred, superimposing a false water level within the wells. However, the possibility of reading false well levels was minimized by re-drilling each well, each week. The detection of perched water tables was also accomplished by observ- ing whether or not lateral seepage occurred from the profile into the well excavation, as drilling proceeded to the phreatic surface. The depth at which notable seepage occurred was defined as a perched table. The largest number of wells was installed on alpha-gley soils and the organic mucks. These soils contained the greatest segment of the study area in lowland hardwoods. The location and number of wells in the stands selecteu .‘—‘_..._... m . {or ground water studies are indicated in Figure 3. Weekly measurements of the water levels, beginning two weeks before and ending three weeks after the 1961 growing season, were recorded for all wells. (Figures 19-24) The fluctuations of the water tables were measured with a nylon line, narked at one-foot intervals and attached to a lZ-ounce plumbing weight (Figure 4). A flashlight was used as an aid in determining when the weight reached the water table. leadings between the marks designated for feet were taken to the nearest inch with a ruler. Although the wells were not capped, the probability of precipitation or surface runoff influencing the water levels in the wells was slight, owing to the small diameter of the aperture and the level topOgraphy surrounding well locations. In addition, the wells were further protected from surface water influences, by elevating the soil around the aperture after each measurement. Statistics MISTIC. All statistical analyses were performed on Michigan State's tape input digital electronic computer, MISTIC. Analysis of Variance. Analyses of variance were performed to determine the significance of differences for soils and water table measurements (Appendix and Tables 13-19). All statistical determinations followed the standard methods and procedures described by Snedecor (138). Correlation Analyses. Correlation analyses were performed to show the degree of association between plot versus point samples for use in forest surveys(Tanle 8,9). Additonal analyses were performed to determine correla- tions netween Species distribution and soil characteristics, and species distribution and depth of ground water (Table 13,20,21). nun Vb ‘ l .‘ . C 03.1 11mm. ‘ V‘ :3; . , ‘. ' _, ‘3. " Vcbbomn ‘ " D 1 N 10“ I m ‘.[g Gran \ M41, 3 an. e (I Crock \ . or . Cunt \ I V111. . r“; \ . \. ; tool. , l ‘ 2 "O. cu.“ . w sausagnup i :' A I i Sou-cu mantras. ' Fig. 3 . Map of Ingham county showing the drainage system, location and number of ground-water wells in stands selected for the ground water study, .womwuam HaOm any zodop mucosa mg we: cones nous: venoum came osu .hfian cu sow: umuwu one mcwusv .cwxdu mm: szmquuozm menu oEwu «no u< .muwm many so uu>oo vascuw mo xoma ecu ouOZ .qu; venom mLu wo xuss vooaaaq sogfimnm ozu cw dams sewum>uwmno wouwz ocaouw < . q .wqm fzaas “i ll VillVf- As a means of showing the behavioral pattern of {an . v r , . . unminant sneCLes oi the study area in relation to the envxhyumental para- meters investigated, a table of community coeilicients was constructed (Table 25). The importance values of all tree Species, calculated for (I: each tand, were used in place of frequency or frequency percent advo- cated by aleason (08) and Kulczynski (95). The basic formula used by these authors may be expressed in algebraic terms as: 2Wfa+b C) n where C 3 the conmmnity coefficient K the sum of the lower importance values for each species in the stands being compared.1 a ' the sum of the importance values of the first stand selected. b = the sum of the importance values for the second stand selected to be compared to the first stand. A common early technique for representing similarities between forest standsis the use of a herbaceous indicator species (Cajander (38) ). Herbaceous plants are also used in ordination techniqinis because of their greater numbers and sensitivity to differences in environmental conditions. however, their use in the present study was undesirable from two points of view. First of all, many areas in which the Quadrats were located were devoid of, or contained only a partial herbaceous COVer. Secondly, her- baceous vegetation is often reflective of temporal changes in lowland standszauiney be indicative only of a particular year or time of year in which the study is made. Trees, on the other hand, reflect relatively long term changes in the (adaphic environment; the individual having weathered the vicissitudes of T - - Lamportance values used in place of frequency or irequency percent. environmental pressures for many years. The similarity techniques in the present study were based on stand composition of the major and minor tree species present in 19 lowland—hardwood stands. Since threermaumres rather than one measure are included in the im- portance value, it was selected as the most worthwhile indicator of communi- ty composition. Using the importance values of the tree species, the mathe- matical expression of community similarity is in algebraic terms: c = gyg x 100 but) since a+b I 600 in all cases, which will further simplify to C 3 l/BW. Expressed in terms of w : W I 3C. When all of the stands were arranged according to their continuum index placement and the table of community coefficients was constructed, it was apparent that some difficulty would be experienced in explaining its content. In addition, the evidence indicated that within the compiled table, a fur- ther separation was needed to arrange the stands in an orderly sequence for comparisons. To accomplish this, a gradient analysis procedure advocated by Bray (23) was used. an arrangement of values was compiled using two stands as reference points at Opposite ends in the segment oi the continuum covered by the study. When the community coefficients for all other Stands were arranged in des- cending order, in relation to the reference stands, a complete table could then be constructed. The rearranged table of community coefficents for the l9 stands indicated the relationship of each stand to every other stand (Table High community coeificients near the 500 value indicate stands which are Very similar in tree composition. low values indicate stands having little simil~ arity to other stands. nvsrgrq Sampling dethods The five leading dominants of the lowland hardwood forests in Ingham County, recorded by the plot methods, were American elm, red maple, green ash, swamp white oak, and silver maple. Dominance was based on the importance values of the species recorded on 150 plots or point samples in 15 stands (Tables 4, 5, b, 7). The same sequence of dominance was evident in the random pairs method. In the quarter and bull Bitterlich methods, swamp white oak the fifth leading dominant, was replaced by silver maple. For the complete survey of IS stands, American elm and red maple had the highest frequency, density, and basal area values, compared to the other Species.1 Although green ash had a greater importance value than either swamp white oak or silver maple, this could be attributed to the species being observed more frequenctly in each sample, with a greater density in the smaller size classes. gonsidering the number of Species recorded by the plot methods as a standard, three additional species were tallied by the quarter and full Biterlich methods. They were American hornbeam (Carpinus caroliniana), bur oak, and butternut. All other species tallied by the plot method were also tallied by the quarter method and the full Ritterlich method. No additional species were found using random pairs, and this method algo failed to tally red hickory (Carya ovalis), black gum, or hackberry. All additional species, or species missed by other methods in comparison to the plot method, had an importance value of two or lower, and an average basal area per acre of less than one square foot. “fi‘hr‘fi-m-w“1“,“"—-*.~'P’~*”"—-R*.~ ‘,..... --———- 1n percent. -55- ~35- TABLE.-$ A summary of forest survey tree data for 15 lowland hardwood forests of Ingham County, based on the 100 meter plot method. relative Species frequency density basal basal area importance area uvauE ted maple, black Interior View of stand M26k on poorly drained Rifle peat, continuum index IAOB. Leading dominants in importance values and in basal area cherry, and quaking aspen. c Fig. 6 -51.- uw>aum .nmm comum .mn¢~ xwvaa auscwuaoU .euw xuwaQMNm ccm .mfiane "mmum Hands :0 can mm3~m> mummu»0asfl aw mucmCaEOn mauvaug .Em0~ GNOum vmcqwuv Aduoom co nwmz vcmum we zog> uoHuuucu . m .me b u M» m 1. H n W» 3. w \. 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II II II II II . -n .. mm .. m a n- I II II II M II II II II II I- " -' m II I- 'I I- -I III In ill .1‘...‘ IIIiIIII I I.Iln| .I I.tl IlInlllnIl-IIUII Espnsu umo< maid: mwfi»fl.wtucu azumu éacu mangapmn an Arab uLfi> a>puhc 6;:21pwfim rd c..W m'nnww anmuflpoEw :caacwm ou-Q wk “wanna bnxxzuumm J mqu p : v. . P. n I .\ JUL 0 A N N “N N M . I. ac HI 9. Q 1 wt 9 H J S H 5 I71'; ‘ 1011 Cw Ucacw mmuuwfim mm;¢ LOW Quow nag 8950 “Oh .w . :IInIIn IIII {NJBQS pmmwumam . , III.|\Inn.b IInII wfimmm 30 Tugmwfiw .xdo mun—£3 953w vzm "mwhfl Hummn Ga 66” mQDAMNr wocfluhaafi Cw mUGNCflEOm. $65?qu NO 3.9.?) NOHuDuCH :Eu cmofiwaa .nmm :wmum .32 x25... 53:53:00 5.032 033.30 vocwmuv 3.30; so .uo~> vumum t 5'43‘0. ,' I' ‘ - ‘. .nmn -4 . .‘l. , l I in . n 11”. I: -55)- 2 .w: I ' '{fl‘nu- I . Snub-".3; .‘ (52.3”: ‘5' 3‘5 ILV‘N‘, ‘ I 0 ‘1"5 ~ ../\'\- Interior view of stand 3238 on poorly drained Brookston soil, continuum index 1827. Lbading dominanCS in importance values and in basal area oak, and shagbark hickory. 11 . Fig. red maple, swamp white «ensure oi expressing species dominance, Since three direct field measure- ments are usel. Nevertheless. as a measure of dominance in relation to stand-age structure, its use has a repressive effect on those species 'wnich are large in diameter, but occur at infrequent intervals with low density. The eight leading species in over-all importance values tor tne entire study area in descending order were: American elm, red maple, green ash, swamp white oak, silver maple, sugar maple, slippery elm and basswood. Within tne boundaries of the study area, 12 species occurred as a lead- ing dominant in at least one stand. American elm was the leading dominant in 35 and 40 percent or the stands, based on importance value and basal area, respectively. Green ash, swamp white oak, and sugar maple were the leading dominants in 10 percent of the stands, considering their im- portance values, and basal areas, while red maple occurred as the leading dominant in 16 percent of the stands. Silver maple, slippery elm, and tamarack each attained a leading dominant position in a single stand. On the basis of importance values, American elm was the leading or sec- ond place dominant in stands rangfiugfrom 1832 to 2035 along the continuum. in stands falling outside this range, it occurred as a leading dominant only twice: once in a stand of floodplain origin and once in a stand located on glacial till. This Species was a third place dominant in at least four stands, mostly located at the upper end of the continuum. It did not attain a dominant position in three stands found at the lower end of the continuum. Red mapiecxrurred as a leading dominant in a rather narrow range of tne continuum, within the Spectrum dominated by American elm. At the lower end of the continuum, this species led in either importance value or basal area ulclosed basin depressions. It was absent as a dominant species in stands on materials of fluvial origin. in addition, it did not attain domin- zznce in four stands at the upper end of the continuum althOUgh its presence was recorded in these stands. Leading American elm, red maple, and Interior view of stand LlOL on shallow Linwood muck continuum index 1832. dominants in importance values and in basal area green ash. Fig. 12 . 73— .3mm... “023m can .xao 333 95.3w :50 amuauucé nmwum .339 c..— vcm ”xmo 323 madam $1.58 .527; :50 amuuuflé ”mad; vacuuuoaeg 3 muSwdflEov mass»; .83 xwvau gnawudou :50.— ofimzum @25th Sheen no mm: ucmum wo 33> .3235 -74- .voozmnwn can .53 noouw :50 .3033 "m9; 132. aw van 3mm even» .392: vow :50 50:82 53.; 35:0me 3 35.553 9:233 :32 V82: 3:53:00 :53 €53 acumxooum vmaumuv 3.30m no 3: vcmum mo 33> 3“.“qu ¢H .m: -7”)- Green ash was a second or third place dominant across the upper two- thirdsrfi the continuum. Where the basal area of American elm was great, green ash although frequently encountered, contributed little to total basal area. One exception occurred in a stand found on deep muck, were the basal area of both species was approximately equal. Besides the stand just men- tioned, green ash appeared as a leading dominant in two other stands: one located on a floodplain of the Red Cedar River and another on a poorly drained glacial-till. It did not attain a dominant position in three stands in the upper part of the continuum. Swamp white oak, attained its greatest dominance in stands within the continuum Spectrum from 1540 to 1830. This species was present to some de~ gree in all but two of the stands. SUgar maple was the leading or second placed dominant in importance value within the upper third of the continuum from 2110 to 2030. In two stands within this range, it also had the highest per acre values for basal area. Basswood appeared as a weak leading domin- élnt associated with sugar maple in the upper portion of the continuum. it also appeared as a dominant Species in the continuum around a value of 1800. Slippery elm was strongly represented as a dominant commorent of the stands in the lower third of the continuum. This was.attributed to its high frequency and density, rather than basal area. Northern red oak was inter- mittently dominant throughout and appeared to increase in dominance at tne upper end of the continuum. Interior view of stand WSC on imperfectly drained Conover soil, continuum index 15 American elm, sugar maple, silver American elm, basswood, and sugar maple. Leading dominants in importance value: 2155. maple, and in basal area: .an :muwuusr .ouw mwum Human aw mucmawfiov mauvmmq .Eam uuaam> uuamuuanH a“ muawsaaov wawvmmg .ooam xmqu vmcfimuv hauuuwummEa so mad: vaMum Mo 3wfl> noduwusa was .xmo muwn3 mews» .uamme u~>fiam amUfiuma< van .mHQME uw>me wxcdag Eaanwusou .chm mado~ o“ .mE :50 50:82 can .xmo non 893.3: .393. umwam "muum admmn 5 ram .332, mountain a.“ massage—on $1."va .33 x32: Ejacwucou .50." 35mm 0x03 92:93 Suoumuaafl no 43: 2:33 wo 30.? “5:33 . S 47: u a .w:' . .. A n. . iLF. .