| E I- “r r ‘1'..—.. A" .‘ A. v I , 1" l ‘ ’" ’,.. -n .1 .» A‘I~. E U ‘ x115§1‘b;'qu‘i'3‘u.s.~i :V ,5; $27!,1 “it?!” .~ aw. 4;. 2qu M . ,,.uuw: P. ,_ ... L 2 f’ 'F‘E'fl A , F . 12" _..'.: 1“” 5"}? II V ' I. 4 "'2’. .d‘ I'V K'K'J ”rm 3.x H: ..< .\—'4 . I ‘ h. wfi-«a Java M." ”1:22;; “32%: \— V ‘ . o L _ , aw}, ‘3 Q ,: up: .. -. _. i ‘ 1;?“ mac?" .ux“ ; ' t; as: r V’ . - ' , . - c- _. .. . 5 .j 3.; . ,, ‘ 3 1 ~ f. L H‘. _ . . ;, _ ..~ r. ' ~zpr§1£555fi9r=uf .1 _L .. “3:3... I ‘ '1‘. :‘AHMW ‘ ‘ H 1' ’- ' ‘ k ‘2? r1?” ’ I i‘vhqfi. 1 L l .1 n‘uanAu-Lul 4-,” A'fli" ' 1’ , l I". u D 'r‘ 2. «r a» 3:. :wwm 1" LSL'JF‘V‘" ' W4 An :0 HJ': ‘ .; ~.‘y.\1~:{.x. 4-“... V?‘ I. Id“- -fl- \- ' This is to certify the! the ‘ . ; the“: entitled A ‘Immmmcamnmncnw ‘1 mp' I.” m 3 3,7 l K: 7 has been accepted towards fulfillment - » of thevrequiremems for . , 4- ? fldegree tum. ’ 3 ! e Major professor Q h 1 '0 “by: l t . Date-m— if!!! "5 . . w‘x Ink“ ___5_ _____________..___ MSU LIBRARIES RETURNING MATERIALS: Place in book drop to, remove this checkout from your record. FINES wiII be charged if book is returned after the date stamped below. ‘i '3 f3. “" J J L U 'l 4 MAY 2 5 2003 6 2m R U '7‘; 1 3 THE OAK UPLAND CONTINUUM IN SOUTHERN MICHIGAN By George Nyman Parmelee A DISSERTATION Submitted to the School of Graduate Studies of I-Iichigan State College of Agriculture and Applied Science in Partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Botany and Plant Pathology 1953‘ THESIS' T22: 02:: been cayenne: 1:: WWW??? 22162222222:- George Lumen Pamelee AN AmTRfi-JZT Submitted to the School of Graduate Stm‘iiee of liichigan State Coll-ego of Agriculture and fimlied Science in partial Wilbert of the mute-cent e for the degree of BETTIE ‘0? Hill; 133?}?! Lopez-teem of Botany and Plant Fethology J! Although oak'uzlnnd forest 09323135 tore orére:w .0 ecrerjo tLan 6:15? 0"3381' notm'al MIMI-l. ty in emit-hem 2:3cz3: 3:52: 3 reel mad Iron: 1 {‘33, t‘fie «almost car may roux-1 o as the lorot Tmom 3.31- :3t our: 2.3.3:: Zty of tire moi, m. Atom-(11:31:, a misizary otjectivo of 1.13 o to “.13- !wo ha—on to record hw'phytoooclological met Bode t l-o'vesculzr corponenta of a regional series of otendo representative of the oak uplcnd ton. In all, (max-at data for troea, mambo 12.32.1‘ Lorre 9m: 3:”. 221333.13 6.13.331“: rated through.l7 oountioe are presented. The extrtI 3oly diverse ejecioa reg?” mute 15.023 halos-ted by fro-2o date waisted that tho eontlnum concept as elamratcfi by Cartie 2.2a Iclntoah (1°51) would provide the moat realistic orirm tnti on of’etoxis for We: of comma-icon and emtraat. Iz'::~.or‘..:~nco whee zero them- fore canned for each tree apeoieo and these Velma were teamed by climax edepmt .on motors to yield contimzm 1:26.411 mater-a rem-2:13;: from 751 to 19:23. Data mfmomatlng indivi.‘ .aell cwgmseate of the ve rims: 9y: maino of end: stand were charted in e eoquence determined by t: .eoo cortisrmm hnéez uuobere. The resulting curves were interpreted.ee.graphlcal expressions of tolerance ranges. Io tondoaey for eucn rnrg ea to occur in patterns eugI eating discrete oombinet -one of ooocioe was ovidczt in any m1... ( Ratl'm, the charts booed on tree dete 1.31mce a that even the moat contrasting etzmdo difformi 192338 by kind of douimnio than I: martian of dwinentofl The majority of 33-3-43. and herb epocioe were found to eroe’def inite correlation with continuum i“? .eI sequence. Among tE-zoee which do not are certain of the more u‘aicplitmo cardamom-ate and tho... which occur so spend... sally t‘. at tromlo are Melanin. Species of cit her of these cotcyorioo are held to have fl Georgo ‘55. Pamela «- 2 no utility as 13365.3.th of site. Too leading noiro and trim of dam-2.3.3133 in on oh atmd wore cor- related with cumin-3:333: index ammo m in orator to determine the volldity of each as boom for mix-3.4231 closefll’icntlon of 09.3: upload coma-o. For this purpose two lowing; chain-moo are hold to be aumrior to tome because of tho fewer mm:- of 123333333311 coted cmbmntiono and the no.1» rm “looping in scale position of the stood groupings character- 1396 13 the some 1.633631% éminanto. The ace-.19 sector bemoan 11313634! values at 1350 and moo um toototivoly recognized as a mason-y ’aegar. 33:23.5; stands having black ow: no one of the too loaf: “I; den“ «2. to from those in which red oak has. a similar position... Pow women laboratorioo” a: 3.3.9 for» anvislomd to- Hagar (1-3-51), in um ab. 3.13: influence of one-factor mimontal mfiation on con- than: 3.11%: amber cow-.31 to nomad, were row/.1. S no dmlorrod on aim of southern man had aitmiflcmzuy lower continua: 1:33 or: 1131:3331?th those analogues! m slopes of northern aspect. Except on north slopes, mm distur’mmoo woo likewise mflocmd by a signifip cont 103an in omtixmufo index anr. Doopito effect We vol-1331.331 of other manl fa com, a {renew-J. comlatim 3.33:3 m'aad bot-won continua: ind-m: nontor and coil mama's. Data 0W1 in this: omcly 1mm cmbbmé vi to imi’ououzticm from other 5033233308 in an arm to toot the validity of the var}. out! «31:32:93 hypotheses as an filled to tho on}: upland cont 13mm in southem .‘r’i ci-‘zfgon. 7339 (:15; .3233: pattom m'potheoia of Lfizittef-‘er (1953) was found to agree mot cleanly with camel-wane facts. Goargze 1:3. Featmleo - 3 , "fit"? -sav “A: E...‘ bi has" all.) Cut-tic, J. 7., and H. P. 2-fc1rtoah. 1"»51. fin wit-ma fore-at cmtimm: 1n the prairie-forest border region at incl-main. lflmlow 32: [flip-1,736. Gyscl, L. $3., and J. L. Erma. DEB. Oak sites. in sunbeam Eflchimn: their classification and «valuation. Erich. 8mm College Agr. 373;). am. Tech. Bull. 236. 57 pp. Rajar, .T. 1951. A fmmioml factorial same?) to phat 600133}. Ecology 32: 3922-412. Emitta‘ter, R. 2?. 1"?53. A mnsiacmtion of the climax thou? «- the climax as petulstion and pattern. Bic-'31. I—‘aonogr. 23: Alp-753i. TABLE OF CONTENTS ACKNOWLEDGMENTS ........................................ . IJST OF TEXT FIGURES .......................................... viii LIST OF TEXT TABLES ........................................... xii LIST 01“ APPENDIX TABLES ......................................... xiv BHRODUCTION .................................................... l IJTERATURE REVIEW ............................................... 3 EEJREGION AND ITS HISTORX ...................................... ll PhySi-ography OOOOOOOOOOOOOOOOOOOO0.0....OOOOOOOOOOOOOOOOOOOO 11 Primary Configuration Features ........................ 11 Secondary Configuration Features ...................... 12 Clj-Inate OOOOOOIOOOOOOOOOOOOOOO0.0.000...OOIOOOOOOOOIOOOOOOOO 14 80118 OOIOOOOOOCOOOOOOOOOOOOOICOOOOOOO00.0.0.0...O...0...... 15 Genera-vegetation .0.00000000000000000IOOIOOI0.00.0.00.0... l8 POStglaCiaJ- HiStOI'y O O O O O O O O O O O O O O I O 0 O O O O O I O O O O O O O O O 0 O 0 l8 Pro-settlement vegetational Pattern ................... 21 Postsettlement History 0 O I O O O C O O O O O O O C O O O O O O O O O O O O O O O O % MEEDDS 00000000000000.0.0000000.I00000000000000.0000.00000000000 27 Reconnaissance ............................................. 27 Stand Selection ............................................ 27 Quadrat Location ........................................... 30 Collection of Field Data ................................... 31 Qllalitative Data 0 O O O O O O O I O I 0 O O O O O O O O O O O O O O O O O O O O O O O O O O 31 Quantitative Data 0 O O O O O O O O O I 0 O O O O I O O O O O O O O O O O O O O O 0 O O O O 32 VOUCher Plant COlleCtions O O I O O O O O O O O O O O O O O O O O O O O O O O O O O 33 Tree-went Of Data 0 I O O O O O O O O O O O O O O O O O O O O O 0 O O O O 0 O O O O O O 0 O O O O O O 34 malytical Indices 0.0.0.000...OIOOOOOOOOOOOOOOOOOOOOOO 34 Importance value cocoooooooctooocoocoooooccc ccccc o 34 Significance value 0.000000000000000...000000.000. 35 n.4- 014288 DF val-ue OOOOOOIOOOOOOOOOOOOOOOOOOOOOOOOOOOOOO... Simple frequency OOOOOOOOOOIOOOOOOOOOOO00.00.00... smthetic Ind-ices OOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOCOO Presence ......................................... Aggregate constance .............................. Summation frequency .............................. Relative DFr value ............................... Graphical Presentation 0......0.0IOOOOOOOOOOOOOOOOOOOOO MULTS AIJD OBSEVATIOIIS OIOOOOOOOOOOOOOOOOOOOOOOOOO0.0.0.0.0.... Provisional Climax Adaptation Numbers for Southern Michigan ................................................. The Continuum Index Scale . Components of the Oak Upland Community ..................... Tree components O...0.0.0.0....OOOOOOOOOOCCOOOOOOOOOOOO I‘Iean Area. BalationShipS O O O O O O O O O O O O O O O O O O 0 O O O O O 0 O Ilnpor‘tance value 0 O O O O O O O O O O O O O O O O O O O O O O O O O 0 O O O O O O Summation Indices for Tree Reproduction .......... Significance values 0 O O O O O O O C O O O O O O O O O O O O O O O 0 BF values for class 2 reproduction ......... Class 1 DFr values 0 O O O O I O O O O O O C O O O O O O O O O O O I O Shrubs andI‘IOOdy Vines OOOOOOOOOOOOOOOOOOOOO0.0.0.0...O Shrub Species Accidental in the Oak Upland COWity 000000000000oooooocooocooccoocoocooo00 The I'bl‘e comic!) ShI‘Ub components 000.000.000.00... Mean area-frequency correlations ............ Relative significance of the indices of shrub representation ................... Species exhibiting no obvious correla— tion with stand sequence .................. Shrub representation displaying correla- tion with stand sequence .................. Herbs 00......0....00......OOOOOOOOIOOOOOOOOO0.0...0... Comparison of Data from Different Sizes of Qlladrats 0.0...OOOOCOCOOCOOOIIOOIOIOO0.0.0000... SWOieS-area relationShipS O O O O 0 O O O O O O O O O O O O O Frequency-area relationships ................ iii. 39 44 46 47 50 57 58 65 69 '73 '74 75 75 86 89 93 96 96 98 98 Ecologically Significant Herb Species ............ Species exhibiting no obvious cor- relation with stand sequence .............. Herb species correlated with stand sequence .................................. Correlation between Leading Dominants and Continuum Index Sequence ........................................... Continuum Index Numbers as Measures of Environmental Influences ............................................... Influence 0f Slope .00000.0000.000000.00000000....00000 Influence of Slope on Tree Dominants ............. Influence of Slepe on Other Community Components 00.00.00.000...00000000...0.0...00000 Tree reproduction ........................... Shrubs 0.0.00.0000.0 0.0000000000000000000000 Herbs 00.00O.0...000..00.0..00000000000000000 Influence Of 8011 00000000000000000..0000000000.0.00000 Soil Variation within Single Tracts .............. Correlation between Soil Type and Con- tinum Index Nlnnmr .00....0...000.00000...0.... Influence Of Past Utilization ...0.0..00...00..00.0.000 Correlation between Measures of Synusial Diversity and Continulml Index Sequence 00000000000000.0000...0.00000 n16 Herb sym-lSia .000...0.000000000000000.....0.0.00000 The Shmb synuSia 0.0.00..00.0.00....000.000...00....00 DISCUSCJIOIJ .0000...0.000000.00....00000..0.0.00.0000.00.....00000 The Vegetational Continuum Index as a Basis for Classification ........................................... The Continuum Index Scale as an Index of Site and Measure of Tolerance Range ........................... The Concept of Climax Adaptation Number .................... Successional Status of the Oak Upland Community ............ The Oak Upland Community as a Test of Climax Hypothesis ................................... Past and Present Trends of Reproduction in the Oak Upland chnmunity .. iv. 101 102 109 117 121 124 124 127 127 127 128 128 129 130 132 137 138 142 150 150 152 155 157 158 167 SUIYQ'MMIDCOIJCLUSIOI‘JS OOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOO....0 LITWTUBE CITED OOOOOOOOOOOOOOOOO0.0000000000000000000000.... APPHJDH OOOOOOOOOOOOOO... ....... O ..... OOOOOOOOOOOOOOIOOOOOOOO 183 186 196 V. vi. ACKNOWLEDGMENTS It is a pleasure to express my sincere thanks and appreciation to Dr. William B. Drew, Head of the Department of Botany and Plant Pathology, Michigan State College, under whose kindly supervision this study was conducted. His persevering interest and patient assistance during the planning and execution of the investigation and the preparation of the thesis have been invaluable. I am also greatly indebted to the following: Dr. Charles L. Gilly, Assistant Professor of Botany and Curator of Plant Collections, Michigan State College, for his willing assistance in taxonomic matters, his unfailing interest, helpful suggestions and constructive criticism during the course of the study, and his critical reading of the menu, script; Professor-emeritus of Soil Science Jethro O. Veatch who con- tributed generously of his time and vast fund of infermation concerning the soils, land and associated vegetation of Michigan; Mr. Philip G. Coleman, Science Photographer for the Experiment Station, Michigan State College, for his helpful suggestions, willing cooperation and superlative technical aid in photographic matters. Unless otherwise indicated, the photographs included in this dissertation represent his work. Sincere thanks are also extended to Dr. Leslie‘w. stel, Associate Professor, Division of Conservation, Michigan State College, and Mr. John L. Arend, Research Forester, Lake States Forest EXperiment Station, as well as various district foresters, for helpful information concerning vii. location of stands. Acknowledgment is also due the staff of the Lands Division, Department of Conservation, State of Michigan for their many courtesies in providing working facilities and access to the records of the original land survey for Ingham County. Finally, full credit is due my wife, Margaret Wilbur Parmelee, for unfailing encouragement and inspiration as well as uncounted hours of field and clerical assistance. 10. 11. LIST OF TEXT FIGURES Map of southern Michigan showing location of stands Studj-edOOOOOCCCOOOOOOOOOO0.00.0.0...OOCOOOOOOOOOOOOOOO0.0... Lowland form of shagbark hickory growing in association with red oak and swamp white oak................ Old growth bur oak of the type which, in Open stands, once characterized the bur oak plains and prairie environs in southern.Michigan................... Continuum index scale for the oak upland community in southern Michigan showing scale position of the Sta-{Ids StudiedOOCOOOOOO00.00.000.00...OIOOOOOOOOOOOOOOOI Importance values of four major dominants in indi- vidual stands arranged according to continuum hidex nmber...00.00.000.000000COOOOOOOOOOOOOOOOOOOO00...... Importance values of 14 accessory dominants occurring in individual stands arranged accord, illgto continlm-In index nmberCOOOOOOCCOOCOOCOCOO0.00.0000... Significance values for five species of low climax adaptation number occurring in individual stands arranged according to continuum index number................ Significance values for six species of intermediate climax adaptation number occurring in individual stands arranged according to continuum index number......... Significance values for seven species of high climax adaptation number occurring in individual stands arranged according to continuum index number................ Class 2 BF values for ten species of low to inter- mediate climax adaptation number occurring in individual stands arranged according to con— tinuum index number......................................... Class 2 DFr values for 11 species of intermediate to high climax adaptation number occurring in stands arranged according to continuum index number................................................ viii. 43 45 55 56 59 61 67 12. 13. 15. 16. 17c 18. 19. 20. 22. 23. Class 1 BF values for nine species of low to intermediate climax adaptation number occurring in individual stands arranged according to continuum index number..................................... Class 1 DFr values for nine species of intermediate to high climax adaptation number occurring in individual stands arranged according to continuum index number..................................... Index values for four more ubiquitous shrub species showing no evident correlation with a stand sequence determined by continuum index nmberOOOOCCOOOOOIOCOOOOIOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOO0.. Index values for four shrub Species showing no evident correlation with a stand sequence determined by continuum index number....................... Index values for five shrub species showing no evident correlation with a stand sequence detenmined by continuum index number....................... Index values for five less common shrub species showing no evident correlation with a stand sequence determined by continuum index number.............. Index values for seven uncommon shrub species showing no evident correlation with a stand sequence determined by continuum index number.............. Index.values for five shrub species evidently most abundantly represented in the more xeric stands of a sequence determined by con, tinumm index number........................................ Index values for five shrub Species evidently correlated with a stand sequence determined by continulm ind-ex nmberOOOOOOOOOOOOOOOOOOOCOOOOOOOOOOOOOO Index values for five shrub species evidently most abundantly represented in the more mesic stands of a sequence determined by continuum index number............................................... Relation between number and.magnitude of frequency records obtained from two sample areas of dif- ferent 8126....OOOOOOOOOOOOOOOOOOOOO0....OOOOOOOOCOOOOOOOOO Curves of frequency for two species of similar habitat requirement showing similar trends of representatiOnOOOOO00......OOOOOOOOOOOOOOO0.0..00.0.0000... 7O 76 78 79 81 82 83 85 91 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. Frequency values for eight more ubiquitous herbs showing no evident correlation with a stand sequence determined by continuum index number.............. Frequency values for nine common herbs showing no evident correlation with a stand sequence detemined by continum index n‘mmr. O O O O O O O O O O C O O O O O C O O O O 0 Frequency values for 11 less common herbs showing no evident correlation with a stand sequence determined by continuum index nmnerOCOOOOOOOOOOOOOOOOOOOOOOOOOOOCOOI0.0.0....0000...... Frequency values for 12 uncommon herb Species showing no evident correlation with a stand sequence determined by continuum index number.............. Frequency values for 14 herbs evidently most abun- dantly represented in the more xeric stands of a sequence determined by continuum index number............ Frequency values for 11 herbs evidently correlated with a stand sequence determined by continuum index mmerCOOCOCOCCCCOOCOO0.0...OCCIOOOOOCOOOOOOOOOOOOOO Frequency values for ten herbs evidently most abundantly represented in the more mesic stands of a sequence determined by continuum index number............................................... Frequency values for 11 herbs evidently most abundantly represented in the more mesic stands of a sequence determined by continuum index number............................................... Frequency values for fourteen herbs evidently most abundantly represented in the more mesic stands of a sequence determined by continuum index mwr0000000.0..OOOOOOOOOOOOOOOOOOOOOOOO00.0.0000...COO... Relation between index position and the two leading Stand dwinantSooooooooocoo-cocoa...00000000000000. Relation between continuum index position and the three leading Stand daninantSOOCOIOOOOOOOOOODOOOOOOOOOOO... Relation between soil type and continuum index position of associated stands.............................. 103 104 105 106 110 111 112 113 114 118 119 131 36. 37. 38. 39. 42. 43. View along the line fence between stands 16 and 22 showing older timber at the right and younger growth, including some of c0ppice origin, at the left................................ Relation between two measures of diversity of the herb synusia and continuum index sequence.............. Relation between three measures of diversity of the shrub synusia and continuum index sequence............. Relation between total density records for the shrub synusia and continuum index sequence................. View of an oak woodlot pasture Showing effects Of prOlonged overgraZingOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOO Depauperate clone of Juniperus communig persisting under closed canopy oaks........................ Old growth white oak with enormous lower branches forming a wide-spreading crown.................... Seedlings of Qgercus rubra develOped from acorns concentrated by gravity on the floor OfEmOj-St raVineOOOOOOOOOOOOOOO0.0...OCOOOOOOOOOOOOOOOO... Sprouts developed at the crown of a class 2 individual of Qgercus albg which evidently died back in consequence of inability to compete under closed canOpy conditions..................... Reproduction of 93ercus velutina under a seed tree growing in a clearing................................. Reproduction ofIQuercgg velutina established on bare mineral soil in an abandoned field adjOinj-ng a. blaCk 085k StandOOOOOOOOOOOOOOIOOO0.0...0..00... Heavy shade cast by a second canogy of mature cornug florid-é.0.0I0.00000000IOOOOOOOOOOOOOOOO0.0.0.0000... Trees of obvious Sprout origin as indicated by mutiple mle developmentOOOOOCOCOOIOOOOCOOOOOOOOOO00...... xi. 134 139 143 145 149 165 169 171 172 173 176 177 II. III. VII. VIII. XI. XII. LIST OF TEXT TABLES Provisional climax adaptation numbers for tree species occurring on upland sites in southern LiiChigaDOOOOOOOOOO00.000000000000000.OOCOOOOOOOOOOOOOOOO. Mean area relationships by size class for total tree representation................................ Mean area relationships by size class for the single most numerous Species in each stand........... Stand occurrences of 100 percent frequencies W Size ClaSSeSOOOOOOOOOOOOOOOIOOOOOOCOOOOOOOOOOO00.0.... Stand occurrence for tree components of the oak upland community by Size class....................... Tree Species most frequently sharing in control of the oak upland community of southern I'IiChianOCOOOOOOOOOOOCOOOOCOOOOOOOOOOOOOOOOOOOOOOOOOOOOOO Relation of quadrat size to number and magnitude Of frequency record-8.00....00....OOOOOOOOOOOOOOOOOOOOO... Synthetic indices for those shrub species showing no clearly evident correlation with continuum index seq‘LlenceOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOO0.0. Frequency data for the 42 most common herbs recorded in sets of 10 quadrats of three different Sizes established in each of seven stands located within a circle of one-half mile radius......................... Relation of quadrat size to number of herb Species recorded in seven stands encompassed by a radius Ofone-half.11111600000000.0000...0.00.0000...00.... Relation of quadrat Size to average Species density of herbs recorded in seven stands encompassed by a radius of one-half mile............................. Synthetic indices for the more common herb species showing no evident correlation with stand sequence....... xii. 49 51 52 54 84 92 97 99 99 107 XIII. ‘Variations in topography and past land use of seven stands of a complex in which soil type, macroclimate and species complement may be considered constant or to vary ineffectively............. XIV. Net positive differences in weighted importance values between indicated stand pairs contrasted to illustrate influence of slope......................... XV. Net positive differences in weighted importance values between indicated stand pairs contrasted to illustrate influence of past utilization.............. xiii. 125 126 135 I——XXXVI. XXXVII. XXXVIII—LXXIII . LXXIV. IXXVII. LIST OF APPENDIX TABLES Summaries of tree data for individual stands, with exact locations for eaCh standOOOOOOO0.0000IOOOOOOOCOOOOO0.00.0.0. Summary of composite tree data for Stalldsaand 31...0.0.0.0...OOOOOOOOOOOOOOOOO Summation indices of trees for iIldiVidual Stands...OOOOOOOOOOOOOOOOOO0.0.0... Summation indices for trees of Stands21f811d31000000000000.0000.oooooooooooo Number of occurrences of tree Species inaJ-lStand-SOOOOOOOOOOODOOOOOOOOOOOOOOOOOOIOO Number of occurrences of shrub species iII all StandSOOOOOO0..OOOOOOOOOOOOOOOOOOOOOOOO Humber of occurrences of herb Species inall StandSOOOOO0.0.0COOOOOOOOOOOOOOOOOOOOOO xiv. 197 233 234 270 271 272 274 'INTRODU TION Oak upland forest was an extensive and distinctive element of the Indmary vegetation of southern Michigan. Today, though probably more nearly obliterated than any other forest community, this type still occupies a greater aggregate acreage than any other natural plant com- nmnity. DeSpite its obvious importance in the original and present vegetation mosaic, the oak upland element currently ranks as the least known plant community of the region. No phytosociological papers dealing specifically with the oak upland community of Southern Michigan have been Imblished and the only maps in which the element has been segregated are that of Veatch (1928b) based on soil maps and those of Kenoyer (1930, 1934, 1940, 1943) based on land survey notes for eleven southwestern counties. The study reported herein was undertaken to obtain data that might be used in describing the oak upland community. The objectives, Scope and procedure were in large measure determined by the relationship tetween available information and the great areal extent of this commun- ity. Thus, in concordance with the views of Yapp (1922), it was felt that the greatest initial contribution would accrue from emphasis on extensive, rather than locally intensive investigation. If, for example, extensive phytosociological investigation should reveal the existence of Iepetitive aggregations of Species, then subsequent autecological and environmental studies would obviously be facilitated, as emphasized by Cumtis and McIntosh (1951: 476). Moreover, though detailed attempts to 2. demonstrate community interrelationships would obviously be precluded by the time factor, advantage could be taken of such fortuitous Oppor- tunities for pointing out coincidences and correlations as chanced to occur. Such observations might be expected to serve as suggestions for future intensive investigation. Specifically, the primary objectives of this study were twofold: first, to record by phytosociological techniques the vascular components cm a regional series of oak upland stands; and second, to organize the data thus obtained to demonstrate or refute the existence of discrete arrays of dominant as well as subdominant components. Secondary objec— tives were (1) to make such inferences relative to oak upland community Chnamics as were possible within the limits of a short period of study; and (2) to note, wherever evident, coincidences and correlations involving stand composition and environmental factors or factor complexes. LITERATURE REVIEW Darlington (1946) has supplemented a review of the history of botanical exploration in Michigan with an extensive bibliography of well over 300 citations concerning taxonomic or ecological studies on the higher plants of the state. The first reference to plant communities in southern Michigan was made by Lanman (1839). He mentioned (p. 251) heavily timbered land, barrens, white oak Openings, bur oak plains and prairies. The heavily tdmbered land was described as composed Of a variety Of species, includ- ing the combination Of sugar maple and beech. The barrens were depicted as "but thinly covered with stunted oaks." The white oak Openings com- priSed a great prOportion of the area Of the state and were described (p. 323) as composed of "white oaks interspersed with black and yellow oaks as well as hickory." They were repeatedly referred to as park—like in aspect, and lacking undergrowth except where they adjoined heavily tfimbered land. Lehman also provided generalized descriptions (pp. 282—- 291) of the natural vegetation of 22 southern counties. Numerous refer- ences to oak Openings, bur oak plains and prairies occur in these descrip— tions. Such emphasis doubtless reflects the tendency of the early road builders to establish routes through these types which were at once easiest to traverse and which occupied types of land most in demand by the settlers. In the preface to their catalog of the Michigan flora, Wheeler and Smith (1881) depicted the upland forests of southern.Michigan as consisting .VS H ;‘ i 4. of beech-maple and oak smiths, with one type locally shading into the other in response to textural variations of the glacial drift. Other early references to forest communities were based on rela- tively localized observations. In his classical report on the dune vegetation along the southern Lake Michigan shore, Coulee (1889: 384—385) recognised three types of dune forest. These were held to be detemined primarily by slow aspect and distance from the shore. In a study of the forests of Kent County, Livingston (1903: 39) recognized five upland calamities which be regarded as comprising a gradational series deter- mined primarily ty moisture relationships. These canmunities in order from most mesic to most xeric were: beech-maple, maple-elm-agrimony, oak-hickory, oak-hazel and oak—pine—sassafras . Concerning the problems of delimiting these types Livingston (p. 39) states that they "often merge gradually into one another so that in some localities it appears that there is a mixture of several of them." His report also includes a list of accessory species which are classified as to relative abundance in the different cosnnunities . Additional early observations were pub- lished concerning the upland forests of the Huron River valley (Brown 1905; Transeau 1905) and Wayne County (Brown 1917). More recent observa- tional accounts concerning the forests of the general region are included in the contributions of Binghsm (1945), Braun (1950) and Veatch (1953). A mnber of workers have mapped pro-settlement vegetation for all or parts of Michigan. Utilizing soil-vegetation correlations, Veatch (1928b) delimited generalized types of vegetation for the state as a whole. The maps of others were based on field notes coupiled during the original land survey. These reconstructions include (1) mall scale maps of the IQ v-I ”N n." ya ”I 5. Lower Peninsula which show the distribution of original swamp areas (Davis 1907) and gross vegetation types (Marshner 1946) , and (2) large scale maps showing the pre-settlment vegetation in considerable detail for 11 south- western counties (Kenoyer 1930, 1931., 191.0, 1943) . Wheeler and Snith (1881) regarded the flora of the Lower Peninsulawa as relatively homogeneous, stating that "probably thre e—fourths of our species are cannon to all sectors." Soon after, however, two areas com- parable to the tension zones of Griggs (1914: 47) were recognized and Species reaching northern or southern limits in them were listed. These comprised the Grand Haven area along the Lake Michigan shore (Bailey 1882) and the valley of the Grand River (Beal and Wheeler 1892). Veatch (1932) recognized a transition zone, based on pedologic evidence but character- ized also by transitional vegetation, centered to the north of these areas along a line from Saginaw Bay in the east, to Muskegon in the west. The ' northern boundary of the oak hickory forest was depicted as being reached in this zone, as were the southern limits of white spruce, and jack, red and white pines. This zone would thus seem to constitute an ecotone separ- ating natural regions of a scope comparable to the biotic provinces of Dice (1931, 1943), the natural areas of Cain (1947), the floristic areas of Resp (1947) and Egler (191.7) and the floristic provinces of Curtis and McIntosh (1951). The boundary between the Alleghenian and Carolinian biotic provinces (Dice 1931) falls entirely within the transition zone, as does the boundary between the beech-maple and hemlc ck-white pine- ncrthern hardwood regions recognized ty Braun (1950). The first report incorporating quantitative data on actual compo- sition of upland forest in southern Michigan was that of Quick (1924) who presented frequency data for 10 stands of the beech—maple type. Qiick 6. regarded the beech-maple community as an I'ecological species" of "defin- ite biotic canposition.” Moreover, he was evidently determined to depict "typical" composition as indicated by his statement (p. 221) that some species ”were so typical of a preceding association as to warrant their exclusion“ from the data. The tree species accepted by him as components of the beech-maple association were: Acer saccharum Carya ovata Fagus grandifolia Prunus serotina Tilia americana Carya cordifonnis Ulmus americana Juglans cinerea Fraxinus americana mercus alba Ostrya virginiana Tsuga canadensis Quercus rubra Liriodendron tulipifera Betula lutea Concerning the 15 species included in this list Quick stated (p. 221) that ”a few such as the white oak were scarcely typical, but occur so often that they were included as indices of the forest type." The first quantitative account concerned specifically with oak upland stands was that of Wood (1930) who studied five grazed woodlots on the W. K. hellogg tract in Kalamazoo County. Black and white oaks and pignut hickory were the leading dominants of these stands but were poorly represented in the reproduction classes, commising only 10 per cent of the total. In an inventory of 45 oak stands in Livingston and Genesee Counties, Monroe (191.3) found the proportion of trees over 15 inches at d.b.h. so low that the basal area of this class was exceeded by that of the 2- to 6-inch group. Young and Sholz (1949) found reproduction of the preddninant oaks and hickories in a Washtenaw County stand to be "almost negligible in amount.“ The most recent and comprehensive paper on the oak upland is by‘Gysel and Lrend (1953), who obtained tree data from a total of 118 sample plots in 61 oak- stands. The plots were grouped into five site 7. classes based on varying canbinations of soil texture, position of moist layers in the substrate, general tapography and position on slope. The proportion of oak in the reproduction class (less than 0.6 inch at d.b.h.) was found to decrease with improving site quality from a maximum of 50 percent on very poor sites to only 1. percent on very good sites. In general, reproduction of black and white oaks was most abundant on both ”very poor" and ”poor” sites while that of red oak was confined mainly to “medium" to "very good" sites. Studies of upland forests in contiguous regions indicate the occur- rence of oak upland types similar in composition and site characteristics to those of southern Michigan. Kittredge and Chittenden (1929) recog- nized a gradational sequence of oak types on the sandy scrub oak lands north of the tension zone across lower Michigan. These appear to be correlated primarily with edaphic influences. They range from the jack oak type, on the most xeric and sterile soils, through jack oak-white oak and white oak-black oak mixtures on intermediate sites, to the red oak type on the most mesic and fertile sandy soils. In a study of 98 woodlots in Missaukee County, likewise north of the tension zone, Elliott (1952) found stands dominated by red and white oaks to be restricted to xeric sands of the Roselawn series. In the glaciated region of northern Ohio, the general descriptions of Sears (1925), Sampson (1927, 1930) and Shanks (1942) indicate a gradational series of upland oak types determined primarily by available moisture. 0f the species daninating these types, post and bur oaks occur on the most xeric forest sites, with a sequence from black to white to red oaks indicative of progressively more mesic conditions. The two 8. charts included in Supson's second paper (1930: 360, 362) are most use- ful in indicating the ranges in site tolerance and abundance of the var— ious tree canponents. A similar chart of site range for the trees of southern Ontario has been provided by Hills (1952). Gordon (1936) recognizes four types of upland oak forest in Indiana. Although he does not attempt to arrange these in a series as regards site requiruents, fran his descriptions the white oak-black oak type would definitely appear to be the more xeric and the oak-maple type the more mesic. Oak and oak-hickory canprise the intermediate types. Potzger (1935, 1939) and Potzger and Friesner (1940b) present quantitative data which indicate that tapographic climate exerts sharp control on forest types in central Indiana. All three sets of quadrat data included with these papers seem inherently consistent except Potzger' a data of 1935 which indicate that M mg: m: (8 9. M) is restricted to a south slope and Q. zelutga to a north slope. Such results are at such marked variance with the other two sets of data, as well as with observational experience in general, as to sug- gest that the two names may have been interchanged in setting up the table. Potzger and Friesner (1940a) and Jones (1952) have compared the herb synusiae of oak upland stands in central Indiana with those of more mesic types in the same region. With one exception, their data indicate oak upland forest to be richer in species than other types. Comprehensive phytosociological reports on the forests of southern Wisconsin have been presented by Whitford (1951) and Curtis and McIntosh (1951). Whitford classified the 26 stands included in his study on the Q l 9. basis of shade tolerance of the tree components. He arbitrarily assigned each tree species to one of three tolerance classes and then determined the preportion of all trees in a stand which belonged to each class. This permitted the ranking of stands in accordance with the percentage of intolerant components. The importance of black oak, bur oak, shag- bark hickory and black cherry were found to decline sharply in stands canposed of less than 75 percent intolerant species. White oak remained high in importance in the intermediate stands commsed of 25 to 45 Per- cent intolerant species, while red oak reached its peak importance in such stands. Curtis and McIntosh (1951) presented data from 96 upland stands which were ranked according to the more inclusive concept of "climax adaptation ," encanpassing the entire assemblage of attributes which collectively detemine the range of environmental tolerance of a given species. Their method of stand analysis represents an admirable example of the inductive approach. First, relative measures of density, fre- quency and dominance (all expressed as percentages) were added to yield a emanation index termed W 1&3- Next, the importance value for each species of a stand was recorded to scale, and in color code, on a strip of celluloid. Similar strips were prepared for all. 96 stands after which the strips were arranged in such order as to yield the smoothest possible trends in importance value for four key species.1 ' Such a sequence of strips was found to indicate independent , although broadly overlapping, curves of importance value for the various stand canponents. It was possible, therefore, to rank each species as to 1mm.floalha.Q-mhmanddpsrissshm- 10. climax adaptation by assigning it a number ranging from 1 to 10 depend- ing upon the position of its importance value-maximum relative to the other species. These 31113,; W nmnbegg were used to weight the importance values. The resulting weighted importance values were then sumed to obtain the gontiguum ind-g; m for each stand. For any given stand this number has a possible range from 300 to 3,000,2 so a W W £922 of 2700 scale divisions is available for classifying upland forest vegetation. Beginning at the xeric (low) end of the scale, the importance value-maxima of the principal oak upland species of southern Wisconsin occurred in the following order: WW. 9. My mm, 9. alias. 9. ram. zsince the importance value index has a constant total value of 300 units per stand. 