‘b; . Q; w..........., J” " ‘r. _ m .. Ht Lt‘?t.lihll'i0ll ()t‘ Sail l l)r()pt‘rlfi(3$ The stands selected for this study occurred on a rather wide variety oi hydromorphic soils, including imperfectly and poorly drained mineral soils, alluvial soils, and organic soils. These were distinguished according to the classification pronosed for hydromorphic forest soils by Wilde (100), and by the Michigan State University Soil Science Department and F. 8. Soil Conservation Service. ‘The toposequence of drainage conditions and parent haterial relationships for mineral soils is shown in Table 10. The rela- tionships of underlying materials and thickness of the organic soils is also included. The results of the field survey and laboratory analyses showed that four of the mineral series were beta-gley soils, three of which (Con- over, locke, and Spinks series) were formed from coarse to moderately coarse textured glacial drift. The other soil was a moderately coarse textured al- luvial soil, the Ceresco series. The remaining seven mineral series were alpha-gley soils, five of which developed from glacial till of stratified sands and gravel to non-stratified fine textured looms, silt loams, or light clay loams. Included in this group were the Barry, Pewann, Brookston, Gilford, and Sebewa series. Two of the series, Sloan and Cohoctah, were fine textured alluvial soils. The three organic series were Carlisle muck, linnwood muck and Rifle peat. O Reta-Ploy Soilst: The imperfectly drained Conover, Locke, and Spinks series are nonostratified uray-Brown Podzolic soils, which formed from cal- careous parent materials of loam, sandy loam, and loamy sand respectively, and are similar in morphOIOgical drainage and color characteristics, when commared to the associated members of their respective catenas. The Ceresco series is an alluvial soil, developed from loamy fine sand. The A0 horizons consist of partially decomposed forest litter, varying itnwn one to three inches in thickness. 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U wEAwwu muaumwos ca mmanmu pmumz vwzouoa vcm wan“ mo nzamuwova: _ nnn. ..v8.~v.honao.o «one. ”“3”“. o. th.o.nhNON>n.OONN~ ‘ Hal-2x. 58>! 5.28 5-5:: :33 52. in: , Id: ‘1 In 1 q I I I I“: ' fl w u L m u z u ‘ u “an“ u I 3 cool-I... w u .utflflnmm , u A u \ m A u 7 I. 7 V W w .I. .. ; m fl u u , I ,, / . .u W \ / \/{Il\\/J \ u/ n 7 x}; lo a o -/\ ~ 1;. xx loll / I ./. .\/x . \ ./.\ J/ . o , \ /,I \ I. \0 \onllo \ I , \ \ 'II’Io/I\1\oI/\\\ [Ix r7 )- ‘0'- 1.1.5 . \ , 1/‘3 4 dB; .3. «(1“ .CN 4:; “Ivan. ("CM“) nnn...vou.~v.hOnhNO-ONONO.N.n0~N~n. o.'Nh. )3... “£80 5.0; ears aatun hggd z . «53w , O'I Osbd. o o/ \. . .2305m Omam mw 32 0;» cu mflqa caumucd sewumum cowumuwafluoua ecu yew Emprumws dacmcwm» Hood mgb tcmumv a madam» mpsumwse cw meanau uwums vacuuma vcm onuu mo mzawhxap g i. N. \ /. .\. / , \. gash on n. O on «N 4.!1 7w 0! J, (Hum) run Inn 03 «um 97. .c3ozm Oman m“ mu vzu Cu mafia qu kumooH sewumum sewumuwawumua mzu »0w EmuxOumwg dgwwcwm» 4cm” us~ .Armz ccmumv c mafimo» wpzumuoe CH wmflnmu uwumB vufiuxaa can may“ go mzmmuprczx . w; .mwh m n~ ». .. v o~ .~ c. s on n~ o. o ~ on a. ~. a on «n n. o . q~ a. n. n s~ on a. o cu - h CHOIUuUO that, asks 53:00 gt )5 : >‘I J. t‘ __.-__._4-.._ _A. A .- "“an (cc-on) .334 ’3‘ (unan) ”on on. u ”an f nn~o...vn~.~ v.honn~o.o~o~ o.~.no-~n. 0.: b.2ngzn.oo-~ uuauuo IUOIU>! Boobs 5850 bid )1 g ’1 J.‘¢ .; c a u N m ,. a l. I u 338.003 m m a v.9... a ... u \/ .. m . a \flaufl 1 / . /./ .0 m \ /. o /0. \Il \/ \o/ \ /o/ l o N M 0/0\0 0/0 O 0/0 o O ' l‘ [I \\ /o\ {0/ \ \ o\ I. O / \ a lo 0 /. .‘ . .1 .m .czoLm OmAm m“ sewumum sewumuwawowua ocu yew EmhmOumwz ~_chwmu Head mgh 32 mcu cu mm~wE ocupu caucus” mvcmumv m wEwwo» wusumwoe CH mwfinm» LmUaB nvcuxwa saw many wo mnampmopcm: gdl“ .Amomz cam mowx .CBOLm Om~¢ ma 3n c-E dxfi ceumucfl :cfiu -Mum :oflumuwmwowpa msu new Emuwoumflz #finwcdmp good mz+ .AUnHZ ccq nwwx mwcmuwv m wEwwmu upsumuge cwwwHQmu gmuw3 Uvsupwa vcm mauu mo msamevax; — n~ o. .. v on .~ 9. a on an o. o u on o. ~. n o~ - n. o . v~ a. a. a an ow n. c on «u 5.893 cal-up! 5.38 5.333. :33 52. I}. :- 43: i m u. m u u ; lame J m .38!§8 m . 5---. . v" I u .6 m fi ‘ . m . m a W Va -315... A in vita; of“ tin} ccnxtori citrus! {OlUB ioz‘ the: Mean', Slxran, avid fpvcarx)re Creek watersheds, and an evaluation of-the hydrographs revealed that the wa~ ter tables under investigation were being influenced by at least six differ~ ent recharge patterns. . The relative position of the water table for any period of the year in these stands is obviously being governed by the amount of runcfif and uncon- fined seepage which occurs. This in turn is determined by contour elevations, drainage outlets, permeability of the lower subsoils, glacial drift mantle, and underlying rock formations. In this respect it was practically impossi- ble to evaluate losses due to these factors, since the discharge patterns in these stands were not monitored hydrOIOgically. The hydrographs for the cur- rent year, however, can be consider representative of water table cond- tions and changes for lowland soils which have existed for a considerable period of time. This in itself is evident in the age of the stands located on such areas. Spring water tables were at the highest level in all soils during the second week of the study. Saturated soil conditions immediately after the spring thaw, the lack of transpirational draft, and the amount of rainfall which occurred prior to this time were contributing factors producing high water table regimes. a general lowering of the water tables ocCurred in all stands, except those located on alluvial soils, during the period from April 29 to August l9. Hater tables in the alluvial alpha and beta-gley soils were being influenced during this period by the rise and fall of the Red Fedar River, which drains all basins in which the stands are located. Ninor fluctuations (1-10 inches) caused by precipitation from Class 2 thifurm storms (Smith and Crabb (537) ), affected :he position of spring and . _ . . . ‘ ‘ / Steamer water tables in all stands during the same period. Several (lass 4 advancedsmorms occurred during the month of June with little effect on Host water tables, except those of the poorly or very poorly drained Sehewa, Gil- ford, Brookston, and Pewamo soils (stands WAS and WEB). Most of these storms produced a total rainfall of less than one inch, which apparently was not sufficient to satisfy the total interception loss plus existing soil nois~ ture deficits in the profiles. laximum depths of water tables occurred in most stands during the last two weeks of July and the first two weeks of August. Mean depths recorded were greatest in the Conover, Locke, and Spinks soils (stands AlQC, Ml9L and M198). Maximum fluctuations in water table levels occurred in the lat- ter stands thronght the growing season. However, a comparable amount of fluctuation was observed in the Sebewa and Gilford soils of stand WAS. The Ceresco soils, one of the alluvial beta-gleys, had a shallower depth to the mean water table than several of the alpha-gley soils. Water tables thronghout the growing season fluctuated least and were most frequently within ten (10) inches of the surface in organic soils. This was especially true for Linnwood muck (stand LlUL) and Carlisle muck. The other Linnwood muck (stand AlaL) and the Riile peat, however, had deeper water tables on the average than the Pewamo soils, Gilford soils (stand L106), and the alluvial Sloan soils. Even though the growing season during which the study was made was one of the driest in 31 years (Figure 25), ending with a minus 3.73 inrhes de— ficit in rainfall, it appears that some Class 6 and 4 advanced storms were of sufficient intensity and duration to produce some recharge of the . . l 1 . . phreatlc water tanles. Storms or these two classes, normally producxng ~.—...-*v-n- g-..- see Idble 1, total departure from normal rainfall. M-“ w.~~ --.~-~ . -w-o— -‘~——.— _\ *l'.‘{l" .manwunfis Hausa uncontae Sen“ oofiuon oxu you once -xo .GOmmom wca3ouw «sauce osu manuso 30M ouoowu 3oc m vocmw_nmumo HooB menu aw H~>od sauna flood meg umnu muoz ca Haw; cofiuo>uomno mace oLu new mam>o~ nouo3 come we ensues ummmumd c4 okwwhfiu ANHUNHW $.ka wCHqu— CLU 5,35 > J-F Ogl a: 0- Coo M800 nuns-um “1" an M “”1 III“ ' «i. i r- 21*". 1"". .* '3‘. i- Tiara i t“ g, 3 n ‘x I int-3;}. gnu" 3.5.? Terms-t: 1 .-. , wé ii ;ornd;uy? a lhlfll«ll rwu marge: of ennlsurflhrc warwxrs ii; hrdldzmllhlfdkhnhj stanuls. Regardless of whether 0. not the soil was anzfltdn-bmy' or bvta“gl€?, recharge of the ohreatic water tables occurred quite soon; a matter of sev- eral hours alter a convectional storm of these intensities. The duration of storms of these classes must necessarily be longer than an hour. Such rapid recharging of the water tables could not be attributed to flow of water into the well aperture, since proper precautions were taken to prevent errors of this type, in addition, a vist to the well.locathms inmmdiately after several storms of these classes did not show disturbances around the well Open- ings due. to runoff, . It was surmised that this situation was not brought about by saturation of the soil profiles immediately above the water tables at any given point, since some of the tables were 80 to 100 inches deep at the time that rapid recharge took place. On SOme of the poorly or very poorly drained sites di- rect recharge was probable. However, the evidence in the present study. gathered inmn numerous observations, indicates that recharge of the phreatic surface may result from gradient flowage. This would be dependent upon the slope of the underlying strata and the permeability of these materials. The upward movement of water tables in these low areas, however, does not neces- sarily mean recharge from gradient flowage. li the water table is in a steady state relationship with gradient flow from above and local evapotran5* piration, any reduction in the intensity of evapotranspiration (in situ) would reduce the drain on the local ground water. Further, a reduction of evapotranspiration upwatersned would increase the gradient flow and a rise in water table level would occur. A strong recharge pattern began in the third week of August, 1961, after a Class 6 advanced storm which registered 3.56 inches of rainfall at one of the recording stations. The total rainfall from this storm at the ”zone M L Q ~ p , - H: l 1‘. 3 i ‘ ‘ K . J), 3.31“! .‘ , . . '-l'..ul'.9L I t .i .'L n 9" ' V ' ' ‘1’ l N 'b‘t‘ ' .' : " '7' *- "u ' ' ’.V‘ 7‘ ; ')r' ‘I ' ' 'I‘QQ.’ -‘ )1“ i")r t". “3"' ’1; '(101 I‘ . t. 1‘s ;’r )’.!l'.() "Jill -1 {v.t)I-\ “113lt(‘| '1 . L L . 1.. Alt S St“ L I'. It] 1.“. ._ L, .)(_l ..1 .k ‘ :16 ' ,Igl. . i; . _ I the growing season. Two exceptions were noted in the lovke and bpinks soils. 'Yhe:nx ism) soi ls, (hirint: the. thitnl wetd< of ;\ugn:st, iati‘wntter trfiiltu; dTl to DJ? inches deep at. rm! time. this storm (teamed. After this period, their tater tables continued to drop until the third week of Uctoher when recharge to tan surface Degao. At the termination of the study on December 3, on];' the water tables in mucks and alpha-gley soils of stands VZOC, LlOL, LlUC, and N49 Were fully re~ charged. The hrookston soils (stand W53) and the Locke and Spinks soils (stand M19L and M193}, were considerably different, being 26, 35, and 67 inches reSpectively below the measurements taken the first week of the grow- ing season. All other water tables were within three to sixteen inches of the level observed when the study was first initiated. fizzynni ate? rux' onciéws UvV(lcwvm?nt Depression storage irnm precipitation, or a rise in the ground water table from beneath during the winter season and early spring, usually reaches a sufficient depth to preclude the survival of all young plant growth. This condition was evident not only on floodplain soils, but also on several of the lacustrine soils. Therefore, the current vegetation of several lowland stands was subjected to a natural annual disturbance. Only during thoseswmams which would be fairly but not uncommonly dry. could woody and herbaceous species develop sufficient root systems and shoot growth to survive the effects of seasonal flooding. Of the dominant tree species which appeared throughout the lowland stands in the county only four: siver maple, red maple, sugar maple, and basswood showed a tendency to be associated with rapid changes in water table levels. silver Maple.--The occurrence of silver maple along small streams and on soils proximate to major streams has been reported extensively (124,150,163¥. Jr: reproduction of this snccina is very pronOUnced on bare mineral soil alongside such waterways (law). this condition was apparent on the flood- plain soils of Ingham County, nhrrc the reproduction was abundant but a rzcgated into groups, Littl: cnwpetition was present from other plants, urn the distributional pattern of the reproduction appeared to be associated v‘Lh overflow depth from thc it; Cedar River (Flaure 26). In reviewing the apportio want of basal area for the canopy members WLthn all stands. it would appear that a relationship existed between the nrvwlopment of silver maple ant fhc rapidity with which drainage occurs. This was apparent not only for floodplain soils but also lacustrine soils. Amorg f? odplain soils all, except the bloan soils, exhibited the maximum variation -l03- ~104- .Luawn mwcu Ou 30kuw>o noumeumn mo muomwww ozu m>w>unm manmnoum flaw: uwufi wzu um wcadvmmm Hawu wnh .wonucH ma we :uawn w cu msucce paucw3 wzu wcwuav vmumc acacnfi mfimuoanfioo mn ~HH3 uoam kwfisufluuma was? .mwzucw nu mm: ucw6aodw>wv wcwavuwm wo wwwum was» um :uamv wanna uMumz cwoe wLH .on_z uzmum we Emoa mucuw smuuonoo mfiu co mwcwfivawm Qmmmwmmmwwmm uwoHHw mo uwama < WV I 5?”, 7‘ —- . om .m: \a—J in water (Hole depth and 1' inclination. The: (ieVeZogtmt-nt of silver maple on Sebewa sandy loam and those Linnwood mucks of similar hydrologic character- istics also support this contention. Egg;§galg.--Of interest was the conspicuous absence of all but seedlings of red maple on the fast draining alluvial soils. The lack of this spec- ies on soils of floodplain origin could not be attributed to variations in soil texture. It appeared that the occurrence of this species was somewhat sensitive to alluvial sites which drain rapidly after flooding. American Basswood.~-American basswood appeared to possess a wider eco- logical amplitude than sugar maple on lacustrine soils where the drainage was poor. Possibly, catenal positions of the soils on which it was found was more important in influencing the presence of basswood than their textural attributes. The presence of large individual specimens of basswood in the lowlands of the county could he a stand-age-structure relationship associated with the interval in time since these stands first became established. imit rs! » rut iv“: 3 '2 .n: .r? " lurk it i 3-. 3. pm: ‘l..31"i-,.o.)»'.21 . 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A“ uu uu uu uu uu u- u- uu m ofi uu nhmzuc uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu mgtwomnegpu m:_:acm uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu “Mmzmunm x aflzumn uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu spa“: xwficw uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu wcfiuflpmfl xfiwfig uu uu uu uu uu d uu uu uu uu uu uu uu uu uu uu uu uu uu mucucufiwu mrflflatm uu uu uu uu uu uu uu m uu uu uu uu uu uu uu uu uu uu uu mpwmc mn:quum uu uu uu uu uu uu uu uu uu uu uu uu uu NH uu uu uu uu uu CSLNLMN mmuumuwrm uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu wwamucckuuc muscumum uu uu uu uu uu uu uu uu uu uu uu uu uu N uu uu uu uu uu .aam mzmmcwcpu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu mfifimu:ovwouc wwuduu uu uu m H uu uu uu AA uu m uu uu uu uu _m mm uu uu uu Encapwauumn umu4 uu uu m m uu uu uu A" uu uu uu uu uu 0H uu uu m uu DH mzflu0uow wscspu -- q m .. Rm c mp HM .. m NM NH m -- -- -- -- m -- uoHcUflo muopmsc o m m cm ma uu ma uu ON on ma Om mfl m mm mm uu uu uu muM5m>Hmmccoa mDquwkm uu uu uu uu uu uu uu uu uu u uu uu q uu uu .u uu uu uu madm>o QALQU uu uu uu uu uu uu uu uu uu uu o uu NH m uu uu uu m uu wum>o crpmu uu uu MM mm uu om uu uu m ma NH uu Mm uu m m m mg uu cunuL mLEH; uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu dem msuxmsc uu uu uu uu uu q uu uu uu uu uu uu uu m uu uu uu uu uu msxuwpmam mscwxmum uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu mufium>~ym cmmmz m uu uu uu uu q uu uu uu uu uu uu uu mm uu uu uu uu uu cups» mSULQDU m uu uu m NH m uu uu On Q ma uu uu m m uu mm Hm uu Eupnap Lmu< uu uu m uu mm c uu uu OH - Nd mm Q Ma uu uu uu uu uu mcmuwumFm mDEH: o q uu m uu uu uu uu uu uu uu uu uu uu o uu uu uu uu mepCVfinpoo m>umu m uu uu ofi uu uu mm HM uu uu uu uu q uu uu uu uu uu uu ccchwaopmu muzwmpmu m m uu uu uu o as uu uu uu mH uu m uu uu uu uu uu uu msmmcwmuw> mxpumc o uu uu m uu uu m uu uu . uu uu q uu uu uu uu ma uu macawumfim mfldwe om < o m uu uu uu uu uu uu uu uu uu uu uu uu uu uu uu mwfiomwvcmum mama: N no mm OH uu an u- my uu u- u- uu N. uu -u uu u- uu uu Enpwguumm “moa uuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuuLmn&DCuuuuuuuuuuuuuuuuuuuuuuuuuuu H H V V 1. ”v .1 S A H H H H on .J .1: 9 .13! uluul T. T. M 1. fly 1. I. M I. 1. 79 Z. M I“ 7. 7t 7“ fl .5 3 ,6 ,b C. V. .t V 0 C, b. .0 .£ 0 7 9. l .c 9 7 Q U u- L. Wu Q“ fly fly 1 H D 8 1. w H“ 3 H U 3 8 H S H m ou.wuwcw q Ou ~.H meMAamm CO m.pmuoEwHU CH mwso:w woman Aucsoo EmzwcH Cw mncwum UOOSvumcuvcwHSOH ©~ cw zuwmcmb wmwuoam .m~ u.ugr>. itzr‘lm‘fil'h \“"~‘“jy blue violet. black sowieroor, eloqueiozl. wild ginger, horse-nalm. wild leek, white a\eos, blood~ root, tin) snikcnarwbs. true {Uld false Stdivmwt's seals, l)lUP~SLPn tuwldenrod. bollwort‘ blue cohosh, and lopsced. shrubs and vines of. rather common ()(‘CHI'VC~:I,‘(.’ on these soils included; strawberry hush, common elder, shadnusn, woodbine, poison ivy, the currants and gooseberries, green osier, sweet Viburnum Viburnum Lentqio), and button-bush. A complete list of the lesser species feund in lowland~hardwood stands ir'zdiczntirn; changes in species importance along, :1 gradient of changing mois- ture conditions is shown in Table 13. -1“)- ¢ 0, ’1'3‘~.?:.I..Ii.-!3 A species list of the ground flora ccmzmon to the lowland hardwood stands of Ingham County showing the dpuree of importance attained 0y each species. m v—u-Qn- ~&~*-~-_Mh.--.—- .. — -._---. ,3. ~-~—m--'~ ~ . . a Soils and nwlsttxfienrcfipqie Species _£gfv we: _ he: hat-mesic Organic Aipha-glcy beta-gley sol;§“__ ”‘351115 533ils o-ooa-oc-Q-a.acan-conntXIn't)(‘-'r‘coponuow-uoua-nun-nu---- Actaea pachypnda - 1 2 Actaea rubra - 0 - Alllum canadense - 0 - Allium tricoccum * 1 3 Anennne canadensis - 2 - Ara]ia racewmsa - O - Arisaema atrorubens ~ I 1 easarum ca nadcnse - A 3 _fisclepias incarnata — 3 - inter sngittifolius w 3 . - imahmeria cylindrica : L3 3 Cardamine pratensis 2 ~ - Caulophyllum thalictroides - - 3 (To! 1 Lnsonia canadnns i s - - 3 Cicuta nmculata - 3 ° CryptOtacnia canadcnsis - ’5 2 Erythronium americanum - - S Lnlium concinnum - 2 l .55 lium triflurum u 7’4 2 Cofanium macularum - 3 A :JwHW canfldense - 2 3 Ulyccria stridta i 3 - féygrtiix In:tu1e1 ‘ ? l .‘2rgn’sti91u: capensis '7' l - h nations pallida S 2 - light" 1 ia ca I‘dina I is - '1) '- lyczuan Jum’rica!njs - I - Lysimacnia thyrsiflora « 3 - Lyfiimachia Ciliata ~ 9 - I.y:tiwuxcluia qLuadx*iik)l_ia 2 - firthrum salicaria - 5 - '4itcila diphylla t 3 - deturtium offiicinalv 9 ~ - ; 3.24:0 2‘ 1112.4 (71.3 ytnni , 7.3 4 '3.):'i:1x)r"-:1i 2:11 lungi s Lvl is - 3 l i." ii if yin": 101151113 t y] is -. Q 2 ¥%}nophyllum peltatum - Z 3 leo 17.51.12 . ‘ 1.3 . . continued Potentilla palustris Potentilla recta Polygonum natans Polygonatum caniculatum Ranunculus recurvatus Ranunculus scptentrionalis Sanguinar 1'3 canadcns is Sanicula gregarla Sanicula narilandica Smilacina racemosa Smilacina stellata Sarracenia purpurea Solidago cacsia Sodidago graminifolia Thalictrum dasycarpum Thalictrum dioicum Crtica gracilis Urticastrum divaricatum Uvularia grandiflora Viola sororia Viola spp. Pteridoghytes Adiantum pedatum Dryopteris spinulosa Equisetum Hymale Onoclea sensibilis Osmunda cinnamomea Osmunda regalis N I L.) b. meO ubL‘lU-J' wr— WINb‘b—‘PQWNNI NNN-C‘L‘uml tNl-‘lb. 9.95.3.9) walwle‘L‘NI-‘I 0)le ?arranged according to a gradient of decreasing moisture. Dnumbers refer to the following scale of occurrence. O-ptesent l~scarce 2~unconmxn1 3-1nfroquent 3~common 3 ~vcry common Biotic Factors and Their Effects on Community Structure Evidence of the destruction of natural Vegetation by biotic influences is visible thoughout the lake States region. Ravages caused by insects, diseases, fires, animals, and especially man has seriously altered natural succession in many types of vegetation. These changes, principally by nan, were most evident during the early days of logging. Fire lnfluences.~-The role of fire as a factor in the ecology of low- land hardwood forests in lngham County has been of little consequence in de- termining its vegetation. Although extensive fires have played an important part in forest succession, evident in the record for the state's fire history, fires of natural origin in this county have been more or less confined to upland sites. Lowland hardwood forests probably escaped the clear cut- ting practices that prevailed in the coniferous and hardwood forests of up- land sites and the ensuing fires that took place after such devastation. On occasion, fires have occurred on the Sparsely forested deep peat soils of the county during periods of extremly dry weather. For the most part, however, the destruction of these primarily deciduous lowland forests by fire would indeed be a rare event since the ground water is so close to the soil surface. Although the soils of most'lowland stands are not permanently saturated with ground water, the extent of the capillary fringe above the phreatic surface serves to keep nest lowlands from reaching a highly combustible state. Moreover, periodic saturation of the soil profiles by convectional storms and resultant ground water fluctuations occurs often enough and with sufficient magnitude to effectively reduce the likelihood of serious fires. During the ten-month period that the ground water study was being con- ducted, the route of the survey was tantamount to a series of random road counts used often in wildlife studies. Although debris burning on land -11:.. ‘ . . ‘.... .. .,- n3. . u .,r, .. .'.- ' l'.~~ .-~ -‘>.--=-~v ‘ -'.)('1.l.l’. CiR‘J'I. “i.“ with .‘wt't :1 I. r“. L' ill il',"{.1t,’.ll I y!) {A}: «I. 5;. 1 4:35.“? lv'l lbl lIC \‘at‘b 1):; at" vexl fr! tile lovilazuj inlrihwinri l n.d t_ypt‘ for“ tin: exitir13 ywulr. From the Va 1 umi msus litt‘rz! I u‘re concern i D}; the ef tects 02‘ lorest if ires on soils and living orgsris s, very few Studies have reviewed the effects of post-fire succession on lowland soils (2,3l,32,108,l>01. The maiority of references on this subiect have discussed the use 01" zire as a Silvi- cultural tool in the management of coniferous forest stands. The practice of setting fires by the indigenous ‘ribes of lndians tor purposes of come nonication or hunting game was never mentioned in the early history of lngham County (63). It is not here inferred, however, that fires of low intensity have never ownnred- On fallow and ungrazed land, where herbace- ous vegetation continually adds to the bulk of potential fuel, the probability of fires whatever the cause is enhanced (Figure 27). Buell and Borman (31) found that paper birch stands sometimes directly resced to a mesic species such as basswood. Quaking aspen, black cherry, and northern prickly ash (Zanthogylum americanum), however, are the most likely species to reseed.lowland soils in Ingham County following fires. (Figure 28). Quaking aSpen, however, is not necessarily linked to post-fire sucession. In thissmudylit was most prevalent on Rifle pears, which showed no evidence of a prior fire history. Likewise, its presence on mineral soils in the study area appeared to be correlated with natural disturbance factors, expecially windthrow of other species. Eindthrow, Rooting Systems and Community Structure.--lhe importance of root systems and their development, as a prime factor aiiecting Species density and eventually stand composition in lowland forests,has been largely ignored in past ecological studies. The tremendous arount of labor and time involved in the excavation of large root systems, not to mention the difficulty of identification while keeping smaller roots intact, -116- .uowwuam HHOm unu weacwemxo scum uaouwmam was vouusuuo wcw>~n whoa mica we oucmoa>o oz vcmfi owned uu>o mouau we uoauuunuuo use cu mucououov ogumauummw mm uom mmocuw .cOwuwvvm aH .uoasovcwH 0;» kn vouwv on has whom ecu uzn .oucouusuuo meow mo wooua m>ww mono uoc nuuonno dune ads scam .xfimaofi>uuq name» coo uuaw zuwmcmucg 3o~ m an amuse no: man» menu umnu quw «so cu macawumou muss mu>fiw umum whom < - amok“ .nm .wam .mequuonm m=Ow>oun onu mp kumowucw muum may swaounu uawzm saga: .uuww on» son uuu smug swam mane . «EnamUfiuoEd Hmmxmmmmmmm v 5mm zdxuwha :uozuucc vamam umuouowa mach»: v xmo was we mocmmuum us» ouoz .m¢; vcmum cu unwounvm Emo~ maunmm mmfiwnusafim can :0 Ammmmmwmmmmw mmmmuva cmamm wawxmav we v:Mum wcsox m .ww .MHm . “‘7 \ 4 . - fi¢.fi ‘ ~5 , . -Il7- fig . a“) 1 ya 1 constitutes a umjor problem. The majority of root system stratiiication and development studies on hydromorphic soils has been devoted to seedlings or shrublike plants (10,12,56,152). Nevertheless, a reasonable degree of confidence on probable root system extension may be established by observ- ing the rooting habit on windthrown trees and trees with partially exposed root systems. While the soils of lacustrine forests in this county are not affected by the homogenizing effects of floodwaters as in floodplain stands, windi. throw as a mortality factor serves to influence stand-age-structure during later stages of succession. This condition is most evident in stands found on organic soils. The loose consolidation of granular or peaty materials was not only subject to being blown away from the forest floor by unusually high winds, but a col- lapse of the organic matter has occurred with tthassage of time (Figure 29). The extensive system of storm drainage outlets installed in Ingham County over the past 20 years may be partially responsible for this collapse. Alternate wetting and drying within the soil profiles, from precipitation and fluctuating water tables, however, also produces this result. Event- ually the supporting medium for larger trees is weakened. Moreover, a periodic re-assortment and deposition of lighter materials caused by wind 1 . and wave action occurs when the stands are flooded. In many cases, the major portion of the anchor roots of dominant species arerartially or almost completely exposed. Root growth, however, probably keeps pace unless decline of the watertablc is rapid., During the earlier stages of succession, cottonwood plays an important part in lowland stands. Because of its perpendicular taproot or heartroot develOpment, this species is better able to gain a foothold on sites having substantial fluctuation in ground water levels. On the Otherlunui. -119— .mucmcgeov wcfivmog ago we muwasuuos on» cu wouusn swuucoU acuumw anuuozouoa a ma mwuwm owunu co Bounuvcwz .xowmu auoEmuam mo mumum3voo~u any Eouu wcwzmwa uvauuom vcm maswwooafi wcauuoamsm we» we owned nHoo cu one .vomoaxa haouufimEOo umoeam no muoou nonucm mo Ewunhm any .oo~> endow wo x055 odmufiumo mean can so voumuoHA53awumsuumm umu~am < .oN .wgm 'L‘l-lvit'I’ willow ;"‘. :jxdlllxti-zf in [110.179 i..."T‘-Y",‘.'-.‘-Iir)ndl. {lino-ms k’lllfl‘il ra.-.l‘.:-.'lif‘. Urnt‘a» or laws permanent Ev we! . l’cc.‘<::;~‘¢: of the plate-like de_-:*J'v.'lo;uw)nt of tn.» root <:\i> 1 'Hi: i_ti T‘lir‘ ii! t 1‘31‘ siz>(*z:f t‘>;, L31 t iii-x'i x1); () f tlzti l i i‘r {Ill}; :‘f):>t si}'s l“l“ !: CUWHQH. Black willow is soon repiuced by cottonuood, espteiallv after a srrienscsf drooping: seasons. "fire 01;: te-sha uni «levclomtnrnt of the root systwin characteristic of certain lowland tree species, such as red and silver maple, also contributes to the seriousness of windihrow on organic as well as mineral soils. The system of llcart'root development in sx-Jezno white oak and American elm is somewhat flattened on soils with shallow water tables. Windthrow does not seem to be of serious consequence, however, inhibiting the eventual dominance of these two Species. American elm seems to possess great stability on such sites, due to its widespread and fluted rooting habit as shown in Figure 30. The rooting system of green ash appears to be intermediate between the strongly flattened heartroots of the maples and the deeper penetrating sys-° tems of other species such as elms and oaks. The mortality of this Species in lowland hardwood stands was quite noticeable although this condition could not altogether be attributed to windthrow. Green ash, a prolific seeder, easily becomes established in small openings created by the windfall of other species. he resultant closure of adjoining canopy species around such openings acts as the vain factor in later eliminating this SPPClQS iron? ens r S l ands . A summary or the Vthflllty for the major and minor tree species recorded by the plot method is shown in Table 1Q. As indicated in too table, green ash is quite frequently represented. however, KDFt of the vvrtality [hr this species ownnruu; in the {our to eight inch diameter classes was related to canopy closure. he mortality values for quaking GSan and blafk willow were also related to stand closure. hindthrow, .aomdom you—:3 on» wcwuov .Eoumxm goon 93 mo «man. on... wcaocaouusm 33.359: #3055 on... no canoe gm...» on... 353 3 won 3.3353 mu mung. .muoou uonuam an? .«o ucofiowemuum “.333 93 09302 $33 bemum uo Eng 9533 on... no wcuguwamsmuwuuam «32:: Ego cactus; 5. 1|: '0‘, '1 . a . .uufll‘” .ictllaj .90 x. \ I: » . \ I I l / ‘| \ a ,1 y . t ‘9 /‘ ‘.. -l';‘l- .OM .wot If) ‘ ~11 mumbaac coHumunmomnxmewoo uo umouo wcmocmumoo so omucmubm mowomom ansoucoo :w swoon wowocoomm so bowzohoo mocnum II II N NM 0 m w mg m H No II m II N m In H II among 3 In In In II II II II II II II II II In In In In In In In «Handbag x ofivoom II II II In II II II II II In II II II II II II In # In mono: xflfimm In In II II In II II II .H II II II In II II II II II II mcfiofiscm swung II II II II In II II II II II II II II II II II II II II mvoHOuflo: wZMDOOQ II II II II II II N II II II II II II II II II II II In oowwc mocwxo»; II II II II II II II II II II II II II II II II II II II Ezoweflm mouwmwmmu .0. I- I- I... I.‘ ‘U. 'I. '- " -' '- I- '- " 'al In- 'lu -' 'I Oanmm WDUPWMUVOWHIHHV '. ’I- 'I o.' It- -' -.I II- 'C " " '- " 'II " 'II ‘lu " a " m...