11. THE REGION AND ITS HISTORY W Except for a few extremely local outcroppings of bedrock, the whole of southern Michigan is mantled with glacial drift of Wisconsin age. The genetic types of this drift - morainal, fluvial, lacustrine -- have been differentiated and mapped in considerable detail by Leverett (1924) . Primary Configuration Features The brief interval of geologic time during which the unconsoli- dated deposition in southern Michigan has been subjected to gradational forces is nearly everywhere reflected in the constructional character of the land surface. Gross constructional variations in the drift mantle permit the recognition of maj or phys iographic and land divisions (Veatch 1953: Fig. A) . In a general way these divisions are correlated with the genetic types of drift which comprise the primary configuration features of the region. In the southern half of the Lower Peninsula, the prevailingly flat lacustrine plains adjoining Lakes Michigan, Huron and Erie are delimited toward the interior by old shorelines marking the highest stages of glacial lakes. Such shoreline traces are located at elevations ranging frat about 150 to 200 feet above present Great Lake levels. The interior highland reaches elevations of approximately 400 to 600 feet above present 12. Great Lake waters and includes three major land types: (1) rolling clay plains; (2) dry sandy plains; (3) sandy hills. The rolling clay plains, consisting of scarcely undulating to gently rolling terrain are closely correlated in distribution with clay till plains; because of the relatively low perviousness of the drift, surface runoff is of common occurrence and, in consequence, even very slight local relief (may strongly influence moisture relationships. The dry sandy plains are most typical of outwash plains, but also include some valley train and terrace deposits; because of the pervious sandy drift, surface runoff is of restricted occurrence and dry depressions, often of a pit nature, are a common feature. The sandy hills conform closely in distribution to the belts of recessional and interlobate moraines. Although an abundance of lakes .is a characteristic feature of this land type, the presence of innumerable dry hollows which lack surface drainage, implies a prevailingly pervious substratum. Secondary Configuration Features The secondary configuration features, which collectively detemine the local landscape aspect, are largely an expression of minor construc- tional variations in the drift mantle. Though valying greatly in abun- dance, size and shape in the different landscapes, depressions which lack surface drainage are perhaps the most characteristic secondary configures- tion features of the region as a whole. Such depressions are, of course, a corollary of the youthful drainage pattern; these range from more sags 01‘ Pits, dry or only intermittently containing water, to permanent swamps, marshes and lakes, often of considerable extent. 13. Though likewise having a multitude of variant forms, the elevated configuration features of secondary rank are in nearly all instances characterized by rounded contours and a complexity of slopes. In canbina- tion such land fonns result in a landscape of typically softened aspect. Slope gradient varies through wide limits. Occasionally, a prom- inence will be found with an abrupt incline approaching the angle of repose; more commonly, however, gradients are of much lower order and frequently are scarcely perceptible. Local relief tends to be of rela- tively low magnitude; generally, it does not exceed 50 or 75 feet, and even in the more rugged morainal belts is very seldom more than 150 or 200 feet. A map of generalized slope and relief is provided by Veatch (19538 Fig. 13). In the light of the above generalizations it would appear that influences correlated with aspect or exposure may be considered minor sources of vegetational diversity in southern Michigan. However, in canpaly with variations in the lithologic composition of the drift, secondary configuration features are of primary significance in deter- mining moisture relationships on a local basis, and, accordingly, may be expected to play an important role in determining the local patterns which collectively comprise the regional mosaic of natural vegetation. Although the multiplicity of detail evident in a given landscape defies canplete cartographic expression, Veatch (1953), by recognizing pattern continuity as regards minor variations in configuration and lithologic canposition of the drift, has succeeded in delineating minor land divisions which are most helpful in visualizing the more or less characteristic landscape aspects prevailing over extensive aggregate a1‘eas. 11.. 3 Clinics The regional climate of southern Michigan alternates between con- tinental and seminarine. The intervals of the latter type coincide with periods of strong onshore winds during which the equalizing effects of the windward Great Lakes waters are manifested far inland. Because of prevailing westerly winds, the waters of Lake Michigan are more influ- ential in this regard than those of Lakes Huron and Erie. Continental climate, with its characteristic pronounced fluctuation in temperature, prevails during periods of atmospheric calm, as well as during those rare intervals when the Lakes are entirely frozen over. A narrow belt extending along the Lake Michigan shore is character— ized by relatively continuous seminarine climate. The transition from this zone of more equable climate to that of the interior is not well defined and, over the interior, differences in topographic climate are even less marked, as may be inferred from the rather narrow range in elevation and local relief characterizing the area. Spatially, there- fore, the climate of southern Michigan may be regarded as essentially uniform. Such effective variations as occur appear to be of a regional nature, and thus not correlated with the rather abrupt transitions in natural vegetation that characterize the area. The mean annual precipitation ranges from less than 28 inches in the Thumb Area to more than 36 inches in the extreme south-central por- tion. Precipitation is fairly well distributed through the year, though the monthly totals in winter average about one inch less than in summer. An average of 16 to 20 inches falls during the warm season between April 1 and September 30. The number of days with snow cover of one inch or 15. more in depth ranges from about 70 in the south to around 110 in the north. The average annual tanperature ranges between 1.5 and 50 degrees, with a July average of about 72 degrees as opposed to a January average of around 21. degrees. The growing season ranges from more than 180 days in the extreme southwest to about 11.0 days in the Thumb region. According to the K‘o'ppen classification of climates as modified by Ackeman (191.1), the area lies well within the Df boundary, which delimits a cold forest climate with humid, snowy winters. The a/b boundary between the subdivisions marked by hot and cool summers passes through the south- ern tier of counties. A more genetic classification by Borchert (1950) indicates the area to lie largely in the transition zone between his climatic regions I and IV. Its climate is thus held to be intermediate between that of the northeastern forest region of deep winter snows and reliable smmer rains and that of the prairies with relatively dry winters and occasional low summer rainfall. The inclusion of the southern tier of counties in region IV is not well supported by the natural vegetation, although some restricted areas of prairie do occur in the southwestern counties (see subsequent section, General Vegetation). €92: Authoritative treatments of the soils and land of Michigan are pro- vided by Veatch (1941, 1953). His latest monograph largely furnishes the basis for the resume below. The mineral soils of the southern half of the Lower Peninsula are derived from gray and yellowish drifts showing a strong influence of shales and limestone from the local geologic formations. In the highland l6. piwsiographic division of the region there is an additional strong influence of local sandstones. The zonal soils of the region -— those associated with upland forest vegetation - are representative of the grey-brown podzolic group and probably comprise about half the total area. Profiles from southern Michigan referable to this great soil group show certain features in common. The organic accumulation on the sur- fatn is relatively thin, varying mostly within the limits of one to three inches; calcium and magnesium carbonates have been almost com- pletely leached from the solum ; sesquioxides have been removed from the sole, or upper gray horizon immsdiately below the mull humus layer. Depending primarily upon the proportions of the different soil separates, three principal variants of the idealized zonal profile may be recognized. These are: (1) profiles characterized by predominant influence of clay; (2) profiles developed from drift of intermediate texture; (3) profiles developed on prevailingly sand drift. The zonal soils of southern Michigan strongly reflect the extreme diversity in surface configuration and lithologic composition of the parent materials fran which they were developed. Accordingly, it is most difficult to generalize concerning the landscape aspects and prin- cipal soil types of such areas as are characterized by predominance of any one of these profile variants. Nevertheless, catenary sequences involving principal soil types and showing a gross correlation with configuration features m be recognized. Such gradational sequences are determined primarily by variations in moisture and texture (see Veatch 1953: Fig. 10). Thus, clay profiles prevail on the rolling clay 17. plains of ground moraine origin where the St. Clair and Napanee types are representative of the less pervious soils of the relatively flat land, and the more pervious Miami type characterizes much of the elevated terrain. As the topography becomes more rolling, and the clay influence less predaninant, the Hillsdale type becomes increasingly important and forms a transition to the intermediate profile variant. This soil also plays a transitional role as regards topography, occurring typically on the more rolling terrain of both till plains and recessional moraines. On the higher, complex ridges of recessional and interlobate mor- aines the Hillsdale soils are largely replaced by the Bellefontaine and Colma types. The Bellefontaine and Hillsdale types are roughly equiv- alent texturally, but the former tends to be somewhat more xeric because of configuration characteristics. Coloma soils are representative of the sand profile variant and thus constitute the most xeric type of the sequence. A more abbreviated catena is characteristically developed on drift of glacio-fluvial origin, such as outwash plains, valley trains and ter- races. In this gradational sequence the Oshtemo type, characterized by a very weak clay B-horizon, is transitional from the Fox soil, representa- tive of the profile variant of intermediate texture, to the xeric sand profile of the Plainfield type. It must be aphasized that the above soils, counprising the princi— pal upland types of southern Michigan, do not occur in extensive con- tinuous bodies but, on the contrary, are to be found in intimate associa- tion with one another and with a large number of other types as well. Moreover, such soil associations in turn comprise a pattern of marked 18. canplexity, as is strikingly shown by the detailed map of soil associa- tions canpiled by Veatch (1953) to accompany his monograph. Finally, even the soil within the boundaries of a single type must be envisioned as of a gradational nature, for as Veatch states (1953: 27) ”no unit small or large, possesses absolute uniformity or homogeneity throughout the space given to it on a map. It usually happens that the divisions shown on a soil map are more often an association of units rather than a single unit.” Ge al e n Postglacial History Gleason (1923) has presented a detailed, and in many respects speculative, account of the vegetational history of the general region. Subsequent reports have been based on, or at least tempered by, the evidence of pollen analysis. Sears (1942b) has charted the migration routes of five important tree genera. In general terms, Querous and Fagus advanced from the south; Carya and Tilia entered from the west; Tsuga migrated from the northeast. Pollen profiles depicting the postglacial clisere in southern Michigan have been presented by Potzger and Wilson (191.1), Sears (1942a), Pamelee (1947) and Potzger (1948) . In general, these profiles support the regional sequences recognized by Sears (1948) and Flint and Deevey (1951), which involve five main phases from deglaciation to the present: 1. Sprum-fir 2. Pine 3. MesOpiwtic deciduous 4. Xeroptytic deciduous 5. MesOptytic deciduous fa 19. All three deciduous phases of this clisere were evidently characterized by a predaninance of oaks, with beech assuming increasing importance in the last two stages. The method of radiocarbon dating devised by Libby, Anderson and Arnold (1949) has provided evidence that the Mankato maximum , which marks the approximate time of initiation of this clisere, occurred only 11,000 years ago. (Flint and Deevey 1951: 261-263) . Additional radiocarbon datings indicate further that, depending on latitude, the pine phase of North American profiles occurred from 6,000 to 9,000 years ago bi .: 272) . No consistent chronology for the deciduous phases of the regional sequences is yet available. The latest estimate regarding the dating of the thermal maximum (phase 4, above) in North America is from 3,000to 6,000 years ago (£913.: 258). There is increasing support for the view that once local climates determined by proximity to vast continental ice sheets ceased to exist, subsequent major climatic shifts were contemmraneous throughout the world (Raup 1937; Deevey 1939; Willett 1949, 1951). In his two papers Willett presents a great amount of evidence in support of the theory that geological, postglacial and secular trends in climate are all determined by the same astronomic control.3 In the northern hemisphere he regards such control as reflected by long period expansion and contraction of the pattern of zonal circtmipolar weather. According to this theory, phase five of the above sequence, indicative of increased moisture, could be correlated with the sub-Atlantic period of Maps initiated about 2,800 3Most probably ultraviolet radiation of the sort correlated with sunspot activity (see Willett 1949: 1.6). 20. years ago. On the basis of widespread and well documented evidence in the northern hemisphere, Willett (1949: 49) notes the occurrence of a . second thermal maximum during the period from 400 to 1,000 A.D. He states that "this period was similar in nearly all respects to the Cli- matic Optimum except that it did not develop quite as far, nor last nearly so long." Evidently this thermal maximum corresponds to the retreat phase of the Cochrane episode of post-Mankato time reported by Sears (1948), and the,lg§g thermal maximum postulated by Transeau (1935). Sears describes this period as ”marked by the northernmost advance of deciduous trees, notably oak, and by the prevalence of oakphickory and prairie in Ohio." It would seem likely that the pro-settlement prairies in southern Michi- gan represented relics dating from this Period. 'Willett (1949) describes the last 950 years as a period of climatic stress, reaching peak severity during the 13th and 14th centuries when the thriving Viking colonies in Greenland were exterminated. Subsequent to that time he states (pp. 49—50) that "only minor climatic changes have occurred although conditions have remained definitely on the cool and stony side canpared with the milder periods, in both Europe and North America. The 17th and.l8th centuries were rather mild, the 19th somewhat colder and stonmier... but the recent oscillations of climate have been relatively small.“ Judging from this general characterization, recent cliseral shifts of vegetation might be safely considered of very small moment. Pro-settlement‘Vegetational Pattern Prior to the period of intensive settlement beginning about 1820—- 1830, Veatch (1953: 196) estimates that forests covered probably no less than 95 percent” of the area of the state. The vegetation map compiled fran land office field notes by Marshner (1946) indicates dry prairies and marshes were concentrated mainly in southern Michigan. However, the aggregate area of such dominant vegetation would seem scarcely sufficient to be reflected in a proportion of forested area in southern Michigan significantly less than that for the state as a whole. Marshner's map shows the original forests to have consisted largely of hardwoods, with areas of swamp conifers scattered throughout, and with conifers5 codominant with hardwoods over extensive areas in the Thumb Region, in the interior north of the southern four tiers of counties and in Ottawa, Allegan and Van Buren counties bordering Lake Michigan. Veatch's (1928b) reconstruction of the original forests, based on soil- vegetation correlations, is in essential agreement with Marshner' 3 maps as regards the areas of hardwood-conifer dominance and, in addition, delimits tentative, generalized types of hardwood forest. In general, the more local variations in forest composition appear to be strongly correlated with moisture relations, which in turn are con- ditioned mainly by surface configuration and lithologic composition of the weathered glacial drift. Veatch (1953: 39) estimates that possibly as much as 1.0 percent of the area included in the gray-brown soil region (orbs remaining five percent is described by Veatch as comprised of "lakes, marshes, bogs, natural prairies, lake beaches, shifting dunes and garden sized clearings made by Indian tribes.” smably white pine and hemlock. 3 22. of southern Michigan is characterized by azonal or intrazonal soils. of these, wet types - bydromorphic, organic, recent alluvium -- comprise all but a.minor fraction of the aggregate area. As a group, such wet soils originally supported some expression of swamp or bottomland forest. 0f the organic soils, peats were characterized by swamp conifers6 with an admixture of hardwoods such as aspen, red.maple, elm and black ash. iMuck soils, representing an advanced stage of decomposition and alteration of the organic parent material, supported a hardwood swamp forest in which american and rock elms, red and silver'maples, black ash, swamp white oak and cottonwood were common components. Swamp forests of hardwoods also were characteristic of hydrumorphic soils. For the most part these were similar to the types developed on :muck. In forests developed on heavy textured hydromorphics such as the Brookston type, bitternut and shagbark hickories, white ash, sycamore and basswood were additional common components; likewise, the diversity of the swamp forests developed on wet sandy soils were increased by pin and bur oaks, black gum.and aspen. Not all the zonal soils supported typical upland forests. On the less undulating portions of the interior clay plains what might be re- garded as types transitional from swamp to upland forest occurred on the less pervious Napanee and St. Clair types. Except for the occurrence of sugar maple, beech and white and red oaks .- usually in slightly elevated sites - these forests were reminiscent of the swamp type develOped on Brookston soils. 6Typically tamarack but occasionally black spruce or northern white cedar. 23. On the more rolling portions of the interior clay plains, as typified by the Miami-Hillsdale-Conover Association (Veatch 1953: 61) the forest assumed a more definite upland character. Regarding this soil association, Veatch writes (p. 62): The original forest cover on the upland probably contained more sugar maple and beech than elsewhere in Michigan, but it is doubtful whether these species were universally present on all the upland, or in greater numbers than white, red, and black oaks, elms, white ash, hickories and basswood. with increasing local relief and soil perviousness, as typified by the catenary sequence of soil types from Miami through Hillsdale to Belle- fontaine, the original forests increasingly assumed the character of oak- hickory uplands with red, white and black oaks and pignut hickory can- prising the major dominants. 