~—mnuaalhtwuoo "MWUMMWU -- -‘ -3. -- '- I- '. -l. -- -' " I- '- " H In- " -nl .' Pia—A—CIHINMWMMUUNNMW “$00an II In ~ II II A II N II II II II II II II m II II II wowsm>dkwccmc macaxokm In In II II II II II II II II II In In II In In II II II mM~m>o ozoou In II II II II II II II II II II II m II In In II II In mw~m>o o%boo II II II II In In II II In II. II II II II II II II In In muons mafia; In In In In In In II II II II In II II II II .II II II II moan mooposu In II II II In II II II II II II II II In II II II II In occowpoEw nonwxmuu II II II II II II II II II II II II II In In In II II II mowuc>~>m mmmxz II II a mg a q m m~ H II m II II II NM II II II II mszfiLch mafia; .5 H H. ...II ...I HI ....I CI. K... .IH ..../. H...” ..\. 0 «4 TI 1.. MI. In .I M l .l c... 7.. uh I. In. Z .d ”A .l e . 1h. (U r... I../ tn. C. .7 nIIJ. 7v AU .9. n1 .L 7.‘ 0.. bl U U To»; I. .vQC n. l .8 .1. 3 1.. 9.. ”l T. h 3 u. H ,I. 5 .3. S H D. o I «3.5.5333 85%.: cm mosey.” UCOEmouazIocitsofi mo CH Cumin ardent”. .Au:mopcfi 9.3593,.“ . a; I.,...t?.....3., novevur, W35 the major factor cansing mortality tor the maples and biick cherry, cspEClfllly on organic soils. EKEQQEQfliC_£Qf1”?QFQ§ and Community Structure.~- fit all the species re- --.—. -..--—-'-‘A .— corded in the study, American elm had the highest nortality rate. Over 95 percent of the nortality for this important lowland hardwood appeared to be attributable to the Dutch elm fungus (Cetatostor Ila ulmi (Schwartz) Bus- iman). The Spores of this fungus are carried to the trees by the elm bark beetles (hyluEEOpinE§_rgfipe§ (Eichh.) and (Scolytus multistriatus Farsh.). It was not unUSual to find some of the finest, largest Specimens oi American elm suffering from the ravages of this disease. No size class appeared to be immune, but total mortality was more evident in clustered groups of trees smaller than ten inches in diameter. The number of dead or dying trees appeared to be highest along the periphery of most stands on organic as well as mineral soils. Moreover, the incidence of attack was practically complete where American elm had attained dominance at the edges of stands. Within most stands, however, the number of trees being attacked was low, and the prevalence of the disease seemed to increase with a corresponding decrease in stand density. Isolated groups and individual trees in open fields or along tence rows were almost always dead or in the process of dying at the time of the survey. Since no cheap effective control has as yet nc:n developed to combat this disease in natural stands, it would appear that an extremely high loss of flmerican elm is to be fiflthlpfltEd in the losland hardwood land type. ‘ ~linuugh American elm has low comuercial value at the present time, the I‘. importance of this species as itand component at considerable worth cannot bi ignored. Should the disease continue to extend its influence in geomet- ch prOportions, American elm may soon become a uszher infrequent species. 'l‘rze van—yam I‘m. (fire-3131:? by {no loss 01' this §,:M_L‘.l(‘.'n 3t t‘m.‘ present Haw- are rmt-‘t likely to be illled by increasing mmunts of green ash and S‘rldhll" white oak on Innera: sci 1.5;. Un organic soils and in the latter stages 0!? sue- tesslon rrd maple is the most likvlv Species to fill the void, replnCiLg inward ash and tt)fuhh9 extant.swunm)1~nite oak, 'Fhe loss ;n_xnmcricarat-hm on wet-mesic sites, where it decreases in frequency as well as density, is not as great a loss to the character of lowland stands as on the very wet to wet areas where it is strongly dominant. Statistics Correlations between sampling methods.-- in order to determine the relative reliability of the various sampling methods, simple correlations were cal- culated between three parameters (basal area, density, and frequency) as determined from each of the sampling methods. These calculations involved all possible combinations anong the 12 parameter-method measurements. The means for 23 species were used as items. The correlation coefficients (3 r) are presented in Table 15. The coeffi- cients of determination (3 r2), which provide better estimates of the infor- mation provided by one method relative to another, are presented in Table 16. The smallest coefficient of determination between any two methods used to determine relative basal area of different species was.95. This was for the comparison between the random pairs and the full Bitterlich methods. In effect, such a value means that the random pairs method gave 95 percent as much information about the Bitterlich-determined basal areas as did the Bitterlich method. The coefficients of determination were all .97 when basal areas determined by the plot method were compared with those determined by any of the other three methods. Similarly . any of the four sampling methods gave 96 to 98 percent as much information about relative density of the 23 species as did the best method. And each method gave 95 to 98 percent as much information about relative frequency as did sampling of ten 5 x 20-meter plots in each stand. The coefficients of determination between relative density and relative frequency were also high, all being greater than .92. Thus, by whatever metho od determined, the data on density gave 92 to 99 percent as much information about frequency as did a direct determination of frequency. The coefficients of determination between basal area and the other two parameters were lower —- between .84 and .93 for density and between .87 and .92 for frequency. ~125- .EOmeuw mo mmwuwwv AN £uw3 ~m>v~ #0. egg um mucmowmwcwwm haw vwnmm: qu. nu mo m3~m> ms» cmsu umummuw mum: mcoHumdouuou Had m mm. mo, mm. mm. mo. mm. mm. qo. m¢. om. ¢m. nosuws uoaa : mm. mm. om. Na. Na. Rm. ¢m. mm. om. no. vocume nowfiumuufim : mm. mm. moo mm. mm. mm. mm. om. mm. vozumE umuumsv : mm. “m. mm. mo. no. «m. cm. «a. venues muama eoncqu mucozcmum mm. mm. mm. qm. qm. om. No. tongue “can : mm. mm. no. mm. mm. qo. conqu zuwauouuwm : mm. mm. ¢m. ca. No. venues umuumav : mm. mm" om. cm. nocume magma Eowcmu xuwmcua mm. mm. mm. nonume uofia : : ow. No. segues zuwflumuudm : : mm. vozums uvupmsv wvum Hmm1m m .m 1 MW nu Lu 1 .d u .D 1 my 3 m w o n. m m m. n. m w 2 3 J D. 3 3 l D. 3 .3 J 0. 1L 1 3 O a 14 O .6 14 0 . .l. 8 m J a w J 8 m m. J .u H” J .L J u. 9 3 .w n... .m M. u. I. u. I. s m m Illtvozuwfil‘ll IIII©O£HOEIIOI IIIIOOCUQEIIOO mu omunmmoE hp kusmmoe mn4nmw3m¢wfi mm.mu:rmmv»m mm muwmcmm mm «mum fimmmm voxumz umumsdpmm m.mcozqu wCWHQEmm psow xn amusmmme mm muwmswv mam .mucmsvmuw .mmum flwmmn .mmwuwam cow3umn mcoflumfiwpuoo . m~ I.MJmna 00. mm. mm. co. co. co. mm. mm. am. Na. mm. vozume uoaa mo. mo. No. do. ¢m. qm. mm. um. no. em. cozume goddpmuuflm : wa.. ma. co. mo. no. um. um. No. hm. wosums “muumsc : am. am. om. om.. om. Km. No. mm. vocuwe mufima sowcmu zocmscmub mm. um. No. mm. mm. #0. mm. sesame uofia : mm. oo. oo. no. mm. mm. noguwe nuflfipmuufim : mm. om. mm. No. «m. gogume pauumsv : om. oo. mm. mm. cozume mhwma Ecucmu mugmcmo um. Na. Na. manage uofla : = so. mo. vogume ;Ufi~uuuUHm : : om. wozuus wouummv menu Hanan d 00 .D J .0 rd .0 1 d H... .b J 1! Tr n B I I. n by I 1.. n P“ 0 3 B H O 3 B u 0 3 e U 3. 3 1 D‘ 3 3 .4 p. 3 3 ,4 D. a 3 O a 3 O A 3 O l 8 w J 3 w r." a m I J I J T... l T. d I. d T: d 3 .8 3 on .J 9 q T. U. T H I. 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E n; my ,9, x .9... .5. m2 m2 m2 my. a ._ .. .3. m2 m2 m...” .02 its me n: n: a. .3. s. .3. a .. : «so an n? m7 m2 m: L? m2 m2 set its as n : : my...“ 0.7 (.7. Or. I F2 Pu? m2. .0? m7. W7 N Z .. r/ 2,, 9, ma « m2 m2 m2 m2 m... a 23333 83.39,. «2 my new .7 m2 m2 m2 02 wz m : : .07 Pt. Q? m2 m7. kn m2 mm“ +V I I n; m2 m2 m2 scan m2 m2 n : : mz m2 m2 m2 m2 u- N : : m2 nz m2 m2 we a macs casuacma m2 m2 m2 m2 n : : wz m2 m2 a : : sea «es m : : *9‘. N I Z . H ucoucoo heap H7 ow m : : +V .. __ m : . N : : . N acoueoo ufiwn m : : q : .. m : : N acoucoo beam 2 a n N cult- m In libiluiwi: p m a m .w a scenic; Guano; cauquo: senate: ca ucoam>m36w ousumwoz cw mmofi ccwuauww Cw ucmucoo hm~o amen pouoEdLom .UmDCHucco . ma c.m4xtp Improvements in Methodology - Sampling Lowland soils-‘The presence of non-_significant correlations between soil characteristics in different hori- zons was great, as indicated by the results from Table 18. It is there- fore suggested that future vegetation studies in the lowland-hardwood land type should ignore the measurement of physical soil parameters at least to the extent of those included in this study. Among the chemical measurements organic matter, expressed as loss on ignition, and reaction (pH) can also be eliminated from studies of this type. The lack of significant correlations for the soil physical determina- tions can only be attributed to the lowland nature of the sites with high water tables. The effects of sand, silt, and clay content, loss on ignition, and reaction on lowland-hardwood vegetation is probably being masked by the selective pressures exerted by recurrent high ground water. This would be especially true for stands found in a very wet or wet environment. It would be true to a lesser extent for stands found in the wet-mesic segment of the environment. culi‘fi]- Ground Water and Precipitation Measurement.--Analyses of variance were performed to show the significance of differences for water tables through- out the growing season. Summaries of the problems for the analysis of vari- ance of perched and actual water table levels are shown in tables 19 through 22. All tabled F values were highly significant at the .01 level. Mean depths of the ground water levels within soils for the entire growing season and least significant differences are shown in table 23. Fluctuations of the Water Tables Within Soils.--The greatest amount of variance in actual water table levels within soils occurred at Opposite ends of the continuum placement of lowland hardwood stands (1394-2429), A parti— tioning of the variances indicated that over 96 percent of the changes in wa- ter table levels occurred within the Locke and Spinks soils, and the Sebewa and Gilford soils of stands Ml9L, M195, and was, respectively. For example, the partitioning of variance for Spinks soils is as follows:1 (a) 240.26 - ve2 V32 error 8 l : ve 240.26.: 1 - avsz 2'- Vs - 14.10, therefore 2 (b) VS 3 98 percent V82 V82 where VS 3 variance due to soils, and V9 : variance due to error I See Table 20, for values used in the sample calculation.. -1¢»2- TABLL.- 19. Summary of the analyses of Variance showing the differences in perched water tables due to week of measurement through- out the 1961 growing season. «an... ibiiroe of” yogi {at ion 5011 series Stand seeks Error F(a) lucke M19L 153.37 5.37 28.58 Spinks M198 190.26 2.22 85.61 Conover A14C 33.24 8.04 4.13 Brookston A148 56.12 5.56 10.10 Gilford 1106 54.45 2.70 20.19 Brookston W53 166.29 2.86 58.14 Pewama W58 115.25 3.57 32.21 Linnwood Al4L 51.02 1.26 40.34 Linnwood L10L 26.31 .51 52.13 Carlisle VZOC 15.54 .10 161.50 Brookston M268 75.97 7.22 10.53 Ceresco M170 29.36 4.16 7.11 Cohoctah M17C 75.23 2.12 35.49 510an N225 36.55 .88 41.33 Rifle M26R 44.51 1.05 42.44 Sebewa W43 17.40 1.94 98.65 Gilford W45 23.69 1.06 233.39 Barry M19L 26.05 .56 46.45 8all were significant at the .01 level 19 Degrees of freedom 57 TAKLE.- 20 . Summary of the analyses of variance shOwing the differences ——. .4o- out the 1961 growing season. u.“ -.--- in phreatic water tables due to week of measurement through- §_urce of Variation Soil series Stand keeks Error F(a) —--Mean squares--- Locke M19L 2727.24 28.18 96.79 Spinks M198 3388.63 14.10 240.26 Conover A14C 467.83 43.03 10.87 Brookston A148 442.23 50.53 8.75 Gilford 1106 422.11 22.30 18.93 Brookston W58 1341.63 23.29 57.60 Pewamo 053 934.92 28.90 32.35 Linnwood A14L 404.18 9.47 42.69 Linnwood LlOL 220.17 5.16 42.66 Carlisle V20C 119.52 .89 134.06 Brookston M268 807.24 97.45 8.28 Ceresco M17C 348.20 54.80 6.35 Cohoctah M17C 709.08 23.02 50.80 Sloan M225 260.73 4.48 58.23 Rifle M26R 331.13 7.01 47.21 Sebewa W43 1357.92 13.47 100.80 Gilford 048 1750.62 8.59 203.71 Barry M19L 270.59 4.26 63.54 a all were significant at the .01 level Degrees of freedom 32 96 - 1 (+5; in TASLH.-21 . Summary of the analyses of variance showing the differences in perched water tables between stands throughout the 1961 growing season. -_~._--. S()i|rt'e of Variation Date of measurement Stand Error F(a) "-Mcan squares--‘ May 6 108.64 4.75 22.87 " 13 143.02 6.22 23.01 ” 20 124.28 11.38 10.92 ” 27 129.19 7.88 16.43 June 3 126.:’ 13.33 9.46 ” 10 115.#0 10.31 11.19 ” 17 133.12 14.08 9.60 ” 24 156.22 16.70 9.36 July 1 139.91 14.45 9.68 ” 8 141.73 12.71 11.15 ” 15 138.02 12.19 11.33 " 22 139.76 12.64 11.06 ” 29 161.08 11.89 13.55 August 5 190.91 11.99 15.92 “ 12 196.16 15.10 12.99 ” 19 206.51 11.08 18.U1 ” 26 188.74 16.20 11.65 September 2 177.52 13.02 13.63 ” 9 165.41 13.57 12.20 ” 16 2411 52 111.21 23.:17 DP&ICCS of freedom l_7_ 51 fi2.111 were significant at the .01 level IAHLK.- 22. Summary of the anglysjs of Variance showing the difference in phreatic water tables between stands throughout the [gel growing season. Date of measurement —.—-—- Source of variation Stand April 22 .. 29 May 6 ll 13 ” 20 H 27 June 3 II 10 II 17 H 24 July 1 " 8 IV 15 H 22 H 29 August 5 II 12 3 19 " 26 September 2 ll 9 ll 16 " 23 ll 30 October 7 ” 14 H 21 H 28 November 4 H 11 II 18 " 25 December 3 172.37 67.14 112.60 151.96 122.30 138.20 143.47 128.21 154.34 166.55 165.87 163.50 160.58 165.50 98.09 222.28 228.68 227.12 201.80 178.28 170.06 241.44 257.09 228.71 290.1)3 314.31 332.72 356.94 317.51 327.10 376.15 379.19 382.21 ——'-"— -.—- _. 0 A11 were significant at the‘ZOI‘fEVCf 17 --_‘._.. --«Hean Squares--- Error F(a) 1.67 103.44 1.02 ffi).05 4.28 26.32 4.96 30.66 11.12 11.00 7.82 17.68 11.26 12.75 10.66 12.03 11.38 13.56 16.40 10.15 13.61 12.19 11.18 14.63 9.48 16.95 10.76 15.38 10.15 19.52 11.80 18.83 12.52 18.26 10.75 21.12 14.29 14.12 13.22 13.49 13.31 12.78 16.96 22.03 10.73 23.97 11.96 19.13 11.07 26.21 10.28 30.57 9.22 36.08 9.19 38.85 7.72 41.13 7.21 45.34 7.08 51.83 6.66 56.98 6.41 59.66 Degreei of freedom fl.-.“- 51 -146- Table 23-. Mean depth of water tables in soils for the 1961 growing season.a E ii ’ 1: .3 u: u 1: :5 13 E ; E: 2: 3 S .§.§ '52.: . : g “ I 3 - g " a :- ‘ 8 9 g i 3 (J35 , 8:. -5--§6 556:5 1 .000 o 4.40 concave I mmhm 1 66 3: 69 5: :9 29 :2 :9 : : 6 25 6 0 so 33 4 2 f 43 39 2: 5 :4 6 :o o o : z: 5 o :: 5 o o 1 46 6 69 29 6 :o 3 :2 o o :o 27 5 o :3 9 o 4 1 34 9 76 36 7 :6 6 :5 : o :6 4: 5 3 :4 :5 4 3 l 36 30 67 5: :2 23 :3 :6 2 : 26 34 :2 5 :6 :9 9 5 1 26 26 63 4: :2 26 :6 z: 2 : 26 39 :2 7 20 27 :7 5 1 30328945934222!:0234::88233429I: 3: 39 74 49 :2 27 :4 23 2 : 26 34 23 6 27 24 :4 7 ‘ 42 46 77 4: :: 25 :2 22 2 o 24 37 26 6 29 :7 2 :o l 59 59 76 54 :4 44 29 26 2 : 30 4o 29 6 26 27 :3 :o 67 52 63 56 29 5: 37 3: 6 6 35 49 39 :5 so 42 26 24 i 73 57 66 50 29 64 46 35 :3 9 47 54 46 2: 35 54 49 29 i 76 65 66 55 37 66 46 4o :7 :2 50 53 43 ‘25 39 59 52 25 79 67 9o 57 3: 7: 50 45 :6 :5 5: 57 47 32 4o 62 57 26 '1 65 72 94 59 36 74 54 59 2: :7 55 56 46 27 42 67 69 :7 1 69 73 64 57 34 62 33 4a :6 :: 56 43 36 :: 45 69 56 :5 1 93 74 67 63 26 65 4: 46 :9 :3 55 45 4: :5 46 66 65 :9 1 :oo 60 92 72 36 72 49 59 29 :6 6: 5: 47 24 49 74 69 24 ‘ 7:6767262::e:25:23343630323:3:n 1 70 7o 69 40 30 26 6 29 :6 6 37 39 34 4 27 29 9 :5 3 7: 66 7o 42 33 29 :: 26 20 6 4o 36 30 4 27 24 :: :2 ; 888272502:36:933:I94239358303263” ¢ 97 69 69 55 26 4: 25 34 :4 4 46 46 42 :3 33 4o 25 24 \ 768:6742IZBSI7263:38363062626nlo 1 95927246:54:23306038363:7283|2:20 J :o: 96 76 5: :6 42 24 so 7 : 44 39 32 6 30 3: 29 2: I :06 :oo 76 69 2: 46 39 35 :: 4 49 39 33 :2 34 36 26 24 1 ::4 :02 60 6: 22 5: 32 36 :3 4 46 36 33 :o 30 4: 32 30 1 96 96 7o 5: :2 36 26 32 3 o 44 35 24 7 26 27 :6 :6 :oo 96 72 56 :2 39 29 33 4 o 45 37 26 6 29 29 :7 :7 1 99 96 7o 46 6 27 9 24 : o 4: 26 29 4 22 :5 6 :: 1 32 96 94 74 43 7 25 5 24 z o 4: 25 :4 z 23 :: 3 6 92225 ll J. 45__ZZ__ul_.Jl_jnL_1§__JL_Jz. ] 4.0597IZIZB4I064.!7I3945754 ; qooll290506lllllz75222!?"56875 J 5 ..