0n the soils deve10ped from drift of glacio—fluvial origin, oak- hickory forests of canposition similar to the above, or forests dom- inated largely by oaks alone, were of widespread occurrence. The latter type was canposed largely of black and white oaks, except in the environs of prairies where bur oak was the leading dominant. The latter areas occur as relatively small tracts aggregating some 85,000 acres in the southwestern part of the state, where they lie in close proximity to the central prairie region. The absence of any indications of local edaphic, topographic, or climatic compensation (Veatch 1953: 1.1.) suggests that these areas represented relics of a once more extensive prairie region and that they persisted up to the time of settlement through the influence of fire coupled with the capacity of grass vegetation to canpete with tree reproduction. Post-settlement History The pro-settlement pattern of natural vegetation in southern Mich- igan has been profoundly modified by the activities of Caucasian man. It'may'be assumed.that a very small preportion of the change dates back more than 150 years since as late as 1817 there were no roads leading to the interior (Tuttle 1873). By 1830, however, heavy immigration was in progress and during the brief interval from 1830 to 1838 the population of the territory increased from 32,000 to 175,000 (Lanman 1839). Hatch— kiss (1898) states that by 1837 at least 433 sawmills were in Operation in the southern part of the state. The product of these mills was pre- ' dominantly whitewood and basswood lumber which seemed to meet best the needs of the settlers. From.the accounts of Lanman (1839), Hotchkiss (1898) and watkins (1900) it is evident, however, that only a small part of the old growth thnber was converted into lumber. {Much oak and black walnut was used for rail fencing and great quantities of timber of all species was burned to provide ashes for glass and soap manufacture. To make way for agricul- ture, girdling or ”windrowing" preparatory to burning was practiced on an extensive scale. watkins observed that: One of the worst obstacles in clearing off the timber preparatory to opening up the land for agricultural pur- poses was the great whitewood trees ... These trees when fallen and green positively refused to burn ... Other trees were felled across then‘and a fire started and kept burning sometimes for weeks until the great logs finally yielded. Clearing for agricultural purposes in southern.Michigan has pro— gressed to the point where in 1950 woodlands were estimated to comprise only about 10 percent of the farm land (Cunningham 1950). In addition 25. approximately two-thirds of these remnant woodlands were reported to be grazed (Anonymous 1950). The pattern of forests and crepland in southern Michigan has been and continues to be strongly influenced by the pattern of natural drain- age. In the early period of settlement the well—drained oak uplands were particularly sought out because of the ease with which reads could be constructed and the land prepared for agriculture (Veatch 1928b, 1953). In consequence, there is little doubt that the preportion of cleared land is greater in the oak upland areas than in any other forest type in the state. In addition to being greatly reduced in area, the oak upland type in southern Michigan has been greatly modified by past utilization. Many tracts are perhaps better classed as wooded pastures than woodlots; stands containing any large preportion of old growth are uncommon. Gysel and Arend (1953: 14) have called attention to the selective cutting of white oak in the last half of the nineteenth century. They further express the belief that hoary cutting on moist sites where oak was originally associ- ated with other hardwoods has increased the present preportion of oak. Further'modification of the oak type in southern.Michigan.may be expected to result from the ravages of oak wilt. In.Michigan this dis- ease was first discovered in 1951 in seven southern counties (Strong 1951). In some locations the initial infection was judged to have occurred about five years earlier. The disease has since been found in additional counties (Foster 1953). A sanitation program designed to control oak wilt by voluntary destruction of infected trees was initiated in 1952 (Smith 26. 1953). It is at present too early to evaluate the effectiveness of the measure, though Foster (1953) does express cautious Optimism that the disease can be suppressed in lightly infected areas such as southern Michigan. METHODS Recopngigggpce General reconnaissance, to attain working familiarity with the vas- cular components of the oak upland community and to locate representative stands suitable for sociological analysis, occupied most of the summers of 1948 and 1949. Ebrploratory field trips were made to districts con- sidered promising on the basis of vegetation maps prepared by Kenoyer (1930, 1931., 191.0, 1943) and Veatch (1928b) . In addition information regarding the location of specific stands was sought from local sources including farm foresters, lumbermen, district conservation personnel and local botanists. In all, well over 300 stands, distributed through thirty- five counties, were critically appraised as to suitability for sociological amslys is . Stgpg §e1§ction As originally conceived, the criteria of stand acceptability for this a"Sudy closely approximated those of the relic method expounded by Weaver aJld Clments (1938: 1.8) . It was hoped that ungrazed stands, essentially Primary in character, or at least of advanced second growth, could be located in replicate on each of the major soil types and natural land divisions (Veatch 1931) which were found to support the oak upland cm- nrllmity in southern Michigan. However, as extensive reconnaissance re- vealed the full magnitude of disturbance and fragmentation of the oak up- land cmmmnity, it became evident that a drastic lowering of the initial 0‘" ,r. O u 28. criteria of acceptability would be mandatory. Thus, at the end of the second season of reconnaissance, an oak upland stand, wherever its loca- tion, was regarded as suitable if (a) it showed no evidence of m posturing, (b) was reasoggbly well stocked with class 5 dominants, and (c) occupied enough area, either in a single block or in fragments, to permit the establishment of ten quadrats within the zones of border effect. On the basis of these revised standards a total of 36 stands were selected for sociological analysis. The approximate location of each stand is shown in Fig. 1, and a detailed description of location -- by township, range and section -- is included in the captions of Appendix Tables I to WI. Of the nearly 300 stands rejected, the most common defect was evidence of recent pasturing. In view of the urgent need for a large and representative set of stands, no ungrazed stand was rejected without thorough appraisal of its minimum possibilities. Obviously, no clear cut boundary separated stands rejected as too small from those finally selected for detailed study. In several borderline cases tracts consisting of two or more apparently homogeneous fragments of older growth, interspersed in a matrix of younger timber, were included in the Study with the aggregate area of the older growth considered as a single ltand. Of the stands selected, only three contained any large preportion of old growth individuals, and only one -- since logged -- was regarded as a true primary remnant. While it is clearly evident that the shift in emphasis fran primary to secondary stands in this study was a practical expedient, this shift 18 not difficult to defend. Today, an overwhelming and constantly “('3 fl,‘ .. .t .Ir 1 qt 1: Pal I. .3... 0 I -‘ .fidwwfiam mwmuew m3 QOHNhooH hfwhOim Gomfl£odé Chox+Dot 90 Q.L H Okh on! . ZuEmum _ I..|u|L.-.|l1-l.I.ll _” Olt . . \ _ .... . . . 3:2; _ zezuemasi c 285:.— zaorio radical 55.3 2.; Cl - l I 'IIII' . u m 0 O. 004 morass. horizon mamas ...: Eases—ranges e96. —. -l-l.!-l-l.| . o . memopm steam mo NoHquo . . _ . _ . _ -....- _ . _ . . _ _ O . . . O mwoopm 039 . . e rmmmg.‘ Bengt—... - wheat: . Efifil - 26mm... TicouL a 9:323 _ .wl . _ .._| .. II. ....II n III a ..II . M _. . O. _ O. _ 683m oflmfim O .3... _ ..-...._.....s....._a - I... - I... 525 .Iummlummmu. II - ll. . m e _ ..l u re a. m . _ ...Lekamsea ...l. . . w . sez.e-<\tr1>oH no .0: Smmmfiou Inflow; noses. across as Eorfioz Essence messages: asses. on: wood poem Nnmoamoooa has: no ommofi to.» on no squeegee oonoofimsoo on has masoeoameoo mofioomm poo opoefidoonooa .NOQNJ Hfiom song: you Noameoo o no common sebum Mo on: wood poem was anaonmomov ma mneapowhob .HHHN mnde 126. TABLE XIV. Net positive differences in weighted importance values between indicated stand pairsl contrasted to illustrate influence of s10pe Contrasting stand pairs Species 22 vs. 28 16 vs. 32 24 vs. 31 Acer rubrum - 203 -— 284 - 137 Acer saccharum -. .. .. .. .. 56 Amelanchier sp. - -- - - - l8 Carya ovalis 85 -- 1 - 108 .. Cornus florida - - -- 22 -— 22 Fagus grandifolia - -. .. -- 47 -- Fraxinus americana - 31 -— - l3 .- Ostrya virginiana - - - - - 241 Prunus serotina 37 - 51 - -- -- Quercus alba 74 -- 130 -- 221 - Quercus rubra - 278 - 553 —- 167 Querous velutina 43 - 159 - 2 -- Sassafras albidum .- .. 22 .. -- .. Tilia americana —- 48 -- -- .. -_ Ulmus americana 32 - -- - .. .. Total net positive difference 271 560 363 859 391 641 Differential in continuum index number 289 496 250 1See Table XIII for stand characteristics. 127. differences in weighted importance values which collectively produce these differentials in continuum index number are shown in the upper rows of this same table. These latter data indicate without exception that Acer Egbrum and Qgercus Egbrg are mostwimnortant 1n thefimore mesigwmember of each stand pair, while Cagya ov§lis,IQ. glba andIQ. velutina are more important in the less mesic member. -. ha .‘ ' 5M .— ...- _— u—II~ Fa —- n— W.— war" A note of interest, and an indication of the reliability of the con- tinuum index values compiled for stands 24 and 31, is provided by data from 10 quadrats oriented on a diagonal line from crest to base of the north-facing lepe on which these two stands are located (Appendix Tables XXXVII and LXXIV). The continuum index number of 1653 computed from the data obtained by this sampling pattern might thus be regarded as represen- tative of the entire tract. That such a number should be only 12 points from the mean of the continuum index values for stands 24 and 31 would therefore seem to constitute a favorable reliability check. Influence of Slope on Other Community Components Tree reproduction. The three classes of tree reproduction tend to show the same general reSponse to lepe influence as do the dominant elements. Correlation involving any given species, however, is less predictable, as evidenced particularly by'§M§££g§,zpbzg, which in all three reproduction classes was recorded in greater amount in stand 16 than in stand 32 (see Appendix Tables). Shrubs. With the exception of three species, the shrubs recorded in two or more stands of this complex show no definite correlation with 128. lepe. The exceptions involve Ceanothus americanus which was not recorded on any north lepe,'Eiti§ gestivalis var. argentifolia which was not recorded in stands 28, 31 and 32, and.Viburnum acerifolium which appears clearly most abundant on north slopes. ‘flggbg. Totals of species per stand (see Table X) and average Species densities (see Table XI) agree in indicating restricted herb representa- tion on north slopes of this complex. On a Species basis, such restric- tion is explained by the relatively few herbs more abundantly represented on north s10pes, as contrasted with the group evidently restricted in occurrence or absent on such sites. 0f the 42 species listed in Table IX, only Uvularia andiflo , Qgggx albuzsina and.Adiantum‘pgdgtgm appear clearly most abundant on north slopes. A.much larger group, notably fielignthus div ic t , Pteridium gguilinum var. latiugculum, Eigig c olinian , Degmodium otundifolium, Legmdgzg waging; and Wm vigginiana is evidently much restricted on such slopes. The evident sparseness of the herb synusia on north slepes of this stand.complex runs directly counter to the tendency for herb representa- tion to increase in both species and species density with increasing con- tinuum index number (see Fig. 37). The implications of this departure from the general synusial trend will be considered later in connection with other deviations from that trend. Influence of Soil Primarily because of differential patterns of past utilization, the influence of soil in contributing to the diversity of the oak upland com- munity is peculiarly difficult to assess with any precision. Attention 129. has previously been called to the need for basing each attempt at such evaluation upon data obtained from a single tract which appears to have had unifom treatment in the past. Soil Variation within Single Tracts Variations in soil profile of differing magnitude may usually be readily demonstrated in any oak upland tract in southern Michigan. Where such variation does not exceed the relatively narrow limits established for a given soil type, the associated vegetation might likewise be ex- pected to vary relatively little. Most surprising, therefore, is the 649 point difference in continuum index nmnber between stands 1.2 and 22, located in a 40 acre tract mapped entirely as Miami loam. SlOpe in this tract is to the west and averages less than 2 percent. There is no evidence that the pattern of past utilization of the two stands has been different. There is, however, evidence of greater moisture retentivity in stand 27 in the form of frequent wet depressions occupied by fiercus bicolor and 9m 21353 (see Fig. 2). It would seem, therefore, that soil influence must be considered the prime variable correlated with the differential in continuum index number between these two stands. In view of such a large differential evidently attributable to variation within a single soil type, it would seem likely that many replicate continuum index numbers would be necessary in order to provide a reliable estimate of the influence of soil variation greater than the limits of a single soil type. The likelihood that the remnants of the oak upland comnunity in gouthern Michigan would prove too fragmental t permit such estimation seems very great. Only“ a few oak upland tracts ‘4~«'IM-lr - I. ‘.'_.'n./'“1.‘ ‘ 130. developed on more than one soil type were encountered in the course of this study. None were considered suitable for measuring differences between soil types in terms of vegetational response. Usually only one of the represented types occupied sufficient area to permit sampling of the associated vegetation by the usual pattern of dispersed quadrats; often variation in soil type was found to be correlated with some ob- viously effective variation in tepography; occasionally the included soil types were found to be so intricately distributed as to form a canplex supporting vegetation representative of no soil type in particu— lar. The latter condition is well exemplified in the 40 acre tract in which stand 36 is located. Over considerable portions of this tract small, alternating bodies of Brookston loam and Napanee silt loam form a mosaic with which is correlated a stand canplex canposed of elements of oak upland, beech-maple, and bottomland forest. Correlation between Soil Type and Continuum Index Number The contrast in continuum index value between stands 10 and 27 suggests that soil variation may profoundly influence the continuum index sequence. Fig. 35 was prepared in an effort to determine whether or not soil influence is of such magnitude as to permit recognition of a pattern of correlation between soil type and continuum index number despite the effective variation of the other environmental factors. Inspection of this figure reveals two facts of considerable significance: (1) with the exception of stands 24, 28, 31 and 32, deve10ped on Bellefontaine sandy loam, no continuum index values compiled for stands associated 131 o .mpswpm popefioomms mo soapfimoa amps“ esnnfipnoo was camp Hflom escapes scaveaom .mm .on woes“ ansnfipsou comm 0@bfl. coma oouH oooH Gama ooqa ooma coma ooaa OOOH com com o m H p, _ b m0 HN b N _ r J. Hum .\ mam mm om o m _ . _ _ Ham mm Hm mm «m mm ma 0H ma «a MH NH w E _ _ p r _ _ p _ _ . Hum cm mm mm mm 0H _ . L P L HZ ha HA fl . L um ma _ HmH pawn mecca nohom II mam om cm 53 some.“ ogmdfi .... Hum p «L Hmz acoa spasm onfispnomoaamm II Hum - mm 38H Hafiz ..I H: _ Hm: efim 3333a .... am on Euoa hensm «Haopme It HmH _ Ham awoa pdfim mondawz II Hmz mm EUOH page “Sufi: at an: _ Ho case uaoHoo nu no Sued paflm nfiufio .pm at Hum auoH henna xom II Hum sued nobosoo II Ho 132. with soil types designated as sands, loamy;§ands °tx§E§§XmE9§§3 exceeded a value of 1500 points; (2) with the single exception of stand 10, develOped on Miami loam, no continuum index values compiled for stands associated with soil types designated as loams or silt loams failed to exceed 1500 points. It will be recalled that the four exceptional stands develOped on Bellefontaine sandy loam are all located on s10pes of northern eXposure (Table XIII). Moreover, in comparison to the index positions of the four other stands developed on Miami loam, the index position of stand 10 appears clearly anomalous (Fig. 35). The general observation would therefore seem permissible that, in terms of vegetational response, index -. fl__ .ficm ..- two broad textural groupings of soils on ...-.....th position 1500 serves to delimit which oak upland stands are developed in southern Michigan. The sequence of soil types in Fig. 35 is not to be interpreted as representing an attempt to arrange the types in accordance with their moisture relationships as suggested by the data Obtained in this study. Such data are far too scanty, and the influence of other environmental factors far too efective, to justify such an attempt. ‘Nevertheless, the sequence, determined.merely by the lowest continuum index number associated with each type, seems in reasonable accord with existing views (veatch 1953) as to moisture relationships of the types represented. Thus, there is considerable support for the view that the edaphic factor may be con- sidered the prime variable in the oak upland environment of southern Michigan. W The full degree to which past patterns of utilization.modify’vegeta- tional expression of other environmental influences cannot, of course, be 133. determined by comparison of second growth stands. However, a well stocked, uneven-aged stand including a considerable proportion of old growth and showing no evidence of heavy pasturing might, with some justification, be considered a reasonable approximation of the primary condition. In any event such a stand should be expected to diverge from that condition in far less degree than one totally lacking old growth elements and show; ing a tendency toward even-aged second growth. 0f the stands compared and contrasted in Table XIII, numbers 22 and 28 agree with the former characterization, while numbers 8, 16 and 32 fit the latter. A.suggestion of these differences is shown in Fig. 36. As previously indicated (Table XIII), the differential past treat- ment appears to be the only effective environmental variable correlated 'with compositional differences between stands 22 and 16, located on south slapes. The same observation applies likewise to stands 28 and 32, lo- cated on north lepes. Accordingly, Table XV was prepared to highlight contrasts in weighted importance value involving the members of these two stand pairs. The 187 point difference in index position between stands 22 and 16 would seem definitely indicative for severe disturbance, under the con— ditions extant in these two stands, to be reflected in a more xeric assemblage of dominants. The 20 point difference in the continuum index numbers compiled for stands 28 and 32 appears of too low magnitude to be definitely indicative of an Opposed trend under the environmental con- ditions common to these stands. It does, however, demonstrate that disturbance does not necessarily favor species of more xeric affinities. The differences in weighted importance values, comprising the body MW. . I‘VRWubui. ,. ‘.‘E‘& M .57? . u Fl“ ; dame»??? . I’MVI'H' . ‘l: u I . t . I. 134. A ...!!l 1.... - 01'. ‘lfiv'pg‘alib Sh“. . 1 1 A. . .....N .n x. . Ringo-lg”. Idllllkvahl‘lllr .II M c _ a .. .. 3...!1 -.l! SKI”..- , x . 1f.‘\\_“\‘tll.ao . n ,1 . \ . V .A.. City-2‘33“. Ii. . _ . , .31.. \ 1“... y . v 4 , . . \t. a c . n. A”! V I... _ IJIQLN‘Ev .. A a)“; ‘31: if. pice origin, View along the line fence between stands 16 and 22 showing older FIG. 36. timber at the right and younger growth, including some of c0p at the left. 135. TABLE XV. Net positive differences in weighted importance values between indicated stand pairsl contrasted to illustrate influence of past utilization _‘_ Contrasting stand pairs Species 22 vs. 16 28 vs. 32 Acer rubrum - 17 -. 98 Carya ovalis 50 - - 34 Cornus florida - - - 22 Fraxinus americana - - 31 .. Prunus serotina 4 - 18 .. Quercus alba 206 ...” 262 .. Quercus rubra 60 - .. 