-. 1 g I A A 6 6 c o o 6 c I: t r r r o o A ‘ O- 1 2 : an Arranged according to soil series and the continuum placement of lowland hardwood stands. -lQ7- The previous sample calculation shows that 98 percent of the changes in water table depth of this soil could be attributed to time. Error variance due to local differences between wells within these stands was correspondingly high when compared with other soils, except for the poorly drained Cilford soils. Only four percent of the variance was due to differences between well locations within a particular stand.l High variances indicated by rapidly changing water levels were also noted for the perched water tables in stands of the beLa-gley soils; however, this was not true for those stands found on two-storied alpha-gleys. Differ- ences in variation between actual and perched water tables could not be en- tirely explained on the basis of existing differences in horizon textural characteristics, or the presence of impermeable layers in the profiles of these soils. Apparently the ground water level is being determined by topo- graphic and subterranean drainage controls, which would be indeed difficult to investigate without calibration of the watersheds in which these stands are located. The soils of these stands were subject to the influences or two different topographical recharge patterns (water regimes A and D) of the Sycamore and Sloan Creek watersheds. Such high seasonal variation and similarity in water table levels of stands so different in species complement would seem to indicate that the arbitrary assignment of stands to a particular moisture regime may be weak when a single observation is made on water table depth. This would be es- pecially true for any ecological study concerned with determining the rate of successional change. IAn average value for all stands. actual water table variation within soils was least in the very poorly drained mucks and certain soils of the alpha-gley type. An analysis of the COmponents of variance shoved that over 90 percent of the fluctuations with- in water tables of the Linnwood and Carlisle mucks (stands Ath, LlOL, and VZOC), Rifle peat, ponded Barry soils, and the alluvial Sloan soils could be attributed to time. The remaining variance could be allocated to differences between well locations of the respective stands. In the remaining alpha-gleys, the variances,due to time. were greatest in the Brookston and Pewamo soils of stand WSB. This was evident for perched ~as well as the true water table depths recorded during the course of the study. Although the soils in the stands was and H58 are under the influence of the same physiographic recharge pattern (water regime D) , the higher clay content in the deeper lying horizons of Pewamo soils is evidently responsi- ble for greater variances in perched tables. Both stands are situated in 'closed basin depressions and receive little water from adjacent areas. Actual water table levels for the complex of stands in water regime 8 showed that approximately 70 percent of the variances could be attributed to time. The Conover soils had variances only slightly greater than the more poorly drained alpha-gley and organic soils. Approximately 30 percent of the variances were due to differences between well location within each stand. weekly variances in perched water tables were greater in the latter soils. This condition might be attributed-to an increasing clay content at lower depths for Brookston soils (stands AlaB) and the clayey parent materials underneath the Linnwood muck of stand AlQL. The high silt content of the Conover profile (stand A140) was apparently not influential in causing greater variance in perched water tables when compared to other beta-gley soils, or even some of the alpha-gleys. -lé+*!- Although perched water tables were not frequent in shallow or deep or- ganic soils, their occurrence was observed when the well boring penetrated into the underlying materials. Thus, differences in variance in actual and perched water tables of Linnwood muck (stand A14L) and Rifle peat, compared to other Linnwood and Carlisle mucks, could be attributed to the signifi- cantly higher clay content of the underlying materials. Differences between the wells of these stands due to local variation was insignificant. Local variation in actual water table levels (over 35 percent) attribu- table to physiographic differences in well location within each stand was greatest in the Brookston soils of stand M263 and the floodplain Ceresco soils of stand Ml7C. Variance in the Cohoctah soils and Gilford soils (stand L106) was intermediate compared to other soils having poor drainage. In these alpha-gleys, approximately 80 percent of the variance was account- able to water level fluctuations throughout the Spring, summer, and fall seasons, while the remaining 20 percent was due to variance between wells. Fluctuation of the Water Tables Between Soils.--The largest amount of variation in actual water table levels between soils occurred during the first week of the study.1 Variance partitioning revealed that over 90 per- cent of the variance in actual water levels for the entire growing season took place from April 22 to April 29, and from October 7, until the end of the study on December 3, 1961. After May 6, when a general downward trend in the actual water table levels was observed, these analyses showed an increasing amount of variance between all soil types. This trend continued until the termination of the study, except for some minor and expected reduction in variances due to I"See Table 22; column of values for F. - l 3(}— ' partial recharge oi the phreatic surface from rainfall. Partitioning of the variance also showed that the lowest amount of variability (71 to 78 per- cent) between the water tables of all soils occurred during the summer sea- son from June 7 until July 12, and from August 19 until the beginning of {all on September 21, 1961. This could be largely attributed to the effects from convectional Class 4 advanced storms, which were insufficient to sat- isfy soil moisture profile deficits. Increasing variances, between soil water tables toward the end of the growing season, might be accounted for in the continuing drop of water 1e- vels in the beta—gley Locke and Spinks soils, which was directly opposite to the trend of levels observed in other soils during the latter part of the study. oldi— ggrgglatiggswfietween Species Distribution and Ground Water.-- In order to study the relationships between Species distribution and water tables simple correlations were calculated between the importance values of the ten most abundant species and ground water depths during the growing season. Similar analyses were also calculated between species' basal area and ground water depths. The species were: American elm, red maple, green ash, swamp white oak, silver maple, sugar maple, American basswood, slippery elm, northern red oak, and black cherry. Data were available from 14 stands. The stands were (M19L, M193, AlAC, A148, LlOG, WSB, AléL, LlOL, VZOC, M268, Ml7c, M225, M26R, and WAS), in which water tables were measured. Stands means were used in the analyses. The items entering into the in- put were: . l. importance value of each of the ten (10) Species. 2. basal area of each of the ten (10) species, and 3. water table depth for each of 28-weeks. There were 12 degrees of freedom for each correlation. Correlations which fell between r.05 and r.02 were disregarded. The correlations as calculated apply to the period April 29 until November 4, 1961. This period was con- sidered to be the most important hydro-ecologic segment of the study. Species Distribution Related to Ground Water Depths.--The correlations of biological significance between ground water depth and species importance involved two species-- sugar maple and American basswood (Table Z6“) When basal areas were used in the calculations, the same type of association was noted. It may be observed from the correlations and ground water depths in tables 20 and 21 respectively, that American basswood was more tolerant of shallow water tables and poor aeration than was sugar maple. For instance; the water tables in the stands where basswood was present was within one inch r 4321.. {AEIL.- 24 . Correlations between ground water depth and importance of sugar maple and American basswood. (An additional 224 correlations for 8 other species were not signiiicant.) —~;————"“ C0 rcLitiOns between depthugl water table and Date of measurement S-ga maple American basswood importance basal importance basal Value area \alue area between .':y 6 and June 17 \5 NS \S hS between June 17 and August 19 High** High** N3 N3 between fr‘ust l9 and September 2 High*** High** NS NS between .-yteober 2 and September 8 High*** High** High* NS between i - . in , V, (‘ ' ,1 > / if)??? A ~ ‘( \H‘ \. u Lemuel ; and hovember 4 “18h Highk: US US Non-significant relationship r greater r greater r greater than .612, than .601, than .780, significant at 2 percent level significant at 1 percent level significant at .01 percent level ~1J3a TABLE.- 25 . Relationship between the range of ground water depth and importance of sugar maple and American basswood- ll Date of measurement l'-.’ater table depth in stands in which Sugar maple American basswood was__fi __k was present absent present absent#_ between May 6 and June 17 9~83 0-41 2-83 0-41 between _ JUdC l7 and August 19 29-100 1‘74 1-100 1-74 between August 19 and September 2 18-71 1-39 1-71 1-39 between September 2 and September 9 29-71 4'40 4'71 4’33 between September 9 and November 4 35-114 O-Ao 6-114 0-46 -154- of the soil surface between August 19 and September 2 (Table 25). During this same period, water tables were at least 17 inches deeper in those stands where sugar maple was important. Such factual evidence reaffirms the known silvical characteristics of these species (Braun 20, Bray and Curtis 23, Maycock and Curtis 110). ~135- Correlations Between Water Table Depth and Date of Measurement.--ft was noted during the measurement of water tables that both the drop and recharge in ground water was approximately equal between similar Stands for each week of measurement. Presumably this situation was brought about by the eifects of local evapo-transpiration and recharge by rainfall. Such a condition was most evident on muck soils which were least affected by gradient differences. In order to study the relationships between water table depth and date of measurement simple correlations were calculated between the dates on which water tables were measured and their recorded depth. The periods of observation on successive weeks were from April 29 until November 4, 196l. ‘Water tables were measured in stands (M19L, M195, A14C, L108, WSB, Ath, LlOL, VZOC, M268, M228, MZGR, and WAS). Stand means were used in the analyses. The items entered on the tape- input were inches of depth recorded below surface datum for each of 28 weeks. There were 12 degrees of freedom for each correlation. The correla- tions which fell below r-.78 were disregarded. The results of the analy- ses are shown in table 26. The results of the data suggest that a strong association existed be- tween water tables in stands from September 16 until the end of the period entered for the analyses. Stabilized ground water levels could be partially attributed to a reduction in the amounts of local evapo-transpiration within stands and the surrounding areas. In addition, most of the precipitation during this part of the year is produced from Class 4 storms of cyclonic origin (Smith and Crabb 137). Such storms became prevalent during the latter weeks in October and the early weeks of November. This type of storm is of low intensity and serves to satisfy existing soil moisture deficits. Throughout the winter season, of course, storms of this type will produce recharge of the ground water atter profile unisture deficits cease to exist. u t ‘)() .-. .caozm sum Hw>o~ Ho. ecu um ucmowwucm«m mums nowzz .Awn. saga woumouw.uv mcofiumauprU #H4m ucmoauwcwsm uox umx am. am. am. em. Nm. m2 m2 m2 m2 m2 a h32262 mm. am. am. mm. mm. m2 mz m2 m2 mz mm : mm. mm. mm. «m. Nm. m2 m2 m2 m2 m2 Hm : mm. so. am. an. em. m2 m2 mz m2 m2 as : am. mm. mm. mm. am. m2 m2 m2 m2 m2 a sesouuo mm. mm. mm. um. um. mm. so. om. m2 m2 an : gm. om. go. um. mm. mm. no. ow. m2 m7 mm : om. mm. mm. am. am. ma. co. om. mz m2 o~ : em. mm. mm. ow. mm. mm. oh. am. mm. mz m : Na. as. mm. em. mm. as. we. om. we we N hostesses am. om. mm. am. m2 m2 co. m2 we m2 om : so. om. ea. ow. em. mm. m“. mm. m2 mz 0H : so. no. so. mm. mm. mm. mu. mm. mm. mm mm : so. ma. mm. am. am. sm. ma. em. as. ma. m umswae mo. oa. mm. mm. mm. mm. mm. ow. mm. mm : am. so. om. om. em. an. mm. mm. Nu : «o. 00. “m. cm. am. am. ms. m~ : so. om. mm. mm. mm. em. ow. w : as. as. mm. we. so. an. cm. H size so. so. am. so. on. Hm. em : so. No. om. mm. fiw. Nd : om. mm. em. m2 ofi : mm. am. on. m2 m mesh so. am. mz.. RN : om. mm. ow : so. cw. ms : NO. 3 N”: am NN no aw as as am em a aw i- $.29 30.4 Ed 0 u so a o so 12.3 NT; 9.3.. >8. 1 3.2%.. ucwEousmmoE who one; edge: IIIIIIDIII- l. ‘ I. 1-1.. 'v‘. -51 Illillllllu!’)lu".i (Ill... [until v: .D-.. I I‘- -l.'l'|lblli .II III 00‘. a: I .I. .l.‘r|l.b|.| . .c3onm use Ho>o~ #0. one an ucmowuacmwm ouoz cowzz .Awm. coco woumouw uv economgowuoo -< a ucwowwflcwwm uoz u m2 umnEo>oz penance umnfiuuamm um3w2< as = as = m.” .— m = s ssae om : NH : o~ : m mash mm : om : ms : 0 sex as. oo.s oo.s oo.s as. am. as. No.1lno. no. so. No. No. a mm. mm. mm. mm. mm. mm. mm. no. No. no. no. em. mu oo.~ oo.d mm. mm. mm. mm. mm. mm. mm. mm. so. ~m oo.d om. mm. am. so. so. am. No. mm. mm. ¢~ so. as. as. as. as. as. Na. Na. mm. a so. as. «a. ma. ms. so. No. ma. on ma. om. cm. mm. so. «a. co. mm co. co. co. mo. so. mm. o~ mm. mm. mm. mm. No. 0 mm. mm. om. cm. N mm. em. mm. oo.