215 Quercus velutina - 126 - 10 Sassafras albidum - 22 .. .. Tilia americana - —- 48 .. Ulmus americana 32 —- .. .. T°§§f22nfizsmv° 352 165 359 379 Differential in continuum 187 20 index number lSee Table XIII for stand characteristics. 136. of Table IV, reveal the detailed pattern of changes correlated with both contrasts of differential stand disturbance. These changes are seen to be of a much higher order of magnitude than might be expected from the size of the differentials in continum index number, especially the 20 point difference between stands 28 and 32. The marked decrease in importance of Quercus alba in the wholly second growth stands is the most conSpicuous feature common to both contrasted pairs. Acer rubm and Q. zeluting are indicated to be more important in the wholly second growth stands. How- ever, the increase in weighted importance value of A. m in stand 16 is seen to be scarcely significant, as is true of that for Q. xelutina in stand 32. The great relative increase in importance value of 9. 33113;; in stand 32 together with that of g. m in the same stand, and of Q. Ieluting in stand 16, is strongly suggestive of the possibility that slaps influence may be augmented by stand disturbance. This suggestion is further sup- ported by the data in Table XIV which show the total net positive differ- ences in weighted importance values between stands 16 and 32 to reach con- siderably higher values than those between stands 22 and 28. Thus, even though evidence supplied by contimrmn index numbers does not appear deci— sive, significant differences in weighted importance values would seem to permit the tentative observation that, in reference to slope influence, second growth stands tend to show greater contrasts in composition than do primary stands. Some conception of the degree to which stand disturbance may modify contimnnn index number is provided by stand 8 (Table XIII). Because of the absence of effective sIOpe influence stand 8 is not comparable with 137. any of the other stands of this complex. However, it would seem unlikely that the primary assemblage on this site should have been.more xeric than stand 22, located on a south sIOpe. The possibility must be acknowledged that the 305 point difference in continuum index number between stands 22 and.8 may in part be attributable to effective soil variation within the limits of a single type.12 However, the likelihood of such influence in any important degree would appear small in view of the high.importance value of Suszgus.zshra in stand 8. In a classification based upon the two leading dominants as deter- mined by the importance value index, stand 8, in company with stand 16, would necessarily be regarded as a black oak-red oak type; likewise, stands 31 and 32 would be designated as pignut hickorybred oak (see Fig. 33). Such combinations of leading dominants have previously been inter- preted as anomalous and probably indicative of stand disturbance. This interpretation would appear valid in the case of these 4 stands. Numbers 22 and 28, representing the least disturbed.members of a complex including these 4 stands, are clearly to be classified as white oak-red oak types on the basis of the two leading dominants (Appendix.Tables XXII and.XXVIII). The low importance of white oak in stands 16, 31 and 32, and its near absence in stand 8, would thus seem strongly suggestive of differential disturbance which evidently reached peak intensity in stand 8. By permitting establishment of a sequence of stands based upon leimilar to that evidenced by stands 10 and 27, considered previously (see Fig. 35). 138. variations in the dominant synusia, the continuum concept provides a convenient means for determining the degree to which such.variation is reflected in the lesser synusiae. Should correlation of this nature be found to exist, it would add, in prOportion to the degree displayed, to the significance of the stand sequence as an expression of a parallel sequence of integrated environmental influences. Accordingly, the measures of total herb and shrub representation obtained in this study have been graphed below in a sequence based on continuum index numbers in an effort to determine the degree to which such correlation exists. e H b ia Based on herb data from 10 x 10 m. quadrats, total species records per stand and average Species densities, are graphed in Fig. 37. Though both curves are revealed to be extremely erratic, with increasing con- tinuum index number, rising trends in herb representation may be recog- nized whether measured by species occurrence or by average species denp sity. Yet, unless some valid explanation for the enormous fluctuations . between certain adjacent curve points can be offered, such fluctuations must inevitably diminish the significance of the overall rising trends. Accordingly, qualitative data have been surveyed in a search for evidence of anomalous environmental influences which.might be correlated with such extremes in herb representation. With regard to positive deviations from the trend of total Species occurrence, the values for stands 10 and 25 are revealed by Fig. 37, A, to be particularly extreme. Both of these stands are located in a single 40-acre tract characterized by many local areas of imperfect surface 139. 7Q Total species complement I - \ I \ Average species density 60 59 49 30 20 81012141618203; 5 16 15 26 25 3b 35 Order of Stands FIG. 37. Relation between two measures of diversity of the herb synusia and continuum index sequence. MO. drainage. Associated with these areas are a number of herbaceous species of high water requirement13 which serve to increase the variety and size of the species complements for these two stands. Somewhat less extreme is the species total recorded in stand 22, located on a sIOpe of southern aspect. Evidently correlated with the consequent greater amount of solar radiation reaching the forest floor in this stand is a species increment made up of intolerant herbs such as Degmodium ggtgndifolium, Lespedeza intermegg, Mogardg fistggga, Gezgrdig flayg, Q. pediculgrig and Q. gigginica. I Negative deviations from the trend in size of species complement reach extreme values in stands 9 and 34. In both these stands abnormally dense understory layers result in particularly heavy interception of solar radiation. The second layer in stand 9 is composed primarily of ngggg florida (see Appendix Table IX), and.where heaviest is correlated with a virtual absence of herbaceous vegetation (see Figs. 37 and 47). In stand 31., a dense understory of suppressed _A_g§;: m, A. W andHEgggg grandifolig grades into the closed canopy of old growth Qgercus glbg and‘Q.|ngg§. The result is an intensity of shade comparable to that cast by a full beechnmaple canopy. In addition to being thus heavily shaded, the forest floor of this, the only truly primary stand included in the study, is characterized by an extremely thick accumulation of litter and duff, which may further act to restrict.mechanically the representation of herbaceous species. Probably correlated also with decreased solar radiation is the restricted herb complement of stands 24, 28, 31 and 32, located on slopes 13See especially the representation of Carex spp. in Appendix Table LXXVII. 141. of north aSpect (see Table XIII). Although the species totals recorded in these stands are shown in Fig 37, A, to fall well within the normal range of deviation from the general trend, the totals are all low for the, scale sector in which they occur and thus tend to depress the slope of the trend. The general similarity in outline between graphs A and B of Fig. 37 suggests that the average species density of herbs in a given stand, as measured by 10 x 10 m. quadrats, is primarily a function of the number of species of herbs for that stand, Moreover, the approximate equivalence in slope of the two trends suggests further that the relationship is inde- pendent of sequence position of the stand. Confirmation of both indications is provided by computation of ratios between total species complement and average species density. Such ratios have a.mean of 3.0 g,0.5 and a range from 2.4 to 4.3, with 26 values clustered within the limits of 2.6 and 3.2. The only deviations in ratio to exceed one standard deviation are all posi- tive, and comprise those for stands 2, 6, l5, 17, 27, 34 and 36. Inspection of the values charted for these stands in Fig. 37, B, reveals that the low herb frequencies reSponsible for such high ratios result in improved correlation with the general trend of average species density in the case of stands 2 and 27, and contributes to serious devia- tion from that trend only in stands 34 and 36. The high ratio (4.0) of total species complement to average species density for stand 34.further attests to the sparseness of herbaceous vegetation in this primary stand. Observational data supply no explanation, however, for the relatively very low herb frequency implied by the peak ratio of 4.3 for stand 36. Nevertheless, when previously qualified deviations from the related 142. trend of total species complement are taken into account, the trend of average species density would seem to supply even stronger evidence of general correlation with stand sequence than does the former trend. ‘More— over, the significance of the trend in average species density is further enhanced by provision for the expression of frequency variations, which results in an ecologically more sound.measure of herb representation than a trend based on species complement alone. The S b S u i Based.on shrub records from 10 x 10 m. quadrats, total Species per stand, average species density, and the ratio between these two values are charted in continuum index sequence in Fig. 38. Although species comple- ment and average species density of shrubs are indicated by graphs A.and B of this figure to be correlated with continuum index number, the trend of such correlation is seen to be counter to that for the corresponding measures of herb representation. Further comparisons reveal additional significant differences between the two sets of curves. Even though graphed on a larger scale, the values for species complement and average Species density of shrubs Show con- siderably less fluctuation than do the corresponding values for herb representation. Doubtless contributing in large measure to such,moderap tion is the much smaller community complement of shrub species. That other influences are involved, however, is strongly suggested by the lack of coincidence, except in stand 34, of peak fluctuations in herb and shrub representation. Mereover, as indicated by the ratios charted in Fig. 38, C, the relationship between Species complement and average species density is considerably less precise for shrubs than for herbs. 16 143. Total species complement “WWW r l 1 Average species density Total species - species density ratio 5 10 1% 26 25* 3b 35 Order of Stands FIG. 38. Relation between three measures of diversity of the shrub synusia and centinuum index sequence. 1144. Furthermore, where there was obviously no correlation between stand se- quence and such ratios for herb representation, in Fig. 38, C, there is at least a suggestion of a rising trend with increasing continuum index number. As suggested by these ratio differences, Fig. 38, A and B, Show fewer points of similarity than do the corresponding graphs of herb rep- resentation. A third measure of shrub representation is provided by density totals obtained from 2 x 2 m. quadrats (Fig. 39). Except for Sporadic extreme deviations, the trend of this curve is seen to corroborate those of species complement and average species density in indicating negative correlation between Shrub representation and increasing continuum iniex number. The extreme positive deviations from the trend of total density are attributable in all instances to exceptional clonal deveIOpment, usually of a single species. Thus, in stands 2, 9, 15, 21 and 36 Parthenocissus guinmfolia rakes up the greater portion of the density total; similarly predominant is Viburnum acergolium in stand 20, and Gaultheria procumbens in stand 30. In stand 27 gm; radicans and Mg racemosa together comprise most of the shrub representation. Far more difficult to interpret are the fluctuations in species complement and average species density of shrubs (Fig. 38, A and B). Except in stand 3“ where both shrub and herb representation is markedly restricted, evidently by influences correlated with the primary condition of this stand, the shrub synusia apparently responds to a given integrated set of environmental influences in quite different fashion than the herb synusia. his is evidenced not only in contrary trends of correlation with continuum index sequence, but more particularly in the failure of Number of individuals counted in 4 m2 quadrats 140 1&0 220 2§O 390 340 3&0 420 mo 20 1&5. ‘g F 16 1E 26 2? ‘7 ,- 3'0 F Order of Stands FIG. 39. Relation between total density records for the shrub synusia and continuum index sequence. 11.6. conspicuous deviations in representation of one synusia to be clearly reflected in that of the other. The quantitative and qualitative data obtained in this study provide little more than suggestions as to what influences may be correlated with the more marked fluctuations in shrub species complement and average species density. That the low average species densities for stands 2, 9, and 15 may perhaps reflect the influence of unusually heavy competition, is sug- gested by the high total density values reached in these stands (see Fig. 39). The failure of the remaining extreme density values to coincide with abnormally low average species densities of course detracts from the credi- bility of this suggestion; such failure, however, might be found attribut- able to the more favorable moisture relations:L4 prevailing in the other stands where exceptionally high densities occur. A possible correlation of unusual edaphic influence with the high Species total in stand 17 is suggested by the uncommonly large number of calcifugous speciesl5 recorded in this stand. Although the collection of quantitative data in this study was restricted to stands which bore no evidence of recent pasturing, allow- ance in almost all cases should be made for residual influence of pastur- ing in the more distant past. That grazing may not only profoundly alter the total mass of the shrub synusia but also effect marked changes in its Species complement, was repeatedly demonstrated by prOperty line contrasts encountered during the reconnaissance phase of this study. One of the most frequently 14As implied by higher continuum index numbers. 15E aea e ns, Gaultheria procumbens, Gaylussacia baccata, Vaccinium tifolium, l. lamargkii, E. vacillans. 147. observed indications of differential intensity of grazing was a marked increase in the importance of ”armed" species, such as ignipgzga communig or W e icanum, as well as of unpalatable species, such as Ggylugsgcia paccatg, Gaultheria procumbens or‘gzuppg vigginiana, at the expense of most other synusial elements. An extreme example of this phenomenon is illustrated in Fig. 40. Because the influence of grazing might be expected to persist longer in shrub than in herb representation (see Fig. 41), such influence might at least in part account for the erratic ratios between shrub species complement and average species density (Fig. 38, C). I. L. . . III in a as r. .1..L a, View of an oak woodlot pasture showing effects of prolonged FIG. 40. overgrazing. FIG. 1+1. Depauperate clone of Junipems cm-munis persisting under closed canopy oaks. 150. DISCUSSION Whamlme ahasiWo la 1c 1 n The last three decades have witnessed the conflict of two widely divergent philOSOphies as to the organizational nature of concrete plant communities. The first holds these groupings to be highly integrated, functional units composed largely of inter-dependent individuals. Dis- tinctive aggregations of such individuals are regarded to be segregated with such regularity and fidelity as to justify the recognition of ab- stract communities. A voluminous literature in support of, and based on this view, has accumulated. Egler (1951: 682) expresses surprise that so basic a concept as the abstract community should have been the subject of so much emphasis. Perhaps this emphasis reflects not unp familiarity with classical precepts of inferential reasoning (as Egler suggests) but rather attempts to establish the validity of such reason- ing as applied to units of vegetation. Suggestive of such an inter— pretation are the many and varied analogies identifying the concrete community with the individual plant, and the abstract community with the species (Clements 1916, 1936; warming 1925; Nichols 1929; Phillips 1934, 1935; Tansley 1935; McDougall 1949) . The second view, embodied in the individualistic concept of Gleason (1926), denies that stand integration exists to a degree which can be determinative of stand composition. Rather, stand.cnmponents are regarded as determined primarily by environmental selection from such available 151. floristic elements as have compatible tolerance ranges. The concept of the abstract community is held to be untenable because, outside of very restricted areas, a given combination of environmental influences and floristic elements is seldom even approximately duplicated. Even the homogeniety of concrete communities has been considered subject to statis- tical verification (Curtis and McIntosh 1951: 481). Preponents of the individualistic concept have long been handicapped by lack of a classification scheme to support and illustrate their conception of vegetational organization. They could effectively chal- lenge the characterization of the concrete community as a complex, or quasi- organism, or as an element of an ecological "species" (Gleason 1926, 1929, 1939; Braun-Blanquet 1932; Mason 1947; Cain 191.7; Whittaker 1951, 1953); they could point to biased selection of stands to support the existence of such "species" (Cain 1947; Ashby 1948). They could not, however, effec- tively counter arguments such as that of Conard (1939; 110) that "there has been nothing in ecological inquiry which.has been so fertile and productive of results as the idea of the association. It is therefore so useful that whether logical or not, I am for it." Now, in the vegetational continuum index, an efficient means for orienting, and depicting the individ- ualistic nature of concrete communities of a given physiognomic type, is at last available. Orientation of compositional data according to continuum index sequence reveals that the bulk of upland forest stands in southern Wis- consin and of oak upland stands in southern Michigan are distinguished not so.mueh by different 339g; of dominants as by different proggrtigps of dom- inants. The explanation of this phenomenon lies in the fact that the 152. deminants of high importance potential have wide tolerance ranges which, while not coinciding, nevertheless broadly overlap. Thus, of the 15 species recognized by Quick (1924) as typical of the beech-maple com- munity in southern Michigan, not less than eight species are frequently found in stands dominated by upland species of oak. Of these, four species attain relatively high importance values in oak upland stands; two are leading dominants of oak uplands. It is obvious, therefore, that recognition of beech-maple or oak upland communities in southern Michigan is Justifiable only ch pragmatic grounds. The continuum concept accords significance to dominant stand com- ponents in preportion to their representation. However, because of the wide overlap in tolerance ranges of ecologically important tree species, the number of different combinations of weighted importance values which might approximate a given continuum index number appears almost limitless. Accordingly, such a number can provide only a gross indication of the importance of a given species in a given stand. For this reason, con- tinuum index numbers, or even segments of the continuum index scale, must not be regarded as comparable to cover types such as those recog- nized by the Society of American Foresters (Anonymous 1940) where diag- nostic significance is attached to the leading dominants only. Rather, a given continuum index number is an expression of the total dominant vegetation of a given site. 9 ve e tional C ntinuum dex Inde of Site and g Meaguge of Tolerancg Eggs Indirectly the vegetational continuum index:may be regarded as an index of site since dominant vegetation is theoretically the best measure 153. of environment. The value of the continuum index as a measure of site is enhanced.by the fact that it is based only on positive evidence. As Billings (1952: 263) has pointed out, the gbgencg of species is not a.valid basis for site evaluation. This follows because environmental control is only one of many possible reasons for such absence. If, as is contended here, the vegetational continuum index established for the oak upland community in southern.Michigan represents a valid index of environment, as well as of vegetation, certain implications are at once evident. The data charted in continuum index sequence become graphical representations of tolerance ranges of the sort envisioned by Good (1931) in his Theory of Tolerance. Such charts likewise support Billings‘(1952: 262) statement that "each species in a vegetation is distributed according to its own environmental tolerances." Charts, included herein, representing tree species show further that the principal dominants are seldom absent from sites indicated by index values to be well within their respective tolerance ranges. It is more difficult to generalize concerning the charts based on data compiled for species of lesser stature. As a whole, they appear less convincing as graphical representations of tolerance ranges than those based on tree data. A sizeable group of shrub and herb species show no evident correlation with the continuum index sequence established in this study. This might be attributed to a capacity of accessory species to respond to minor environmental variations not reflected.by the canOpy layer, as Pbtzger and Friesner (1940a: 168) believe. Such a view does not appear to be tenable in the present case, however, Rather, a reverse relationship is indicated, at least for the more ubiquitous and widely ranging members of this group such as fiibes gyposbgti, Egzthenogiggug 154. guingugfglgg, Smilgcina,racemo§g and Geranium.maculgtum. These Species are evidently capable of thriving under a great diversity of edaphic and microclimatic conditions and as such might be considered characteristic species of unistratal communities or unions (Lippmaa 1939). Such evident adaptability would likewise appear to support Cain‘s (1944: 22) contention that as a rule a species "consists of thousands of biotypes . . . sorted out on a combined habitat-tolerance basis.” However, the majority of non— correlated accessory species are of sporadic occurrence. While the fre- quency and DFr curves for such species are in consequence scarcely sugges- tive of a multiplicity of adaptive biotypes, neither are they indicative of narrow tolerance range, since most of them range over the greater por- tion of the continuum index scale. It may thus be observed that while the non-correlative species of either ubiquitous or sporadic occurrence do not directly support the con- tinuum concept, neither do they in any way refute it. They emphasize the futility of attempting to recognize discrete types of oak upland on the basis of phytosociological indices such as presence, constance, or fidelity. A further contention, supported here, is that the absence of a given species from a given stand may be considered attributable to reasons other than environmental control when the index position of that stand lies well within the limits of the frequency and/or DFr curve for that species. This view permits the majority of accessory species of the oak upland community to be regarded as correlated with continuum index sequence; likewise the frequency and/or DFr curves of such species may be considered expressions of tolerance range. Such ranges ferm.typically an overlapping gradational sequence similar to that of the tree dominants. 155. The lower synusiae of a forest community exist in an environment greatly modified by influences attributable primarily to the canOpy layer. There- fore, the fact that the tolerance ranges of most of the components of the lower synusiae should occur in a sequence paralleling that of the tree dom- inants constitutes support from a different quarter for Kittredge's (1948) repeatedly emphasized view that site, overstory—composition, and forest influences are strongly interrelated. Such a parallel, of course, supplies further evidence of the validity and.value of the continuum concept. It likewise reaffirms the utility of accessory species as indicators of physi- cal environment, though there appears no reason to suppose they are in any way superior to the tree dominants in this regard. On the contrary, the most widespread and abundant accessory species are evidently of no value as indicators. MOreover, many species correlated with index sequence are of limited utility because of their restricted occurrence. The Copggpt of 011mg; Adaptation Number Perhaps the most controversial assumption involved in the continuum concept is that all the adaptive functions which collectively determine the relative degree of "climaxness" of a given species may be expressed by a single number. The climax adaptation number of a species reflects the relative position of its sector of Optimum development in a subjectively determined order of stands. Such a sector does not necessarily reflect Optimum physical environment for the species concerned. In the case of a leading dominant it could, and probably more often does, represent a range more or less removed from the sector of Optimum physical environment by the influence of competition from other leading dominants. In consequence, 156. the sector of Optimum develOpment for some species is located near one end of the tolerance range. In view of the biotypic and ecotypic richness of species in general, it would seem scarcely credible that the whole p0pu~ lation of such species could be represented by a single climax adaptation number. Camp (1950) has indicated that the wide distribution or Eggug m. _ifg_l_i_.5 and Age; W is not a reflection of uniform climate, but rather is a manifestation of the great ecotypic diversity of these two species. Each of the three ecotypes of E. W recognized by Camp appear so distinctive in their ecological requirements as to merit a separ- ate climax adaptation number. Note has previously been.made of apparent ecotypic variants of m m and W W (in southern Mich- igan. Similar habitat forms of these species evidently occur in Indiana (Dean 1940), while in neighboring Ontario only the moist site variants are found (Anonymous 1949). The strong correlation between site and type of root system develOped by m m has been emphasized by Toumey (1929) and Kramer (1949). Such adaptation doubtless contributes to the unusually wide tolerance range of this species, one of two species which occurs in all the forest types recognized by Sampson (1930: 362) for northeastern Ohio. The capacity of red.maple to develOp well over a wide range of light intensities (Billings 1938) further complicates the task of designating a climax adaptation number for this Species. A suggestion of two trends with considerably displaced maxima has been noted previously in the curve of significance value for Qgggug flgzidg. The possibility must be considered that these are indicative of the existence of two biotypes having differ- ential potentialities for vegetative reproduction. The interruptions in 157. the curves of importance value for M gpgpdifglig and W tglipifggg are of obscure significance. Camp (1950) has indicated that the pOpulation Of beech north of the glacial boundary is genetically highly complex, a condition not conducive to the isolation of beech eco- types noted elsewhere. Tulip poplar in the oak upland community is appar- ently at the xeric edge of its tolerance range (Sampson 1930) and could therefore be expected to occur Sporadically. Moreover, as previously noted, its absence in some stands may be attributable directly to the sel- ective cutting which occurred on an extensive scale in pioneer days. Never- theless, the possibility that the curves for these two species may reflect ecotypic isolation remains. From the above examples it is obvious that if climax adaptatign num- here are to have maximum significance, studies directed toward determines MFCW“ ..m—uwmn ”...— --r-W' ‘ 'M.... tion of the physiological and.morphological characteristics of ecotypes and seeds must be made for each species. Camp‘s (1950) analysis of the american beech could well serve as a pattern for such studies. H Succession t tu of the U land unit Data and Observations bearing on the successional status of the oak upland community in southern Michigan have been presented in previous sec— tions of this report. It is the purpose of the present section to integrate and complement these data and observations with pertinent information from other sources. As Curtis and MOIntosh (1951) have said, the word fplimax" has a plethora of meanings. This ambiguity, together with interpretational dif- ficulties of other nature, has prompted some workers to abandon the term 158. and sometimes even the concept. Such a negative approach, however, seems scarcely justified. Certainly succession is not a universal process; wherever it has culminated in a vegetation in essential equilibrium with the other components of the ecosystem, a descriptive tenm to desig- nate that condition of near-stability is meaningful and necessary. NO term seems more apprOpriate than climax to denote such a condition. An Obvious need exists, however, to isolate the term and the concept from deductive theory of the sort which Egler (1951) has referred to variously as speculative philOSOphy, ecological dogma, and one-factor ecology. Accordingly, an attempt has been made below to assemble a background of pertinent data by which the oak upland community in southern Michigan may serve as a.measure Of the Objective reality of the various climax hypotheses currently used in interpreting vegetation. The Oak Upland Community as a Test of Climax Hypotheses Braun (1950), a preponent of the monoclimax hypothesis, indicates almost all of southern Michigan to lie within the boundaries of the beech- maple climax. This is to be interpreted as indicating that on well- drained, but moisture-retentive soils of intermediate texture and mixed mineralogic canposition the primary forest is usually of the beech- maple type. As Cain (1941: 193) has so aptly said, the climatic climax is indeed "merely the edaphic climax of non-extreme sites." Implicit in the monOclimax hypothesis, however, is the added proposition that the natural vegetation of all other sites in southern Michigan represent successional stages trending toward the beechemaple type. According to this assumption, the Oak upland.community represents the subclimax stage of the xerosere. The question naturally follows: through the action of 159. what influence or influences is the oak upland community in southern Mich- igan supposedly destined to be replaced by a beech-maple climax? Granting stability of climate and tolerance ranges of the dominants concerned (as the monoclimax hypothesis requires) plant reactions might conceivably effect such a change. Within what interval of time might one reasonably expect the potentialities of plant reactions to be fully expressed? Major (1951: 398) believes that successions of the type attributable to plant reactions are capable of stabilizing "most vegetation in less than 1000 years." In a study of succession initiated on abandoned upland fields in North Carolina, Billings (1938) found that reproduction of climax hardwood species was well established in little more than a century and in position to replace members of the pioneer overstory whenever a natural Opening should occur. Moreover, correlated observations of soil profile develOp- ment revealed that in as little as 30 years all evidence of a plow layer had disappeared, a typical forest floor had accumulated and a humus-rich A1 horizon had develOped to a thickness of three inches. Concerning the recent intrusion of forests in the Ozarks, Bellman and Brenner (1951a: 261) write: "This time-elapse study of only 12 years revealed a speeding succession of plant species not at all approaching the accepted trial- ’ andperror elimination which is supposed to set the pattern for our forest areas.“ Of course, a primary succession nearing the point of approximate stability must be expected to progress at a slower rate. Nevertheless, if the potentiality of a given oak upland site could be raised through plant reaction to a level such that it could support a beechemaple type, there is no reason to believe that such change in potentiality would re- quire the cumulative reactions of repeated generations of oak upland dom— inants. 160. Dominants of the oak upland.type do not produce reactions directly inimical to repeated occupancy of a site by others of their kind (Braun 1947: 216). Accordingly, the only influences conducive to possible in- vasion by beechpmaple elements would appear to be those which tend to better the pgzgiggliihialgsisal, and chemical canditiqn of lbs 8011- Such amelioration is effected primarily through the medium of the forest floor. The amount of forest floor under hardwoods does not increase indefinitely (Kittredge 1948; Ovington 1953). Kittredge (1948: 171, 174) indicates that in a given stand a balance between accumulation and decomposition may occur at about the time of culmination of growth. After that period the amount of forest floor may actually decrease. The forest floor serves as the principal source of the humus which becomes incorporated in the upper>mineral soil. It would seem, therefore, that any tendency toward stabilization of the forest floor would be reflected in a tendency toward equilibrium in the.A horizon between melanization on the one hand, and eluviation and decomposition of organic matter on the other. This cone tention would seem borne out by the fact that in upland forest sites in southern Michigan the melanized.A1 horizon very rarely ever attains a thickness of more than two or three inches (Veatch, 2i.£l¢ 1941: 36). Thus, the view of Quick (1924) that the beechpmaple association is :)_",5 ncapable of occupying all the soils of theI§EEEE:§E:EEEE].1.W. [because]. . . . the water retaining qualities of the soil may be increased or decreased by the addition of humus . . ." YEE£E_EEEE~ES—£§9kra factual basis. ‘ Concerning chemical influences of the forest floor, it may be said in general that, except for potassium, the leaf litter from oak upland 161. dominants return relatively low amounts of the majoerla t nutrients to -~—u "' .1 “In" ..lm the soil (McHargue and Roy 1932; Alway, Kittredge and Methley 1933; Alway, Maki and Methley 1934; Coile 1937; Broadfoot and Pierre 1939; Chandler 1941; Kittredge 1948). It follows, therefore, that occupancy of a site by such dqmipgnts,-no matter how long, could hardly be expected tg~increase‘sgil_£ertility to the level rquEEEEQEZMPE?Ch andsugarlmaple. On the contrary, the findings oszorham (1953: 148) would seem to indi- cate that plant "pumping" and flushing from above does not even keep pace with the acid production and leaching promoted by litter from oaks. Ad- mittedly, however, certain understory Species, notably Eggpgs florida and gstgyg‘vigginiana, tend to produce lEEPQE~§i9§m19 minerals (McHargue and Roy 1932; Broadfoot and Pierre 1939; Kittredge 1948). Yet, the yield of litter derived from such species must be small compared to that of the overstory species. As Billings (1952: 261) has said, cumulative influences are attributable in the main to the principal dominants. Furthermore, Kittredge (1948: 176) emphasizes that Species yielding litter high in calcium, nitrogen, and phosphorus "are those which are associated natur- ally with fertile soils usually well supplied with calcium." It would seem necessary, therefore, to regard such species as tending to maintain fertility rather than to build it. The above considerations, viewed in the light of the pollen record for southern Michigan (see section on postglacial history), clearly indi- cate that the past has provided.more than an ample test of the capacity ...-...- 1 ll.“— . FM-..“ m w of plant reactions to effect succession from oak upland to beechemaple forest. The fact that in southern Michigan oak uplands remain today the most widespread forest type, and beechsmaple the least (Gysel and Arend 162. 1953), supports the contention held here that such capacity is in general laij-ng o A second possible means by which the theoretical monoclimax in pa. 7:: ...—urgi- southern Michigan might be attained is through the cumulative effects of weathering and podsolization on oak upland sites. Lutz and Chandler (1946: 85) state that soils developed on Wisconsin till are frequently immature. However, there is no reason to believe that soils supporting oak upland forest are further from that theoretical condition of near stability than are those of beechpmaple sites. On the contrary, in.view of the prevailingly greater perviousness of oak upland soils and resulting greater leaching, the reverse should be true. Such a contention is sup— ported by the findings of Gorham (1953). Nevertheless, the processes of weathering and podsolization continue, and according to Wilde (1946: 63), both tend to increase water holding capacity. That the influence of pod- solization in this regard.may be less than generally supposed, however, is suggested by chemical analyses of Michigan soils which indicate that most of the clay in the waterbretentive B—horizon is either inherited from the parent material or is the result of weathering in place (veatch 1953: 27). Moreover, even weathering has failed to produce any semblance of a clay horizon in certain widespread soils of the region. These include the Cglgma_and~§1ainfield types formed from highly siliceous parent materials. It seems inconceivable that any process short of removal of the present surface through geological erosion could raise the potentialities of the afiééflOflmhhéfihWLB—mh figoped ’00 themes anyways. Furthermore, even geological base-leveling does not necessarily pro- duce vegetational uniformity. Cain (1947: 193) emphasizes that "even on a peneplain there are site differences and corresponding vegetational 163. differences." Thus, even within the dimension of geologic time there is ”V” ”’9‘”- ' “TM" 5. ....~_ .- no reason to suppose that the influence of climate will be expressed over —— w' I H I I“, JI"_-"m" UH'M’MWN‘” all the local habitats of the region in the form of a Single vegetational *mmF—“M “hm-'9‘-— -.-t .- --<- H» .w- ...,“ u- u! ""' *‘fi- "myfll. ... '~Mevw'- ”4' Itzpe. Mereover, the monoclimax hypothesis ignores the simple truth that climate is clearly the least stable of all the major environmental vari- ables. Veatch (1938) has noted the occurrence in southern Michigan of relic podzol soils dating back at least to the pine period, as well as relic hydromorphic soils antedating the thermal maximum. Leverett (1909), in considering the effectiveness of geological erosion and stream dissec- tion on Cary drift, estimated that "scarcely one-tenth of the surface has been reduced below the original level as a result of drainage." In con- trast to these indications of edaphic and topographic stability, Sears (191.3: 331) has listed a total of six general climatic shifts since recession of the Mankato substage of Wisconsin glaciation. Surely Cain (1947: 193) is justified in his belief that "the monoclimax hypothesis has been not so much an ecological touchstone as a.millstone." Mm If the monoclimax is untenable, what of the one-factor climaxes recognized by the polyclimax school? The lack of any great physiographic contrast in southern Michigan precludes any possibility of the differentia- tion of physiographic climaxes in thiswpartmpf the state. Here, local relief features, insofar as they influence soil moisture, are far*more limportant than physiography as a determinant of natural vegetation. In general, the oak upland type__ in southern Michican is Ioevelooed ’C -'-..v-~\-\-V‘I' on.more pervious and less fertile soilsI than thosIe supporting beech-maple. .va—gvrzh --'v--"" . « W'- 'fib" use ...” Yet, certain soils, such as the Miami and Hillsdale series, are pivotal * s.- “gm-”b n,” -..-mung- n-f'" —~...- I in that they may support either oak upland or beech-maple types. Moreover, ...-- .... h—qm—..,..