# mm. mm. . n 4 mm um ed s om mm 0H m N mmilima mg w some sumo ouwc noncocc ponEoumum umnmdm -1 r. eucoz .cmscwucoo . ucquusmmmE w0 some om-.mgaae 458-. During the period from August 5 until September 16 relatively slight changes in water tables occurred as indicated by the correlations in the table. These slight changes could be partially attributed to the effect of increasing or decreasing evapo-transpiration locally as well as in the sure rounding watersheds. The ground water depth was also affected by convectional storms of moderate to severe intensities during this part of the year. Changes in water tables for the period June 17 to August 5 were also compli- cated by storms of this type (Class 6 storms,Smith and Crabb 137). Non-significant relationships between the water tables in all stands occurred from about May 6 until June 10 when water tables were decreasing or increasing differentially. These changes were most strongly at variance during the earlier weeks of this period as a result of thawing and freezing of soil water which caused differential recharge of the phreatic surfaces. Immediate and continuing effects from precipitation, intermittent changes in the amounts of water being transpired by trees, and temperature differences probably added to confusion of ground water levels during the early part of the year. Since 2376 measurements of water tables were made during the present study a subsequent series of measurements with a minimum effort should yield similar trends of fluctuation. In addition, a series of yearly trends could be established in ground water levels for subsequent studies in lowland hard- wood areas. -131}- improvements in Methodology - Sampling_Ground Water.-*The relationships indicated by the results from the correlations between water table depths and the date of measurement suggested that precise data may be gathered by in- termittent sampling of ground water depths during the growing season. Con~ sidering the annunt of time and effort expended in collecting such information, a reduction in the number of field measurements would indeed be desirable. From the analyses of the ground water correlations for 1961 the follow- ing information was derived: a. If one could afford to sample ground water depth only once during the growing season the best time to sample would have been between June 17 and July 15. The lowest correlations between samples taken on any of these dates was between r'.78 and r. .81 b. Sampling twice during the growing season would not have given more information on water table depths than sampling once. c. If one could afford to sample ground water depth three times during the growing season the best time to sample would have been on May 20, July 29, and August 19. The lowest correlations between water table data taken on those dates and intervening weeks was .83. Correlations for additional samples taken on ground water depth are shown in table 27. -TOU— TABLE.- 27 . Optimum sampling data in 1961 to give maximum information on water table depth at minimum effort. i£_one could afford to sample the best time the lowest correlation between this many to sample measured water table depths in times ___ was intervening weeks was (a) number ----- ---date------- 1 between June 17 and July 15 .78 to .81 2 (b) ------------------- ..... ----- 3 May 20, July 29 and 4 May 20, July 29, August 19, and August 26 .85 5(C) - ______________________ _ ...... ---- 6 May 6, May 20, June 10, Julyl. August 19, and August 26 .90 10(d) April 29 and May 27-then on June 17, July I, August 5, August 26, and September 30 .95 'YS) A11 correlations were significant at the 0.1 percent level. (b) No combination of two dates gave better results than the best sampling on one date. (c) No combination of five dates gave better results than the best sampling on four dates. (d) If one desired 90 percent (r2) information on water table depth, sampling would have been made on these dates. Stand Similarities From the values within the table of community coefficients,(Table:28) several relationships are suggested. it is evident that on peat soils the tree composition is quite dissimilar to that found on mucks and mineral soils. in addition, the canopy vegetation may be just as dissimilar between peat soils. The strongest disparity is indicated, however, between the vegeta- tion types found on pests and alluvial soils as shown by the extremely low values at the base of Table 29. A comparison of the remaining community coefficients indicated that no other abberations were present within the table. Since 23 different hardwood species were used in the construction of the table, the continuing homogeneity of lowland hardwood stands is demon- strated by the small differences between most values. As has been previously mentioned, an attempt to show stand similarities through the use of the basic community coefficients proved that the stand alignment was distorted. The use of the gradient treatment suggested by Bray gave a better perspective of the spatial relationships between stands. The Spatial separations along a single axis actually represents the compres- sion of a two dimensional relationship into one dimension (23, 43), The points at which each stand is located along the two-dimensional axis is conmunly referred to as its W value.L An attempt to portray relationships based on stand to stand similari- ties and the data for the environmental measurements were included in a series of graphs. it was decided that the best method of representing the continuous nature of the gradient would not require any additional or arbi- trary segmentation of the data. Since the water tables of the stands were actually measured, rather than inferred, the arbitrary division of moisture classes would be presumptuous. 1a coefficient olr-communlty slfilldrlty., QC (1. mcfi -Ib2w OOH Omd mflm omm Nm . Nod mm mm 0 mm Rafi ad mm NHN wflm M¢M OH“ QNH “@fl Omfi do 0 cud oo~ mqfi cnw mmw _m~ Or“ any ~w~ mod mm~ NNN 0mm 7mg ooN Nfim «cm om_ 0mm mam mmm mmm q~m cam mmm wnm mam qu dqm ban qcfi nxm mew mmm mom 40m wqm RAH mum com com m_m mm_ mm; “om 0mm mum mom ANN mam CNN mom mom nfig mqm 0mm 0mm mwm mmm new 4mm mad nnn NHN “mm mom mmN nmm wmm 0mm cmm omfi wmfi mtw com mom mfim on“ «mm m¢m Nflm mwm cam mam mom Nmm mam nnn mam mmm Nmm mgm wax m¢m mwm oflm mmm 0:“ 0mm wmm Amw mnm mfim 0mm mmw Adm mam mm“ m qaw mow mam cmfi qmm New 0mm mam omm 3mm mmm emu mew qmm cxm 'AJII «((0 UPC N N3 34 m: z: 2: 2: 98 No“ 3“ SN 0: EN 93 cow IN «8 hi. . ll (\ “A.” ”H“ C 1 ”IL ”NV 1 H A N ”W1 V 7 v VH1 1‘ S .6 1..” «7. Z I. .l H. Z 0 Z Z .r/ 1.. M” “w Ht U U 3 m]. In 4n. r? \l!‘ .r/ C 0} (L O .20 no 7 (J (3 ha (0 Lu. 9 V cm U m - 3 C. no ,r: n: 0 8 W. D on an H ,1” 3 S n... u b 4 k n. uLu C M .muzsou EnchH mo mncmum vOOBVLmLuwcmHBOH o~ mowumam my»; «a mwsfim> mucmupanw mcu co vwmmn ADM "3v mucwwufiwwwou xuwcsEEou w~.«.ygmflgv +55- The relationships of lowland hardwood stands based on their N values and the various soil measurements are shown in Figures 31 through 40. dean values of the edaphic parameters are indicated along the ordinates, while w values for stand arrangement are shown along the abscissas. At each stand position changes in the importance values for {our of the leading dominant species are also shown. This was accomplished by using single lines scaled to actual importance values at the cardinal points of stand position. Any attempt to represent a species pattern through the use of line dia- grams was not suitable for the following reasons: a. the number of stands used in the ordination was small, and lines connecting stand positions to show species changes would have resulted in identical patterns. b. the use of actual importance values attained by the species ap- peared to be greater value in gradient treatments, than the selection of an average importance value to define species pattern. _§tand Shift Related to the Number of Stands Sampled.-- The spatial distribution of the stands along the abscissa, representing similarities in tree compo- sition between stands, is a resultant of the number of stands sampled. It is apparent that the inclusion of additional stands in the present techni~ que would produce an entirely different distribution on the gradient. Likewise, this condition would be equally true for stands which were omitted from the technique. The shift in spatial relationships which occurs when a different stand is selected as a base of reference is shown in Figure 31. It nay be observed that the magnitude of shift is least for those stands containing large amounts of silver maple and small annunts of red ample. Since an index of similarity formed the basis of the gradient treat- ment, stands most unlike the reference stands would be displaced the least. .mocoummop we omen o no memo venom mo woman :« oouumaom ma mom: venom coca musuoo Lowna .mvcmum voozvhms vcma3o~ we mafigmco«uo~ou amuumdn cw uuwsm egg . Hm .wum ‘ OHDJ<> I (SP RN DDN if. J a... 8... O. u. 8.. t .2... 8a. I. I u .NEHWLEM Lalo 1 M . at. on. 3:. iv. a e v .e m . in m m m r“ \l Ollllllll. uc~> W Oll« «Is. a .( an. .6... .- I lllld’lll l.llllllllsl Although most of the stands located on mineral soils were shifted downward, all stands located on mucks were shifted closer to each other. The remaining graphs were constructed on the basis of the original gra- dient technique since many combinations of reference points could be included in a treatment of this type. Stand Shift With Changes in Organic Matter (loss on ignition).--A comparison of the changes between stands with changes in organic matter, or loss on ignition from the soils, shows a distinct separation of the stands into mutual groups (Figure 32). It may be noted that red maple is most strongly represented at the left of the gradient on peat and muck soils. An analogous situation prevails for this Species on the mineral alpha-gley soils. Al- though this species decreases in importance as the w values increase, it is most weakly represented on the right side of the gradient. The three other species green ash, American elm, and swamp white oak, enter the gradient on Linnwood muck soils. Swamp white oak is somewhat better represented than green ash on Linnwood mucks. However, green ash usurps the importance of the former species on alpha-gley soils in this sec- tion of the gradient. The synecological requirements of these two species for sites possessing high amounts of light and moisture is rather well known.(125, 150, 154) Although the two species occur at about the same stage in sucession, green ash appears to be more tolerant of wet extremes in moisture and full sunlight than does swamp white oak. The differences in amplitude between the two species in this segment of the gradient may be due to differences in such conditions between the stands. The trends for these species are uncertain toward the right as both Species continue to be approximately equal in importance along the remainder of the gradient. Although American elm attained a high level of importance in the ma- jority of stands, it reached its maximum deveIOpment toward the right of the gradient. .ucwwpmum onu macaw csonm Oman mum mofiooam ucmCMEoc usou new moadm> mucmunoaaa ecu ca mmwcmnu .mw:~w> 3 new Acofiuacww co mmOav mHNOm wzu mo ucwucou nouums owemwuo on» so pause mpcmum poozpumn,pcma3o~ we awnmcomummwm.mm .wfim 83.; - 8n F on. 8. .8.- ‘ SN. .0. N u n N . u .0 ,0 E. >- gl 2.!!- o m I I . as a 0 ) a; m 3 m all He .1 .83 w Id Ice—ll- .n 53 I... 3...- II. on #3 ~ .1 In! .- g as! nil-I... v ,6 ~10!» The fact that this species did not show up as a major species at either end of the gradient satisfies the assertion that the gamut of lowland hardwood stands within the country had been sampled. EEEB§_§E££E,E££h_ghifl§33difl_flfi” The use of mean soil reaction as a parameter showed a strong rise to the right from about 4.5 to 6.2 on the acid peats to mucks. Slight increases toward the right were more oe less continuous in a band of values from 6.7 to 7.9 for most of the remaining stands eventually becoming asymptotical at the end of the gradient (Figure 33). Of the four species plotted for this ordination, only red maple appeared to be tolerant of extreme acid conditions below a mean soil reaction of 4.5. The vacillating decline of this species toward the right of the gradient may represent the initial side of the bimodal curve reported by other authors (54, 110). Likewise, its entrance at the extreme right may reflect the initial upswing of the second peak. These changes are not necessarily the effects of soil reaction, except where this species exhibits a clear tendency to attain optimum importance at the lower end of the gradient. Even so, red maple may simply possess a wider ecotOpic amplitude for acidic peat soils compared to other lowland hardwood Species. Stand Shift_with Changes in Sand, Silt, and Clay Content-- The distribution - v-w~!I-——— of the stands also showed noteworthy patterns of change when mean values for textural variation within the soils were plotted (Figures 34, 35, and 3b). The bands of vertical separation decreased proportionately in width, from sand to silt to clay, between the 12 stands sampled for soil texture. The coarse sand fraction, beginning at 47 percent at the left of the gradient, showed an increasing width in the band of values of approximately 25 percent. A median value value of 57 percent was reached on the far right. «lbs- .mm:~m> 3 can magOm one we me one co co vmwmn epcmum poospumzlpcma3o~ we nonmc0wum~om .mm.wwm .9344) I new: I‘lll OM14. OF - &. TII II J. . at" our .v ‘9 law :25: .n ace :2.- alcum o~ 8.4 (cure 0. and: evenness. av-d now. (001) m a... L ~369- ~«0m one aw meow uceuuwm «no :0 comma mpcmum coozpumsupcm~3ofi «o nfizchwomHmm .wo:~m> 3 use 3’- use: also-g an :3 Ha...- allo on '49... 5’ it... an? “fivotflfiw (some) on" -121]- .mo3~m> 3 can mdwOm ecu cg ufigm ucooumq as» so momma mosque poospuenupcmHBod mo nnnmco«uw~ox 8m 8... law 8. 8. on 0! u: c an “09 09 (mama) an 0!. at} 8.. .v M 3m .3253 .n :3 at... I...» .N m a: :38 .. 1 SJ; Hush:- 00' .mm.wae m 0 . # a can an eon 8. 8. 02¢ ov- 8N. 0C 4.71- as 0!. 09 (mum) mo 33- one . a Id 33.5.! .v 3 at... ale-n .M x: 338 .. ~33. 35:83. a 00' -1,’;{- The intermediate soil fraction, or silt content, showed a slight de- crease from 32 percent on the left to 25 percent on the right along the en- tire gradient. The widest part Of the silt band was coincident with the range of w values for sand but was somewhat narrower having a width of 23 percent. The trend of change for the fine fraction or clay content of soils deviated least showing only a slight increase from 18 percent on the left to 21 per- cent on the right. The maximum width of the band, only four (4) percent, was considerably less than that of the other soil fractions. Within the stands sampled for soil texture, three Species increased in importance toward the right between w values from 200 to 250. American elm - and swamp white oak showed stronger importance values than green ash, while red maple declined in importance for this portion of the gradient. _§tand Shift with Chagges in Moisture Equivalent-- The use of the moisture equivalent in the treatment (Figure 37) showed a continuous rise from 25 to 32 percent, which was preportionately opposite to the trend observed for the intermediate soil fraction. An abrupt break, unlike the trend for silt content occurred at a w value above 250, which corresponded to the two stands found on sandy soils in_the upper segment of the continuum. Changes in the importance values for the four species plotted in the graph followed the patterns described for soil texture. .mosdm> 3 man adieu oz» we ucmfim>wavw ouaumwoe ecu no women museum noosrumzuccm~3oa uo mwnmcofiumawm .mm .wam 33: - 8n Bu can on. 8. S N u n “.14: s u a I‘I fl u 8.. m 3: a J . ‘ .3. m mi. 1 a m a a, Us .08 o. W m 3a .3252 .n ( :3 2:.- I25 .u x: :88 .. 9 I5: 213:. t m c n . _§tand Shift with Changes in hater Tab]es-- A graph of the lowland hard- wood stands based on mean water table depth and similarities of tree composition is shown in FigurestSand 39. The catenal relationships of water tables between the soils in the stands is also shown. A distinct separation of the stands was again indicated by using this parameter which was similar to that shown by using loss on ignition. The stands on both alpha-gley and beta- gley soils are arranged into a paracentrical figure resembling a paraboloid which points downward toward the right of the gradient. The taposequence of water tables indicated by the directional lines connecting the stands_serves to confirm the segregation of moisture regimes by soil profile examination. Within the stands located on mineral soils, American elm increased to- ward the center of the figure decreasing on either side of the gradient. This species also showed a general increase in importance with a rise in av- erage depth of the ground water to within 20 inches of the surface. Red maple, with the exception of alluvial soils, was better represented within the area having decreasing ground water depth from 55 to 23 inches. Swamp white’ oak increased in importance within an area from.24 to 38 inches in av- erage depth. Green ash was best represented along the gradient when the ground water came closest to the soil surface. Although this Species was almost as well represented as swamp white oak in several stands, it in- creased in importance on either side of the vertical axis where swamp white oak was absent or deficient. The stands on organic soils are clustered into an elliptical shaped group in the figure. The ellipse or band, which tapers to zero, is 25 per- cent wider on the left of the gradient than on the right. Within these stands the importance of red maple decreased from the left to the right of .maaOm one 3 can sense afinmu amass name no woman avenue vooscusnnvanasofi mo magm:0wusdmm Hand: Cum ov 3.5 33.5.2 .m .20 HS:- sacnm .N x: run-3 .. and; 32.2.2... v cwnuuz madame nous; no endeavomoaou as» mcwsonm munaw> Inuit) I \ s: oo~> ONN' ao.a 0| (Bluall) 310v; I311. JO Midi. ura- .mm .waa .c0mdom mewsouw ecu wcqusw cauumnuonfim manna ueumz we mwcmu ecu wswsocm mes~s> 3 can sumac manna woum3 cmefi :0 women museum coosnumcnvcmfi3ofi mo awnmco«um~mm .mm .wwm QUDJ<> D con on~ com AH. oo. ‘ 00 40.8 8.4 ~l7bo or a... . m n'.< 0'. Oh. I + M mONa . k cs 05 (neat) non sun .00 «mo an. co... on mNNI cJ i_ ,7, do... 08> '-177- the gradient while green ash and swamp white oak increased in this direction, Changes in the importance of American elm, even on some of the mineral soils, were inconsistent with changes in the importance of the other three species. This could be attributed to the fact that the graph shows only present day community conditions. All stands within the study area were not at the same developmental stage in succession. The degree of importance at- tained by American elm, within any stand must necessarily be due to the amount of elapsed time since the species became important in the intermedi- ate phases of secondary succession. Because of its terminal position in low- land succession, a nearly pure stand of American elm would represent the finite stage of development on very poorly drained soils. A graph based on mean water table depth and the continuum placement of lowland-hardwood stands (Figure 40) was constructed for comparison with the values derived from the formula method. By referring to the respective figures, several similarities as well as differences are apparent. It may be noted that the paracentrical figures derived from using the continuum as an index of similarity are not greatly unlike the figures derived by the formula method. This is evident for stands found on organic as well as min- eral soils. Within this representation four additional stands, each lo- cated on contrasting soil types and moisture regimes, were plotted from sev- eral observations of ground water depths during the course of the study. The chief distinction between the two graphic treatments is that the continuum does not represent a two-dimensional compression of similarities in vegetation but rather one dimension, Even so, the use of the continuum values along the abscissa lends credence to the thesis that a strong corre- lation exists between tree composition in lowland-hardwood stands and the mean depth of their respective water tables. Evidence in support of this thesis is shown in the continuing decline of stand position with increasing .cOmmom weasouw ecu weaken Auden mans» nouns wwwuo>m one name noomfin xoonw Ensnmunou wanna no woman museum noozounnuvnmgsoa we awemco«oefiox .oq .mwm szusuoqaa 5.2. 232.58 B kw! MN MN pw LN up D. C D» h. h. h. k. .. bob. O o 0 do.l IL 0 n 3 V . .‘ )3 C 7 m u .i .u. . 0 J m I u moan-ah... .3 was: 5.:- o u m u u mON, 05.: Q'- . 3 32:. 8.. .v New 3w 23.5... .n SN: 0 w. :3 wt:- aacnm .~ H. x: 23.8 .. m 23; marquees... o 3 '4'! m came an: mNNS n . uo~> -yin- ground water depth toward the lower right. Of some importance is the fact that the use of the continuum failed to show dissimilarities between stand composition on alluvial soils compared to peats. As a means of strengthening the ordination by the formula method, an additional calculation was performed resulting in a change of stand order along the abscissa. This procedure involved the integration of units in ground water depth and fluctuation with the w values obtained from the table. The multiplication of respective values resulted in an increasing spatial assignment for stand position which could be best represented on a logarithmic scale. The abscissa, or original w values axis, was expanded to 5.77 times the original range. An example of the construction using percent organic matter or loss on ignition is shown in Figure 41. The catenal rela- tionships of water tables are also indicated. Although a strong separation of the stands is again evident, the prin- cipal distinction lies in the shifting of stand position. Compared to the former technique, where the position of each stand along the axis was based solely on vegetation similarities, a more logical sequence with reSpect to the continuum is now indicated. This is most obvious for stands on mineral soils. Those stands on alluvial and beta-gleys have been shifted consider- ably toward the left side of the gradient. Stands found on the alpha~gleys of ground moraines and basin depressions have brought closer together. Stands located on beta-gley tills having better drainage conditions have moved upward toward the right edge of the gradient. It may be noted that some stands on organic soils have been shifted downward a considerable distance while others have moved upward on the gradient. Within the organic soil group, red maple is now somewhat better represented to the right of those stands containing American elm, swamp white oak, and green ash as the major species. Such evidence would appear .:0wusauu:~« manna noun: we «made x canon canon woos: ewes x muaan> 3 we wanna uwanuwuawod n onsAnOHuanww no mundcm~«0m one we unsound segues ownmwwo one no cause avenue voosouwnuvnofisoa mo magchwuwfiom 30.93ng5 ‘ 6". I I: 81' I .02...) O ~180- a: (IIIOUOO) Illavl 900'... g at 0' Id ecu-I'c on do 3.... and: .N ‘1 g o. :3 Hidi- Q 08. :J 8. 00.: 0. g ‘ _ d..- .3 .m: ~181- to be in conflict with the expected ordering of stands on the basis of com- munity resemblances. The positions of the stands along the gradient, how- ever, are being influenced mainly by the water table regimes within the stands. The entire spatial displacement between stand points is indicative, therefore, of a compromise between the present composition of lowland hard- wood stands and the trends of gradual or rapid alteration in composition to be expected over a period of years. Relatively rapid changes in tree com- position would be indicated by a substantial displacement along the horizon- tal axis while stands in juxtaposition would be similar in their trends of change._ Succe§sion in the Lowland Hardwood Complex-- It is obvious that certain w-.t~- . V0 difficulties are inherent in trying to portray community relationships brought out by the foregoing techniques. The rise or fall of stand position along the ordinate is entirely dependent upon the changing values in a selected parameter of the environment. The position of each standpoint. is actually within a sphere of values within the environment and each stand point is a result of sample size. Such relationships especially those con- cerning succession may be more easily visualized through the construction of a three-dimensional model. Succession in hardwood stands of the lowland survey, implied by the most significant variables of the environment, is represented in Figure 42. The construction is based on present stand positions in the continuum, depth of the organic mantle in organic soils, mean moisture equivalent of the alpha and beta-gley soils, and mean water table depths and fluctuations which occurred during the 1961 growing season. The figure in its entirety indicates a probable succession of 15 lowland hardwood stands. Five of these stands are located on organic soils, six on alpha-gleys, and four on beta-gley soils. The soil types are represented by three colors; red for organic soils, green for alpha-gleys, and orange for beta-gleys. .coHumsuuaau new guano wanna noun: name we muwca van .mugc: ucoan>uavo awaumwoa can neuuua ounnwuo .cowufimoa EBacuu ucou ucomoun uwocu no woman meadow-voosuuw: can~30~ ma new cowuuauumcou 4 .~¢ .wwm -182- ~Id3- The construction may be envisaged as proceeding from the center of a vortex outward along three axes. The stands are represented as a series of interlocking triangles which by definition are either isosceles, equilateral, or oblique. Each stand or triangle is tilted at a slightly different angle depending on the plane from which it is viewed. The three axes have each been divided into comparative units of 30 parts as follows: a. axis O-X, proceeding along a line from the geometric center of the conscruction to the foreground and representing the present position of each stand in the continuum (range 300-3000). b. axis O-Y, proceeding along a line from the geometric center of the construction to the upper left and representing: as. depth of the organic mantle in organic soils (lower third of the axis, 1-3 levels or units) bb. mean moisture equivalent of the alpha and beta-gley soils (upper two-thirds of the axis, 3-30 units), and c. axis O-Z, proceeding along a line from the geometric center of the construction to the upper right and representing units of mean water table depths times range of fluctuation (0-30 units in logarithmic scale). The position of each stand along the continuum (axis O-X) reflects the progression of successional change through time to the present, as defined by stand-species composition. Along this axis the penetration of planes, representing stands on organic soils, into the segment of the con- tinuum occupied by stands on alpha-gleys is attributable to the elapsed time since the origin of each stand. It may be noted that all stands on or- ganic soils are represented by a cluster of red triangles having their fur- thest extension along this axis. The group as a whole resembles a small pyramidal viewed from the terminal position in the continuum. The diminutive -1ga-, base of each triangle is caused from two repressive factors, which may be considered inseparable and essential to the subjugation of lowland hard- wood succession on organic soils. Most important are the suppressing ef- fects of ground water remaining close to the soil surface. Of secondary im- portance is the curbing effect of increased depth to the mantle of organic matter inherent in such stands. Those stands growing on alpha-gley soils, represented by a cluster of green triangles, are tilted toward the upper left (axis O-Y) and become al- most congruent when viewed from the terminal position of maximum moisture equivalent. The fact that alphaqgley soils possess higher moisture equivalents than beta-gley soils indicates that conditions should be quite fa- vorable for the advance of lowland succession toward a meSOphytic climax. This is dependent of course upon the trend of future drainage patterns. Such a situation could only be brought about by increased ditching of adja- cent wetlands, or a series of droughty years causing a general lowering of phreatic water levels. At the present time, however, changes in the Species complement of this particular group of stands is subservient to the smother- ing effects of high ground water. It may be observed that the planes repre- senting certain stands on alpha-gley soils are interlaced with stands on or- ganic soils, while others are intermediate along the O-Z axis. The fact that the pyramidal figure representing stands on alpha-gley soils appears to tilt upward to the left is a result of the continuing pressure exerted by recurrent periods of innundation. This condition of course serves to limit further extension along the O-z axis. Those stands growing on beta-gley soils are represented by a cluster of orange triangles having their furthest extension along two axes (O—X and O-z). This results from the fact that phreatic water tables were much deeper in these stands compared to others and the nearly terminal position occupied -15»- by these srmuls along the continuum. The intermediate extension along axis n-Y is indirectly a result of coarser textured nnterials in the soil pro- files, compared to the alpha-gley group. When projecting the line of sight to the focal center of the figure, it may be observed that between stand se- paration is greatest on the beta-gley soils. in addition, the narrow base pyramid representing this group appears to be falling into the center of the figure. This is an optical illusion created by the crossing of planes at the t0p of the figure and the lower position of stand Ml7C along the 0-2 axis. As a whole, stands of the beta-gley group are actually tilted upward to the right, a result of greater fluctuation and depth in ground water. The degree of Spatial separation along the O-Z axis, between this group and the others, is indicative of a somewhat faster rate in progressive successional change for stands on these soils. A review of comparative values between the present position of each stand in the continuum based on its present vegetation, with a projection of prob~ able successional changes indicates the following: a. stand WAS will proceed toward some point close to a mesoPhytic climax at the fastest rate. b. stands H176 and M263 will proceed at about two-thirds the rate of the preceeding stand, but one-third more rapidly than stands