--. \mwfl“ -"—-’~I-‘ Malibu-d iw ./ 7'1"" ... i 164. the lee lepes of sand dunes along Lake Michigan support vigorous sugar maple-basswood stands (Cowles 1899). Whether or not such phenomena are ‘ attributable even primarily to physical or chemical influences of the soil has not as yet been established. Thus, any reference to the oak upland type as an edaphigflglimax.must be made with reservations. The role of fire in maintaining the oak upland type in southern _‘ ‘W m.— ' ‘ ' " " ‘ _ yaQfi" - u m. Int-'1 h’“ ’ *"h-uuwflW-m an... - .."h'hé‘nzi Michigan“ is obscuie. Lanman (1839), Hubbard (1887) and Beal (1904) speak casually of the occurrence of early fires as though they were established facts and without need of confirmation. The first two authors imply that they were annual events during the period of Indian occupancy and had the effect of keeping underbrush in check in the oak Openings. Lanman (1837: 324) states: Each kind of Opening is subject to what are called grubs. These are formed.by the fires which annually run through the woods, and burn the tops of the vegetation, leaving a root sometimes three feet square, and is firmly imbedded in the soil. Six yoke of cattle are frequently required to tear up these grubs, which is done by the plough. Certain inferential evidence still extant provides additional support for the belief that recurring fires may have been at least locally effective in modifying forest composition and structure (see Fig. 42 and further in the next section). On the other hand, the land survey notes for Ingham County, com- piled during the period of Indian occupancy in the years 1825 to 1827, include frequent reference to dense underbrush in the oak upland areas and.make no reference to fires. Moreover, the occasional occurrence of witness trees of such fire—sensitive Species as Eggnug serotina, Qgtgyg {gigginigpa, and Acezngbzymbin the oak upland areas suggest further that ‘ n .fh} . \Qb This tree, b.h. and probably more than 300 years 01’, eeent closed 4 ore the Dr so e-spreaolng crown. .. 1 ing a Wih f (Photo by the author.) Old growth white oak with enonmous lower at d evidently reached maturity be . 42. branches form FIG can0py developed. 52 in. 166. fires were not important in determining forest composition in this par- ticular county. It seems necessary to conclude, therefore, that the ‘ influence of recurring fire was not universally manifested in the oak .._ J...” ...-nu, .. V._.__— ' 'ma—n—er—I— -- uplands, but rather was probably confined mainly to the more xeric areas -n.’ \—-I- as determined by topography and soil. The intervening areas, by serving -v-v-—— rum-N“ """“ as barriers to fire, doubtless tended to prevent regional conflagrations of the sort which evidently occurred regularly in the prairie-forest border to the west (Marks 1942; QQEEFm’1949; Braun 1950; Beilmann and Brenner 1951a). It is evident, therefore, that ngmgeneral reference to _ the °§§;E£$E§§43XREH n southern Michigan as a fire.Qlimax is permissible. From the discussion above, the general conclusion follows that the éoak upland community is not subject to interpretation as a single-factor W climax of any type. To attempt such interpretation would reflect disre- gard for the basic truth expressed by Billings (1952: 263) that "vegeta- tion is an indicator of the whole environment and not just of climate, or parent material, or fire, or any other single factor." Comprehending this fundamental truth, while at the same time pro- viding for the "local relation of community gradients to environmental gradients," as well as recognizing the relativity of the concepts of suc- cession and climax, the climax pattern hypothesis recently advanced by Whittaker (1953) would seem to fit best the facts as revealed by this study. 'Moreover, it has the necessary conceptual breadth to relate the oak upland continuum to other elements of the vegetational mosaic both in time and space. It has therefore been adOpted as the working hypothesis for interpreting the inferential evidence assembled in the course of this study. 167. Past and Present Reproduction Trends in the Oak Upland Community Rejection of the monoclimax hypothesis in favor of the climax pat— tern hypothesis permits consideration of the possibility that the oak upland continuum may;repr§sent¥climaxiyegetation”forithehrange of sites wow-n- on which_itmi§w§ev§lgpeév The fEEEHEh§t oak uplandmforest is today the most wideSpread forest type in southern Michigan, together with evidence that this ranking has been.maintained continuously since the close of the piggnsriosi, clearly attefis .130--Eh?.-PaPa931131.“??thiS..J‘Jpe tommaintaqin itself indefinitely. It would thus seem to satisfy the qualifications ‘_ —— of a climax; ‘Yet, the results of this study as well as those of others (Wood 1930; Young and Scholz_l949; Arend, §t_al, 1950; Gysel and Arend 1953) indicate that reproduction of the principal dominants is in general strikingly low compared to that of the accessory dominants and understory trees. Nevertheless, in the light of the obvious success of the oak up- land type in persisting up to the present time, it would seem exceedingly presumptuous to infer from these results any marked decline in the capacity of oak upland forest to maintain itself on present sites. On the other hand, any serious attempt to support the contention that reproduction of the leading oak upland species is adequate to ensure continuation of_the V'— present level of dominance of these species can readily degenerate to Special pleading. It is evident, therefore, that inferences concerning future composition based on present reproduction trends are subject to severe limitations. There is no intent here to disparage inferential reasoning as such. The point is that to be really effective as an ecolog- ical tool it must be used against a background of ecological fundamentals of the type outlined by Pelton (1951). As yet, little autecological 168. information of this sort is available concerning the components of the oak upland community in southern Michigan. Without it, only tentative conclusions may be reached. In part, the dearth of oak reproduction may be attributed to rela- tively low seed production compared with that of other trees (Korstian 1927: 105). In more important part, it is due to extremely heavy utili- zation of acorns by insects, rodents, and birds. Insect infestation is a prime source of acorn destruction in southern Michigan (Allen 1943; Gysel 1953). Gysel's data, obtained from two Clinton County woodlots, show the following incidence of damaged acorns: white oak 84 percent, black oak 62 percent, red oak 43 percent. Of the damaged acorns, 81, 78, and 67 percent, respectively, were insect infested. To the toll of insects must be added the consumption of viable seeds by rodents and birds. Experiments conducted in a Washtenaw County woodlot by Cahalane (1942) indicate that by spring, squirrels may recover as much as 99 percent of the nuts buried the previous fall. Evidently only a small prOportion of acorns survive to produce seedlings. Yet, the results of this study, though based on too short a period of observation to be really conclusive, suggest that oak reproduction is limited not so much by small yield, or great destruction of acorns as by failure of most seedlings to survive beyond the class 1 stage. Actually, class 1 seedlings of the three leading species of oak compare favorably in number with those of other species (see Appendix Tables XXXVIII to LXXIV). Under especially favorable circumstances, seedlings of this class may even be abundant (see Fig. 43). In size classes 2 and 3, however, there is a progressive and marked decrease in oak reproduction relative to that ' , :rf' / FIG. [3. Seedlings of Quercus rubra developed from acorns concentrated by gravity on the floor of a moist ravine. 170. of other species. The latter chass which, theoretically at least, should be most likely to presage future stand composition, includes far less oak reproduction than any other class. There is also a marked ten- dency for oak reproduction of classes 2 and 3 to be of low vigor. Indeed, much of the class 2 reproduction of Quezcus alba represents seedling sprouts, including some of the second or even third order (see Fig. 44). The decline in number and vigor of oak seedlings in the more advanced reproduction classes is not a universal phenomenon. In larger clearings within stands, or in abandoned fields bordering oak upland stands, reproduc— tion of all three size classes is usually ample and vigorous (see Figs. 45 and 46). White and, eSpecially, black oak is able to invade such areas even in the absence of litter which Korstian (1927) holds to be so important for survival of oak seedlings (see Fig. 46). The conclusion would seem to follow that oak reproduction in general has a low survival capacity under the conditions of low light intensity and strong root com- “ "“*-1---.._.....,,..,..,,,. i,_,__,..m..,... .. .t . . -- ...-............ ..A :..— ...-...- --.‘- petition prevailing under closed canopies. The fact that somany class 1 seedlings of oak are to be found would appear to be explained by Baker's (1950: 140) observation that seedlings derived from large trees are com- paratively tolerant for the first few years because of the initial advan- tage of a large food reserve. Such a conclusion raises the question of how closed canopy dominance by upland oaks is attained and maintained. There is ample evidence to support the view that the canopy of the primary oak forests in southern Michigan was, in general, of more Open character than that of present secondary stands. Such evidence is to be found in historical accounts (see Literature Review), in the original land survey notes, and in the FIG. 44. Sprouts developed at the crown of a class 2 individual of guercus alba which evidently died back in consequence of inability to compete under closed canopy conditions. FIG. 45. Reproduction of cus zeluting under a seed tree growing in a clearing. (Photo by authorj FIG. 46. Reproduction of @9rcus velutina established on bare mineral soil in an abandoned field adjoining a black oak stand. 174. general observation that with advancing age a canopy intercepts less solar radiation (Shirley 1943, Kittredge 1948). Acorn production under such conditions must have been far heavier than now, since trees having ample crown space fruit much more regularly and far more abundantly than . those growing in close stands (Toumey and Korstian 1937; Baker 1950). Beilmann and Brenner (1951a: 273) report that even in woodland rated as "understocked" many oaks have produced no fruit in ten years, while iso- lated individuals may bear heavy crOps in alternate years. It seems very likely, however, that then (as now) production and utilization of fruit were in close balance. Well-documented reports of enormous squirrel populations in pioneer days (Allen 1943) suggest that, even when the great reduction in forest area is taken into account, squirrel pressure on the acorn supply was no less in presettlement times than at present. Any advantage to oak reproduction through more Open growth would seem to be restricted, therefore, to better survival conditions for seedlings. Conditions for survival of reproduction in the oak Openings were evidently far from optimum in the light of the accounts (Lehman 1839; Hubbard 1888) indicating that these areas were swept by periodic fires which kept down the undergrowth and favored formation of sod. On the other hand, the relatively sparse crowns of old growth oak canOpies may have admitted sufficient light for effective development of dispersed oak reproduction. That decreased interception<>f solar radiation with advancing age of stand may effectively influence reproduction of some species, at least, is indi- cated by Shirley (1943: 341), who speaks of the vigorous growth of repro- duction possible in aspen and pine forests "after they have passed their prime and.begun to Open up naturally." Kittredge (1948: 51) indicates 175. that the maximum interception of solar radiation in well stocked stands coincides approximately with the culmination of annual increment and thereafter decreases. The greatest annual increment for most Species occurs within 60 years after establishment (ibid: 32). While, in general, the stands included in this study are older than 60 years, the overstories remain relatively dense, and eSpecially where associated with well developed understories of Qggggg florida andlggtryg virginiana, the amount of intercepted radiation is great (see Fig. 47). In these second growth stands occasional evidence of Sprout regeneration of canopy individuals is to be observed (see Fig. 48). However, the great bulk of the dominant individuals seem to be of seedling origin. This fact suggests that such individuals represent elements of the first regeneration following removal of the old growth, since recutting of second growth could be eXpected to result in a preponderance of sprout reproduction (westveid 1939). The added fact that in a given stand so many of these individuals appear even-aged suggests further that reproduc- tion of oaks was well distributed and established at the time of the original timber harvest. Such reproduction need not necessarily have been of high vigor. Beal (1888: 76) has called attention to the extreme tenacity of seedling sprouts of black and white oak, and Baker (1950: 6) has stated that such Sprouts may develOp into trees comparable in stature to those originating directly from seeds. As Gysel and Arend (1953: 14) have suggested, heavy cutting could be expected to result in compositional differences between old and second growth stands on a given Site. The net effect of such cutting would seem Inost likely to be reflected in a lowering of the continuum index number for the site, owing to the initial advantage provided less tolerant Species. FIG. 47. Heavy shade cast by a second canopy of mature Comus florida. Where such a canOpy layer is developed under an overstory of oaks, herb and shrub representation is scanty or absent altogether. snuurmmu 1, ’1 lift; .1 Nita—w. 1,. 3. I . d... ... Trees of obvious sprout origin as indicated by multiple bole 480 FIG. development 178. Evidence in support of this contention is to be found in Table XV, where the prOportion of black oak in an entirely second growth stand is seen to be considerably greater than in an adjacent stand containing considerable old growth. It must not be inferred, however, that black oak was an uncommon tree in oldpgrowth oak uplands. A check of the records of the original land survey for Ingham County reveals that, in the oak upland areas in the southern part of the county, black_oak was second in importance to white oak. Significantly, however, in almost all cases where all witness trees for a corner were black gaks, the distance from stake to trees were considerably greater than at those corners where white or red oaks served as witness trees. Many of the distances from corner stakes to black oak- witness trees exceeded two chains (132 ft.). It seems obvious, there- fore, that the density of black oaks in present stands is much greater than it was in old growth stands. The resulting greater interception of radiation in such second growth stands doubtless accounts for the almost complete failure of the reproduction of this Species to attain class 3 size in well-stocked sites. Advanced reproduction of certain accessory dominants of the oak upland continuum tends to be as conSpicuous by its abundance as that of the oaks is by its scarcity. In the majority of stands Qgryg,gzali§, Aggr rubrum,,§runus serotina, and.Fraxinu§ gmericana collectively comprise the bulk of such reproduction. All four of these Species are capable of developing deep and extensive root systems in the characteristically per- vious soils supporting the oak upland type (Van Dersal 1938; Harlow and Harrar 1950). Supposedly, therefore, they Should be able to compete 179. effectively with the dominant cake for nutrients and water. Yet, the results of this study reveal the importance potentials of these species to be decidedly low compared to those of the oaks (see Table VI). Data provided by Curtis and.McIntosh (1951: Table 1) indicate that these species are also to be regarded as minor dominants in the upland forests of south- ern Wisconsin. The failure of Ergngg serotina and Fraxinug americana to attain high importance values despite abundant reproduction of all three size classes may be attributed to inability of these species to withstand sup— pression, except when young (Harlow and Harrar 1950; Gunther 1950). Indi- viduals of these Species which had attained class 3 and even class 4, size before dying were repeatedly encountered in the course of this study. The low importance potentials of m M and Age}; m cannot be explained so simply. Both Species are frequently well represented by vigorous individuals of 8128 class 4. Evidence of this sort, as Billings (1938) has pointed out, seems strongly suggestive of high survival capac- ity leading to greater importance in the future. Evidence concerning the status of pignut hickory and red maple in the primary forests of southern Michigan is inconclusive. Witness trees of these species were recorded only sporadically in the original land survey of Ingham County. On the other hand, Beal (1904) described a virgin oak woods in this county in which red maples up to 24 inches in diameter formed a conspicuous element. The possibility must be considered, therefore, that red.maple, at least, may be in the process of returning to a former level of importance in the oak upland continuum. Although Steyermark (1940) has described a white oak-red maple association for the Ozarks, it is doubtful, however, whether 180. red maple will ever merit recognition as a major upland dominant in south. ern Michigan. Otis (1931) describes this species as a tree not exceeding 50 feet on upland sites in Michigan. This limit is well below the upper canopy formed.by its upland associates. It would seem more logical, therefore, to regard it as a secondary tree component Of the oak upland type, a view previously expressed by Cain (1936), Billings (1938), and Costing (1942). To the uncertainties involved in inferential evidence attributable to lack Of information concerning the ultimate fate of trangressives must be added those arising out of the relativity of the concepts of succes- sion and climax. Cowles (1901: 81) has referred to succession leading toward a climax as "a variable approaching another variable." This com- parison highlights with particular clarity the difficulty experienced in attempting to interpret a given inferential trend of reproduction as evidence of succession or simply as ”fluctuation about an average" that represents the climax condition (Whittaker 1953: 61). The Opinion is expressed here that for the most part reproduction trends for tree com- ponents of the oak upland continuum are best interpreted as of the latter type. With few exceptions they appear to involve conserving or consolidat- ing species in the sense of Braun-Blanquet (1932: 316). This would appear true of the trends for Qggyg‘gyglig and App; gpbggp considered above. Similarly the xeric maximum in the curve Of significance value for Qgercus glbg (see Fig. 7) probably merely reflects a trend toward the compositional pattern which presumably prevailed in the primary stands on sites that now support stands largely dominated by'Q.'velutina. Definite evidence of succession involving dominants potentially 181. destructive of present compositional patterns was obtained in a few instances, however. The most striking examples are provided by the class 3 and 4 representation of App; saccharum and,§gg2§ grandifolig in stands 30, 34 and 36. All three stands are develOped on soils which most frequently support the beech-maple type. Since stands 30 and 36 are secondary, the possibility may logically be considered, therefore, that present reproduction trends may, in these two instances, merely reflect secondary succession toward the type which prevailed before timber removal. Stand 34, however, was in essentially primary condition at the time quadrat data were Obtained. It has since been cut over and whether or not beech and sugar maple would have assumed dominance in the course of natural events will, of course, never be known. The evidence, however, particularly the suppressed but vigorous understory of sugar maple, strongly supports the view held here that such a change would have ultimately occurred. Braun (1950: 319) has interpreted a strikingly similar canOpy condition in oak-dominated parts of the Russ Forest in Cass County as evidence of succession toward beech—maple. Like stand 34, the oak-dominated portion of the Russ tract has been little disturbed by cutting and is develOped on soils typically associated with beech—maple cover. That both sites concerned have the capacity to support beech-maple as dominants would seem.adequately established by the vigorous understory of these Species which seem in position to assume dominance whenever a canOpy Opening should occur. The most puzzling aSpect of these stands, latherefore, is not that they should be trending so strongly toward beech- Inaple at present, but rather that dominance by these Species should have been deferred so long. The action of some retarding factor or factors 182. seems indicated. The occurrence of fire-scarred, Old growth individuals Of Liriodendron tulipiferg within stand 34 and.in a tract adjacent to the Russ ForeSt suggests the hypothesis that fire may have been instru- mental in retarding the development of beech-maple on these sites. Sup- port for this hypothesis is provided by Beal (1904), who attributed the evidently similar invasion by sugar maple of an oak woods in Ingham County to cessation of fires. In summation, the inferential evidence considered in this study suggests that reproduction trends in the oak upland continuum prevailingly reflect the sort Of variation around an average associated with the climax state. In those few instances where definite evidence of succession exists, the forces involved seem to be traceable to the activities Of man and to have been set in motion by events of the recent past. 183. SUlvfl'vMRY AND CONCLUSIONS 1. Examples of oak upland forests in 17 counties Of southern Michigan were studied by the quadrat method of sampling. Data on tree, shrub and herb representation were Obtained from nested quadrats located in 36 ...