H228, was, and Ath. 0. stand MZGR will proceed most rapidly toward the stand character rep- resented at the present time by stand Alah. d. stand AlAL will show the snallest rate of progressive change, to- gether with stand VZUC. e. stands LlUL and L100 will continue to prOgress very slowly from their present stand character. They may or may not be dominated by American elm, depending upon the eventual progress of the Dutch elm disease. If this species nzves Luiy to tiw'cfiisea923 {We stflrnfiéx4111 h *(Knnluatttll)y red nuu)Ie. f. Since Stands 026R and MlQL torm the relerence stands, a shift from one direction to the other is not apparent. It would be expected, however, that stand M26R will progress rather slowly to the stage of vegetation now exem- plified by stand M26R. Stand M19L should continue toward the mesophytic clinmx at a slightly faster pace than either stand M195 or Alhc. Discussion-- In most cases, the lowland hardwood stands represented by the trend lines of succession in the ordination are relatively stable com- munities. The host of extremely complex interactions, within the sequence of generations that produced these stands, has been instrumental in deter- mining the character of the stands. The stands represented by the cluster of green triangles in the center of the construction are probably the most stable of any group, because of the greater numbers of species present within their boundaries. By the very nature of their age-structure, these forested communities are practically immune to invasion by other Species. In general, the continuing pressures exerted by high ground water serves to set the limits within which the composition and structure of the stands can vary. Although these communities are by no means as diverse in spe- cies composition as their counterparts on upland soils, they are diverse within their own realm. Outside and below this area of green triangles in the ordination dia- grwm, the group of stands represented by the set of red triangles are less diverse forested communities with few species. In these stands the more rigorous environment, deep organic mantle, low pH, and high ground water, are probably instrumental in maintaining a lower species diversity than in the stands on alpha and beta-gleys. The area in the ordination occupied by the series of orange triangles presents a somewhat different picture. Since this zone forms an area of ecological overlap between mesic and wet-mesic sites, it contains numerous Species from both lowland and upland areas. With this resultant increase in diversity the variability from place to place is greater. Although the normal develoPment of these sites is not expected to bring about drastic changes within the immediate future, water table fluctuations control the 4.323.- wealth of species. These communities are therefore seasonably unstable and subject to the periodicity of continual or irregular hydrologic variation. Taken as a unit, the lowland hardwood complex represents a relatively homeostatic situation. The successional changes outlined in the proceeding are dependent, therefore, upon an assumption that future land use and drain- age patterns will remain essentially unchanged. S UWJR RY A considerable acreage occupied by lowland-hardwood forests in southern Michigan is considered to have a storage and stabilizing influence on ground water supplies. Since little is known about the composition, ground water hy- drology, successional relationships, and general ecology of these forests the present study is an attempt to provide preliminary information on these sub- jects. Measurement of the number, size and distribution of tree Species in the lowland-hardwood stands was accomplished using four sampling methods. A plot method was used as a standard for the study against which data were compared from other methods. Non-areal methods included random pairs, the point- centered quarter, and the variable plot-radius methods. In order to determine the relative reliability of the various sampling methods, simple correlations (r), were calculated between three parameters (basal area, frequency, and density), as determined from each of the methods. The relationships indicated by the use of coefficients of determination (r2) showed that any of the four sampling methods gave almost as much information about the three parameters as did the best method: The quarter method was re- commended for further use in lowland-hardwood ecological surveys. The same measures (basal area, frequency, and density) were used in de- scribing the composition of the stands selected for the study. Continuum in- dices were calculated for all lowland-hardwood stands. These indices indicated a rather complete coverage for the spectrum of lowland sites. The range of continuum values varied from a low 680 to a high 2430, Most of the stands were clustered within a segment of the range from 1800 to 2200. -189- Importance values calculated from the three parameters proved to give the best measure of species dominance. The leading dominants in the study were american elm, red maple, green ash, swamp white oak, silver maple, slippery elm, and American basswood. The effects of fire and pathogenic influences, windthrow and rooting systems on community structure of lowland-hardwood stands were discussed. The role of fire in the ecology of these forests appeared not to be of consequence in determining its vegetation. Windthrow and rooting of certain lowland tree species served to influence the eventual stand-age-structure of lacustrine forests. This condition was most evident in stands found on or- ganic soils. Thedevelopment of the understory and ground flora ranged from one extreme to the other on poorly-drained soils. As drainage conditions became more favorable the density of the flora increased. Differences in frequency and density of the ground flora could be partially attributed to the effects of seasonal flooding. An examination of the soil profiles showed that the majority of soil types were alpha-gleys which were poorly drained. Soils were analysed for total sand, silt, and clay content. Other analyses included moisture equivalent, loss an ignition, and pH. Differences in soil characteristics between the stands were evaluated statistically. Simple correlations were calculated between the importance values and basal areas of the ten most abundant species and soil physical properties. Two species showed some tendency to be found on soils with certain physical char- acteristics--black cherry and slippery elm. Because the presence of non-signif- icant correlations between soil characteristics in different horizons was great, it was suggested that future vegetation studies in the lowland-hardwood land type should ignore the measurement of physical soil parameters. 1.0a kl . :rv.:sz:r«rn;'ttm..lm inlormation on water table} 600:“ coultituc sunstantj;ally reduceth as a means 0? depicting the behavioral pattern of lowland—hardwood 19'- ests tne stand to stand relatiorshlps based on vegetation SlmlldTIL;PS and {szilv>nnuultul rm;as\nw&s werx‘ inclzjde(l in :1 sernxn; of ;;ra:nu=. lrzing {Err ctr~tixux;w as an index of similarity has not unlike the figurts derived from a iormula method. Jflvtefislon in lowland-hardwood stands implied by tne most signilicawt gnvironmvntal measurements was represented in a dimensional figure. 10. ll. 12. 13. liTHRATURE ClTEU Afansiev, M. 195?. The Bitterlich method of cruising -- Why does it work: Jour. Forestry 55: 216-217. Ahlgren, C. E. 1959. 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APPENDIX I ~205- TABLE.»29 Profile means for alpha-gley and betaobley soils, showing differences between soil texture, moisture equivalent, loss on ignition and reaction.8 Soil_grogp Ehysical and chemical parameters Soil series, __ Content 0f ‘_ Moisture Loss on and stand sand Silt clay equivalent ignition pH ............... percent-aa-—--—-------- number Bli'l'A-GLEYS 16690 57.1% 24.95 17.90 18.12 5.53 7.29 .;:%ks 77.3' 12.85 9.80 14.93 4.19 7.13 1:; 1'1: #:chr 50.7: 13.05 16.25 23.49 5.18 6.98 Ce:esco 67.75 19.00 12.75 25.48 4.81 7.98 “170 F.I.3'lL'\-CLF.YS Brookston 52.60 29.20 17.65 29.35 11.66 6.77 Elilord 72.25 13.96 13.55 26.10 10.63 6.71 P iiéim. 45.05 31.35 23.60 30.12 10.74 7.29 ;:ookst0n 47.15 31.95 20.90 25.10 6.52 6.49 cgfigctah 58.30 29.10 12.60 19.43 6.66. 7.12 Elli. 43.50 36.85 19.65 31.60 7.80 7.32 :gégwa 54.00 29.55 20.45 27.74 4.76 7.93 gilford 68.75 20.20 10.95 15.89 4.41 7.73 u4s 1.5.0. .05 8.70 5.33 5.02 7.23 2.98 1.63 1.5.0. .01 11.60 7.08 6.66 9.58 3.96 2.15 5.arranged according to soil group and the continuum placement of lowland hardwood stands. 131111.13 . - 31’) , Horizon means for beta~gley soils deveIOped from glacial drift, showing differences in soil texture, moisture eq- . . . a uivalent, loss on ignition, and reaction. Physical and chemical parameters. Soil type, Soil ___Content of Moisture Loss on stand horizon sand silt clay equivalent ignition pH --------------- percent--------------- nighct Locke A1 57.0 29.2 13.8 23.7 11.0 7.0 2“”4y loam A2 53.7 fi6,6 14.6 20.7 8.5 4.7 B1 63.? I-“‘-.8 18.0 12.5 3.1 7.‘ 82 56.2 21.0 21.8 ' .l 2.8 7.0 C 30.' ”7.0 21.3 1’.8 2.4 ‘..‘.?.:‘;i:..‘; A1 81,“ 17.0 6.2 :‘".0 10.7 0.7 ummy sand A2 73.1 4.4.0 7.2 1.".0 6.2 6.) H198 Bt 85.5 9.0 5.5 5.0 1.4 7.4 AZBt 79.3 9.5 11.2 10.5 1.0 7.3 C 61.3 13.7 19.0 10.2 1.8 7.5 Conover loam All 45.5 40.0 12.5 36.6 11.3 6.7 1 A12 48.0 32.8 13.2 30.0 7.2 6.7 Bl 48.4 32.3 19.3 19.4 2.6 b.’ 82 50.3 30.5 19.2 18.2 2.1 6.7 C 59.3 23.7 11.0 13.3 2.7 7.8 1;}.ij .05 113 "8.8—1.4 10.6 6.8 ‘6" L.».D. _01 NS 11.2 10.7 13.4 9.0 .7 7farranged according to soil type and the continuum placement of lowland hardwood stands. ~ 35 horizon means for 21.911.14.111); soils da'~\.'elt1ped from glacial drift, showing differences in soil texture, -moisture equivalent, loss on ignition and reaction.8 Soil type, Soil Content of loisture Loss on stand horizon sand silt clay equivalent ignition pH * ............... pe rce nt --------------- n umbe r Brookston All 56.7 31.8 11.5 47.3 29.6 6.3 sandy loam A12 51,5 34.5 14.0 49,2 19.6 6.2 A149 8218 50.6 31.2 18.2 16.6 3.5 6.7 8228 54.8 24.7 20.5 17.2 1.9 7.0 Cg 52.0 23.8 24.2 16.4 3.7 7.7 Gilford A1 66.7 24.3 9.0 66.2 35.9 6.0 sandy loam 681 79.4 12.6 8.0 17.9 6.8 6.3 L100 GBZ 76.7 10.3 13.0 13.2 3.8 7.0 G83 66.1 11.1 21.8 20.2 4.3 6.9 DE 72.1 11.6 16.0 13.0 2.4 7.5 Pewamo loam A11 74.5 36.3 16.2 46.4 23.0 7.1 058 A12 43.8 33.0 18.7 36.4 15.9 7.2 8218 48.0 28.3 23.7 23.3 7.1 7.4 8223 35.8 32.0 32.2 25.8 4.7 7.3 Cg 45.5 27.3 27.2 18.9 3.1 7.5 Brookston A11 49.0 39.0 12.0 36.8 12.4 6.2 loam A12 74.5 36.0 16.5 28.2 8.6 6.0 M268 8218 48.5 31.5 20.0 22.0 5.9 6.5 8228 4.15 30.7 27.8 21.3 3.7 7.7 Cg 49.5 22.4 28.1 17.3 2.0 7.1 Sebewa loam A 51.6 33.2 15.2 49.9 13.2 7.5 W48 G 1 54.6 24.0 21.4 28.2 2.8 8.0 G821 45.6 26.7 27.7 26.7 3.1 7.9 6822 52.6 24.2 23.2 23.2 1.1 8.1 Dg 65.8 19.5 14.7 10.8 3.7 *8.3 Gilford A 66.8 25.2 8.0 29.4 11.9 7.5 sandy loam G 1 66.1 22.1 11.8 15.3 3.7 7.6 W43 032 67.5 18.5 14.0 15.1 2.6 7.7 083 66.2 20.0 13.8 12.5 1.8 7.7 DE 77.0 15.2 7.8 7.1 2.1 8.2 1.5.0.,05 NS 8.8 8.4 10.6 6.8 .6 L.S.D..OL_* NS 11.2 11.2 13.4 9.0 .7 3 arranged according to soil type and hardwood stands. the continuum placement ofvlowland horizon for aiinvial 60113 showing differences 1 NJ ‘_) in soil texture, moisture equivalent, loss on ignit- ion, and reaction.3 Physical and chemical parameters Soil type, 5011 Content of Moisture Loss on stand horizon sand silt clay equivalent ignition pH - --------------- percent? -------------- numbe; 7 Ceresco A1 56.0 30.0 14.0 43.2 10.5 7.5 sandy loam CI 60.2 25.5 14.3 32.9 7.1 3,0 M19L C2: 62.7 21.0 16.3 24.4 3.3 8.0 C3g 73.0 14.6 12.4 19.7 1.5 8.1 C43 78.0 6.3 6.7 7.3 1.2 8.4 Cohoctah A1 61.1 30.4 8.5 29.7 16.5 6.6 sandy loam 61 57.8 32.9 9.3 23.1 8.5 6.7 M17C 02 55.6 29.7 14.7 15.6 4.1 7.5 G3 55.6 28.9 15.5 15.4 2.6 7.4 Cg 61.4 23.6 15.0 13.5 1.6 7.4 Sloan loam 91 42.4 42.0 15.6 51.4 17.1 7.1 'MZZS C18 43.4 413 15.3 37.0 12.3 7.2 C2g 43.4 34.5 22.1 23.4 4.5 7.5 ng 41.7 34.5 23.8 24.1 1.5 7.5 C48 46.7 32.0 21.3 22.1 3.6 7.4 L.S.D. .05 NS 8.8 8.4 10.6 6.8 .6 L.S.D. _;QL, NS 11.2 10.2__ 13.4 9.0 .7 3 arranged according to soil type and the continuum placement of lowland hardwood stands. 1&8L8.- 3;, Proiilc and horizon means tor organic soils, showing differences in loss on ignition and reaction.a - -——- .. ‘- .~- -d—‘C\ Chemical‘parnmeters Soil type, 8011 Loss on 1&n1tion __“__~2H stand horizon profile - horizon pro{i.oiifi?§;33 Liming-15:1 muck 01 59.4 5.6 .311". 1.. 02 62.1 6.1 03 41.7 54.8 6.24 6.3 04 29.2 6.4 mg 3.1 6.0 tinnwnod muck 01 52.3 6.4 1.1-.11. 02 59.6 6.6 03 38.77 54.5 6.76 5.6 0:. 25.2 7 6 D8 2.2 7.7 Carlisle muck 01 42.1 6.9 V20C 02 58.2 6.7 03 32.58 51.8 7.27 7.0 04 7.0 7.8 05 4.0 8.0 Rifle peat 01 63.6 4.5 M26R 02 73.6 3.9 03 44.15 71.5 4.50 3.8 04 . 9.3 500 D8 2.8 5.2 L.S.D. ..05 3.34 13.2 1.63 .6 L.S.D. “QQ1 4.47 17.4 __ 2.15 .7 3 arranged according :6 soil type and the continuum prCement of lowland hardwood stands. ll .3'! TASLR.~%..Horizon means of underlying nuterials of the beta-gley, alpha- gley, and organic soils showing differences in texture. a Soil group Soil series, Soil Content of and stand horizon sand silt clay ----- percent--------- locke C 50.7 28.0 21.3 M19L Spinks C 61.3 19.7 19.0 M198 Conover C 59.3 23.7 17.0 A14C Ceresco C98 87,0 6.3 6.7 6117C ALPHA-GLEYS Brookston C8 52.0 52.0 24.2 A148 Cilford Dg 72.1 11.6 16.0 LlOG Pewamo Cg 45.5 27.3 27.2 W58 Brookston Cg 49.5 22.4 28.1 M268 Cohoctah Cg 61.4 23.6 15.0 M17C Sloan C48 46.7 32.0 21.3 M228 Sebewa Dg 65.8 19.5 14.7 W48 Cilford Dg 77.0 15.2 7.8 W43 ORGANIC SOILS Linwood Dg 53.4 23.0 23.9 A14L )- .1 i '\ ’ ‘ '17-'51) 1.11:. . 51:1 _ 131‘) {11. i nued Linnwood DE 77.1 12.0 10.9 LIOL Carlisle Dg 81.0 13.0 6.0 VZOC Rifle Dg 43.9 25.5 25.6 M26R L.5.U. .05 17.6 9.1 11.3 L.§.D. .01 23.3 12.0 15.0 hardwood “'h-W .ma—“n -. v- -M arranged according s t:1vitls . a... o—- 4—-———— o-q .a. .- to soil group and the continuum placement of lowland VIT\ ixilgex‘i lxiasic: 15r:;u:it (kuujidate for tine Degree of Doctor of Philosophy Dissertation: The Lowland Hardwood Forests of Ingham County, Michigan: their Structure and Ecology. Maior studies: Forestry Minor studies: Soils and Statistics BiOgraphical Data: Rirthdate: January 3, 1928, New York N.Y. Undergraduate studies: McNeese State College, l949,-1950. ionisiana State University, Forestry Dept., 1950-1953. 8.8. 1953. Graduate studies: Louisiana State University, Forestry Dept., 1954. M. S. 1954. Michigan State University, Forestry Dept., 1959-1962. Experience: Research biologist, Louisiana Wildlife and Fisheries Commission- Fish and Game Section., 1952-1953. Refuge biologist, Rockefeller Refuge, Grand Chenier, La., 1954. Assistant Professor of Forestry., McNeese State College, Dept. of Plant Science, 1954-1959. Graduate Teaching Assistant., Michigan State University, Forestry Dept., 1959-1962. Assistant Professor of Forestry., McNeese State College, Dept. of Agriculture, 1962 to present. Member: Alpha-Zeta X1 Sigma Pi Society of American Foresters ~213- 11001.1 USE 01.31! _ a ’ 71" ,’ 1 : '7’ . l . 1., H I , ,1