-i stands. 2. Importance values were computed for each tree species in each stand. These values were weighted by climax adaptation numbers to yield continuum index values ranging from 751 to 1999. 3. Data representing tree, shrub and herb composition of each stand were charted in a sequence determined by continuum index numbers. The resulting curves were regarded as graphical expressions of tolerance ranges for the species concerned. NO tendency for such ranges to occur in patterns suggesting discrete combinations of species was observed in any habit category. 4. The charts based on tree data reveal that stands Of the oak up- land community in southern Michigan form a continuous cline differing, even at the extremes, less by kind of dominants than by prOportions of dominants. This was regarded as a result of the broad overlapping of tolerance ranges Of‘most tree species. 5. Trends in representation Of the majority of shrub and herb Species Show definite correlation with continuum index sequence if indicator significance is accorded only to the presence of Species in 184. a given site. Among those Species which Show no evident correlation with continuum index sequence are certain of the more ubiquitous species, as well as those Of too sporadic occurrence to yield conclusive trends. Such species are held to be of no utility as indicators of site. 6. A gross correlation was observed between continuum index sequence and the number and frequency of Species in both the Shrub and herb synusiae. Within the limits of the continuum index scale established for the oak upland community in southern Michigan, the number and frequency of com- ponents of the herb synusia varies directly with continuum index sequence; an inverse relationship is indicated for the shrub synusia. 7. The leading pairs and trios of dominants in each stand were cor- related with continuum index sequence in order to determine the validity of each.as bases for empirical classification of oak upland stands. For this purpose twO leading dominants are held to be superior to three because Of the fewer number of unduplicated combinations and the narrower overlapping in scale position of the stand groupings characterized by the same leading dominants. The scale sector between index values of 1350 and 1400 was tentatively recognized as a boundary separating stands having black oak as one of the two leading dominants from those in which red oak has a similar position. 8. The influence of certain environmental variables on continuum index number has been noted as follows: a. In those few instances where other environmental influences could be judged to be constant or to vary ineffectively, stands 185. developed on lepes of southern aspect had significantly lower con- tinuum index numbers than those developed on slopes Of northern exposure. b. Under similar conditions of l-factor variation, and except on north lepes, severe disturbance was likewise reflected by a significant lowering in continuum index numbers. c. Despite effective variation of other environmental influences, a general correlation between continuum index number and Epilmtextnre waswnoted. Such gross correlation permitted the designation of index position 1,500 as an approximate boundary, in terms Of vegetational response, between sands and sandy loans on the one hand and loans M and silt loams on the other. 9. The results Of this study were supplemented and complemented with data from other sources in an effort to test the validity Of the various climax hypotheses as applied to the oak upland continuum in southern.Mich- igan. The climaxepattern hypothesis of Whittaker was found to agree most closely with observable facts as Opposed to speculative theory. 10. Few reproduction trends indicative of important changes in stand composition are evident from the results of this study. Such as occur seem attributable to events Of the recent past and not to developmental trends produced by plant reactions. 186. LITERATURE CITED Ackerman, E. A. 1941. The Kbppen classification of climates in North Allen, D. L. 1943. Michigan fox squirrel management. Mich. Dept. 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A phytosociological study of the herbaceous plants in two types of forests in central Indiana. Butler Univ. Bot. Stud. 4: 163.-180. . 1940b. What is climax in central Indiana? A five-mile quadrat study. Butler Univ. Bot. Stud. 4: 18l-—195. , and I. T. Wilson. 1941. Post-Pleistocene forest migration as indicated by sediments from three deep inland lakes. Am. Midl. Quick, B. E. 1924. A.comparative study of the climax associations in southern Michigan. Pap. Mich. Acad. Sci. 3: 211,.244. Raup, H. M; 1937. Recent changes of climate and vegetation in southern New England and adjacent New York. Jour. Arnold Arb. l8: 79--ll7. . 1947. Some natural floristic areas in boreal America. Ecol. Monogr. 17: 221-234. Sampson, H. C. 1927. The primary plant associations of Ohio. Ohio Jour. Sci. 27: 301-309. . 1930. The mixed.me80phytic forest community of northeastern Ohio. Ohio Jour. Sci. 30: 358-367. Sears, P. B. 1925. The natural vegetation of Ohio. Ohio Jour. Sci. 25: 139-l49. . 1942a. Forest sequences in the north central states. Am. Jour. Bot. 29: 684p-691. . 1942b. Postglacial migration of five forest genera. Am. Jour. . 1948. Forest sequence and climatic change in northeastern North America since early Wisconsin time. Ecology 29: 326—-333. Shanks, R. E. 1942. The vegetation of Trumbull County, Ohio. Ohio Jour. Shirley, H. L. 1943. Is tolerance the capacity to endure shade? Jour. of Forestry 41: 339-345. 194. Smith, N. F. 1953. Oak wilt project. (Mimeographed Report) Mich. State Cons. Dept. 1 p. Steyermark, J. A. 1940. Studies of the vegetation of Missouri - 1. Field Mus. Nat. Hist., Bot. Ser. 9: 349-475. Strong, F. C. 1951. Oak wilt in Michigan. Mich. State College Agr. Exp. Sta. Quarterly Bull. 34: 41--47. Tansley, A. G. 1935. The use and abuse of vegetational concepts and terms. Ecology 16: 284-307. Toumey, J. W. 1929. Initial root habit in American trees and its bearing on regeneration. Proc. Intern. Congr. Plant Sci. (Ithaca) 1: 713—- 728. , and C. F. Korstian. 1937. Foundations of silviculture upon an ecological basis. Second Ed. John Wiley and Sons. New York. Transeau, E. N. 1905. The bogs and bog flora of the Huron River valley. . 1935. The prairie peninsula. Ecology 16: 423-437. Tuttle, C. R. 1873. History of Nfichigan. R. D. S. Tyler and Co. Detroit. Van Dersal, W. R. 1938. Native woody plants of the United States. U. S. Do A. MSG. Pub. 303. 362 pp. Veatch, J. O. 1928a. The dry prairies of Michigan. Pap. Mich. Acad. . 1928b. Reconstruction of forest cover based on soil maps. Mich. State College Agr. Exp. Sta. Quarterly Bull. 10: 116-126. . 1931. Natural geographic divisions of land. Pap. Mich. . 1932. Soil maps as a basis for mapping original forest cover. Pap. Mich. Acad. Sci. 15: 267-273. . 1938. Pedologic evidence of changes of climate in Michigan. Pap. Mich. Acad. Sci. 23: 385-390. _ . 1941. Agricultural land classification and land types of Michigan. Mich. State College Agr. Exp. Sta. Spec. Bull. 231. 67 pp. . 1953. Soils and land of Michigan. Michigan State College Press. East Lansing. 195. , 23 Q1. 1941. Soil survey of Ingham County, Michigan. U. S. D. A., Bur. Pl. Industry, Div. Soil Surv. Ser. 1933. Warming, E. 1925. Oecology of plants. Oxford Univ. Press. London. Watkins, L. D. 1900. Destruction of the forests of southern Michigan. Mich. Pioneer and Historical Society, Hist. C011. 28: 148-150. Weaver, J. E., and F. E. Clements. 1938. Plant ecology. McGrawaHill. New‘York. Westveld, R. H. 1939. Applied silviculture in the United States. John Wiley and Sons. New York. Wheeler, C. F., and E. F. Smith. 1881. Michigan flora. Mich. State Hort. Soc. Ann. Rept. 10: 427-—529. Whitford, P. B. 1951. Estimation<>f the ages of hardwood stands in the prairie—forest border region. Ecology 32: l43-146. Whittaker, R. H. 1951. A criticism of the plant association and climatic climax concepts. Northwest Sci. 25: l7-3l. . 1953. A consideration of the climax theory -- the climax as pOpulation and pattern. Ecol. Monogr. 23: 41-—78. Wilde, S. A. 1946. Forest soils and forest growth. Chronica Botanica. Waltham, Mass. Willett, H. C. 1949. Long-period fluctuations in the general circula- tion of the atmOSphere. Jour. Meteorology 6: 34,-50, . 1951. Extrapolation of sunspot—climate relationships. Jour. Meteorology 8: 1-6. Wood, L. M. 1930. Silvicultural management of the oak-hickory forest type in southern Michigan. Unpubl. M. F. thesis. Michigan State College. Yapp, R. H. 1922. The concept of habitat. Jour. Ecol. 10: 1-l7. Young, L. J., and H. F. Scholz. 1949. Some results of selective cutting in the Eber White woods, Ann Arbor, Michigan. Pap. Mich. Acad. Sci. 33: 39—-56. APPENDIX 197 . 5.2. Ha... m6 R SH «:3 2. m.$ EH 3:8 I I «.0 H I I I I I I «A H OH I I ence.—Hose nae—B .H.H H.H mé on I I I «Hm o 3 <23 m 3. ~.m mm 3.63.? Hangman «.8 mix“. «.0 mm «do mm 03 H.HN e on I I I «.0 H £3933 36.35 I I «.0 H I I I I I I I I I ~.o H manna 26.85 o.~ c.H «.3 5 RH H a.mH m 8 EH mH ow m.oH 8 and. 3688 H.o Hie cam 92 I I I I c I «H w on ER HmH 8323 SEE l I «.0 H I I I I I I I I «.0 H can. 239:. I I. «.0 H I I I I II I II I «.0 H .mm newcoveho o.H w.o o.o New I I n.0H m om 5.3 mm 00 v.9 mom 3.?on 3980 I I o.H o I I I I I 9m m 0H w.o e. 33.8w commando H.~ b.H n6 om m.H _H H.HN .H on 06 o 3 man mm 318 shame I I H.H b I _I I i I I I I I m.H 5 gen .82 l . . e i o l . o l o o o «t a i a a i o_ a a a a a a n Eamon ._._wzmo >.:mzmn , >._._mzmn >._._mZMO >._._mzmo m H. m .e. m4g~m6g66 g m 2 so s:aaoagsn~s~* g 6 ago" 8888888338838 m H H 3:3. 3 M g 2.1 3 u: E? g h- m P 3. $45 no t: g: o m hdmwhrw. Cancun: wmmfhflu Crataegus sp. hmmanflma mamadm finaunmma Uhmamfium Amrflhm muwamWa 201. O 1. v C a ‘o {I u! . n D . o.) O O .I‘n .1. t .. 0 Incl . .l . .'..I O‘ ol- 4. o .0 O I.) :0 5A 0 . ... o I: . n vllun 6 s., R. 15 m, 1H; NIH section 33. TABLE VI. Smary of tree data based on ten 100 m2 quadrats fran stand 6, Cass County, La Grange Twp., T. Acer saccharum Amelanchier up. Carya ovalis Coma florida Crataegus up . 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A o Ir-Iul Onlu‘ -l (0 11. 1 3., n. 2 11., 811i 111: 8 auction 17. Mary of tree data baud on ten 100 n2 quadrats fre. stand '7, J ackaon County, Haterloo Yup. , TABLE VII. tn NN coo~m~tm m E88 dléfiléfisgél —l _l : go. «8 HH «038283118 H O t- éldé'éndsd 3 ... HNQPMOHOMI‘F 3 588 gasfiagaasgg :1 go 88888888888 5:; O. m 6 E8 838§8§§2§§8 8 8 888 8 g5 III3|IN~0~KQ'I co go 111°11~8811 8 6 E8 11181188811 ” _ ° H 8888 8 "(‘E§o fii'gljgb‘n'" ~11 U) 2 :81: ”I!Sl‘*°‘°’"‘1l 8 I- d m E8 81181888811 (I) < o b. qt: "3“) o. . "2 .1 g& grangwligi 9. u 2 En 21°”lfi“ll‘°l 8 Ill : 8&0 81881881181 U) [I- 8 88888888888 8 E5 gonno§MMH8N 12 CD Ea S”2$"$33°§S g a' 538° 828§8§888§8 a 33 Egg :3 0'5 “$00 3 ... g grammes“ .— 8 geos>° 8% e stgaééééa .p 32503 58385 203 . lull. 1;»! Q“. 204. .83 03 «.HH 3 «6 .N m.«H mm 0.2. «8.... 35.3 I 1. n6 « o« .... 1. .... I .... .... «H H OH as H 3 38225 388 I .... «H o 3. .... ..1 .... .... 1. .... 1. 1.. .... H.« o 3 38205 8SE H.o H.o m6 on om 1. .... .... .... .... 1. «.m « oH m.«H R 8 .8831 2838a 0.3 0.3 0.3 3 8H H43 o« 8 man m 3 .1. .... 1. «:H «H on «5an? 3536 H.m« «6H «.2 8H ow H.o« «H 2. «.3 H on «.m « OH H.« o« 8 «BE 86.33 Hé o.« <.~ 0H 2. «.0 m on m.o« m on w.« « om I. 1.. 1.. 8.1 28930 «.0 «.0 EH 8 8 .1 .1. .... «..H H S m.HH c on $3 .3 om 3388 3&5 .1. ...... «.o H 2 .... .... ..1 .1 .... .... .1. I 1. «.0 H S 559:.» 83.835 .1. l «.o H S .... l I 1. .... .... .1 .... I 3. H S 33228 «squash 1... ...... n4. on cm ..1 11 11 ...... ...... 1... 11 11 .1. «.3 on ow .9... 260398 9m 9m «.8 3H 8H 3. « o« o.«« o 8 Eoo R oo «.3 o«H 8H 318 «baa 1. 1. «.m H 8 I I l .... .... .... «H H 2 «..H «H 3 do noHfifiHoaH «.0 H6 m6 « S 1. .... I .1 I I. m.n « 3 .1 1. .... 53E .83. 88. 8t 88. a . 88. 88 a ck. 88 9 cm. HR. a 85 fix. a 88. $593 (um: .22 >._._mzmn dug... >._._mzua dam... >._._mzmo dug... >._._m_zmn dumb. >._._mzmn .05.... m H H m « m41:sz due... E0200 3:... £0sz SE Emzmn SE £sz .30... m _ 0 a u 0.22.0» 33:0 04<.H0._. 00.7.0 mu; IIIIIIIIIIIIIIIIIIIIIIIIIIIII .11 Ill” IlhhflIIIIIIIIHIII .8“ 8308 «.2 .....H H .m :0 m ..H :3 H83 .3500 88818 5058138.“ 38830 «80383888331083.0055 -883 . I... 'lut Ill- 0 1 .5 o I. ‘- p III I :4 .1 .‘IO ... ‘I- O I‘ . 0 ll .-IIV 1": .1. oil vtvl III. 0 .la ...: . .. I c 0 941.1! .Id“ 1 ...I n o‘ .n . . all. III. .Ir‘ .I’ .I“ 9‘-.. ., a. I Ola" 207. H.00 000 0.0 00 H.0 mm ...0 00 0.00 000 0405» «.0 H.0 0.0 0 0H I I I I I I ...m m 0H I I I 3.335 330 I I H.0 H 0H I I I I I I I I l 0.0 H 0H 3820a,. uHHHa 0.0 0.0 0.00 0.8 8 I I I I I I 0.0H 0H 00 0.00 000 8 5031 02.0880 0.00 0.8 0.0 ..m 00H 0.H0 0m 00H H... m 8 I I I I I I 03038 2508.. 0.0H 0.0 0.0 ..n 8 ...NH .. 0.. 0.0 H 0H 0.H H 0H 0.0 00 0.. £02. 03026 ...Hu 0.0H 0.0H 00 8H 0.8 0H 00 0.00 0H 0.. 0.8 «H 00 0.0 00 0.. 3H... 03.83 0.0 0H H.0H ..0H 8H I I I 0.8 .. on 0.8 0H 0.. 0.0H 8H , 8H 03388 35.... I I 0.0 8 00 I I I I I I 0.H H 0H ...0 0m 00 8823:. 35x8... I I 0.0 8 0.. I I I I I I I I I 0.0 8 0.. do 308080 I I 0.0 0 0H I I I I I I I I I 0.0 0 0H «SSH... 80.80 I I 0.0 0 8 I I I I I I I I I 0.0 n 8 8.08 0.50 0.0 0.0 0.0 m. 00H 0.0 m 8 H... m on 0.9 .. 00 0.0 H0 0.. £18 «.50 I I H.0 H 0H I I I I I I I I I 0.0 H 0H .aaHnHHofio 35050 0.0 H.0 0.0 0.. 8H I I I I I I 0... 0 8 H.HH 0.. 8H .8 noflofiHsa 0.0 0.0 ...0 8 00 I I I 0.0 H 0H 0.0 0 8 0.0 0H 00 age... 080 8 «E. 8 0 8 8 0 8 8 0 8 8 0 8 8 0 8 00.080 ._._wzuo .Omm... >._._sz0 due... >._._mzmn dun... Chm—mm— .Ommh— PEmzuo Gum... 0 0 0 0 0.330... 00.0000 0.2.00. 00...... 0:0 .H 8380 a. 00 :0 .. ... .... 0 .0 3.3 33005000 .3380 05300 .NH 05.... 8.... 000.330 «a 00H 08. 8 083 3.0.. 8.3 no 0.25:0 .8 0000... ..IO O:.6 «‘1‘. II. to ..Oll ii I. ‘ .... r . v. ‘5‘ II ... .108 .0. ‘0 "I‘ 208 . 0.0m HH.. 0.0 00 0.0 00 0.0H 8H 0.0.. H00 0.5.2 I I. H.0 H 0H I I I I I I I I I 0.0 H 0H 0:82.... uHHH... 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I I I I I I I 0.0 0 8 3.858.... 080 I I 0.H 0 8 I I I I I I 0.H 0 0H 0.H .. 8 £500.. 080 8 0..... 8 0 8 8 0 8 8 0 8 8 0 8 8 0 8 00.008 (9.2 4503 >._._mzmn dun“. :32”: .31... >._._mzmn .32... :3qu .OMm... >._._mzun dun... 0 . 0 0 0 m._<._.o._. mm_uw._m mm<._o mn_m & 00...»...0» .0H 8308 +30 .... H .0 .... m. .0 :08 5005 .5080 3.008 .0H 08% 39c 3880.. «s 00H 08. go 083 38 a...» .3 .5550 .80 3000 IIIIIIIIIIIHmHflHflH" t‘ I..- ' q . .‘ o... '5'- 00‘! . ll; -.ol. " I. VI 7. I-.- .III‘ 33 ‘} section 5. 8mm of tree data based on ten 100 m2 quadrats flan-stand”, Hflladale County, Camden Twp., 1‘. 8 3., R. 4 IL, mu XIII. O Hv—IL‘QMH \OwawNN :3 gig 03Io°c5cr§c3c3<5 éér-Iéfixo'o'I 5 §~ Q ddaifid fifififlflflfl Q o If a oomoooI00I-Iuoago g l- H 0 do31g23§ e Es~wagg§33§8§§§§ §§§§go.z§§a§53§§§ 209 . Iqu- low: ...-lo -11 0‘ 1.1.] I. ‘ u‘ c 1‘ O. I x ‘ I o . IIII .II I I II I ~. ... Jr . ~ 0 I ‘I'. 'P‘ r In. .I‘ .‘1 -- cl 4 I‘ll vII O 06 .av I. I l I ...l OIIIJ . I . . I . V . . . . . . x .1 ill! 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In me‘hnbOQOM In 3 $59 o'odoaoio'b'gio'to' —l : §~. §|~m._._mzmn dun... :5qu dun... >._._mzun dumb. >._._mzun dumb. :5sz dumb. m c n « m0 0 9Q§Qiflfififi931€9§fid9Q9 3 :E\ deOROOHOOOOONOSO¢ma U) _ 2* 3 go amabgHQwHQMHmamgflggg g n. “ gee §888§888888888888888 Inl— cow N >0 0:0.- e n—8 IIIIIII3H'IIHIllm I no _ NM m z «30 Illllllmflllfllllfiflfifil 53 d “48° IIIIIIIRSIISIII8S 8| 8‘. m 0 mo 00 0 mm H >- e ee .0 e 0. e 1 tag IIIIHIHHIHmclllgllgH . m P z 0.,“0‘0 IIIIHIHHIHMIIIIRIIS” % I'- d m #5" IIIISISSISSIIIIRIIRS Innh In be H H H H H p < >° cc. 0 e e e e e e 4 EE\ 30HI3IHIIHIIIII®IHIH n 0 2 J80 8°H|81HIIHIIIII8IHIH 8 Inlnnl N fig SSSIRISIISIIIIISRISIS Inl- m H“ “N9...§ . .923 993 9 t§3 ddN NHgo oooIoIIom NIMHR g E z m mm OHNH N m8 com o «an H H“ a I II mflIN 3 a d 315$ §888§383I3I|8888|838 ll- 0 en 3 .gag 88 a 5 'H v'l 85 8838 §°§ E 8. a fi' ‘4 n Od-on m E .§°.85385§8 8838888 4 g g°°§g a sfizu :gusaag 5 “’ Egg Iggg’fissflaaaa *‘ annéfiaififiaagggwn §§oossmafiammm 58838 fll -.-. 19'- V... 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H.00 0H 00 0.HH 0 om 0.H HH 0H. 318 93.0 I I 0.0 m 0H I I I I I I 0.0 H 0H H.0 m 0H E3530... asfifiuo m.0 H.0 méa H.HH 00 I I I I I I .w.HH 0 on 0.0.0. mMH om do 8300305 H.H 0.0 0.0 mm 00 I I I H.0 H 0H 0.0 m 8 0.0 mm 00 55.... .83 0.0 «H... 0A. 0 HR 00 0 § 0m. 0 «A. on. 0 on. .x. 0 HR. mmBEm <52 .293 E0200 .05.... >5sz .005: r5200 Sm... >550 SE :5sz due... 0 H H 0 N 041.0... 050:0 .r 0.2.7:: mm<._u 0:0 I .0H 838.. a. :0 « 32 ..z m .m ..z 0 .0 SE .830 .5550 03:30 .0H 05% g 30.630 N... 00H 03 8 083 8.3 can no 5550 .HH mafia ata frcn stand 21 1' section 3. Smary of tree data based on ten 100 m2 quadr St. Joseph County, Iockport Tap” T. 6 8., R. 11 $1., NE TABLE m. : . H bflflmoqaedwi to 0:5 oIc~I<5c30°u§m~ooooo I- z . {3 §~. H addadiawdQI d .2 I: o Nooomgboooo 8 =1.. 3 E5~ HHdHH5§H040mm U) m "" Z a 3 a mooaomgo‘te 515$ g I. ‘” fig sggsssgggassg ll. xowo M 1‘ I10" I II I I ' g INI I013: 01 no I 30 IIHIIIIPI'SWIHI fig IISIIII§88ISI II. "’ an «N: o m L; 1E0 tummmlqmli‘; .. m z :30 new I”‘&‘I"‘I"I“‘ 9 P o .. g0 Ileslsalazans II. In o~¢~ma~mc~~o 05‘? O I: E03 HAOH6£§III$£§ d m III N 35° sassasalulaaa " E U) account: coco bCRI-I \q EZ§° NH? #HIdOK‘I’Ibfl' 2 39a ‘3 Id 3:: Haflmlamfilsng g g 88888I888l888 I I5 I a a a ...-Ira I; gigsaasfiwgg g a: «:me ..«aaaaééé. 385 §§§°§3335 44030 a {DD 217. ‘. .00 ‘1 r..- 0.2. mom 0.» .3 9m .8 0.3 no a.mb m3 335... «.0 H.0 wd m m I I I mé H OH 0.H H 0H 0.0 n S .53qu 3&5 I I m6 mm cm I I I I I I I I I n3H mm ow E631 39.33...” «.0 ca p6 .H on mam .\ on I I I I I I I I I 2333 08.8.5 H.HH bén a.mH mm 03 0.3 H om M3 H OH “.0 o 8 H.HH Hm 02 8a.." 32.3 3% 5.8 H.0H 8 8H H.Hm .2 om I I I H.HH b on .3 on ow 3H. 22.3 I I 0.« NH ON I I I I I I wé m om m...” m on 333." mg m.o 0.0 Nam «m om I I I 0.0H H on 0.H. m 3 «.« mm om 3338 3&5 I I «.0 N 0.H I I I I I I I I I «.0 N 3 aqflfimflb macaw—”8:. .... I ad 8 om I I I I I I 0.H H OH m.~H R 2. .5388 Sfifié I I «A b on I I I I I I I I I men 5 on .3» «$333 H.0 H.0 «3 Hm om I I I I I I .2. o 3 in mu 3 .383 3&8 H.HH 0.0 «3 SH SH m.mH o 8. 0.8 .HH om Ham 2 8 o.m~ 8H 8H 31:5 ago «.0 H.0 fin Hm 2. I I I I I I S. m on «6 on 2. an uoflofiHoaq n6 «.0 H.H.H 8 02 I I I as H OH a.mH 0H 8 0.0H 3 8 gen .83 .5 «..E fix. a fix. on» 9 “RV on. a on. “N. a “No an. a fix. $595 $5 .. c .Q Q QQdQQ Q h5~ IIIIOIIIIHIIHHoMHIII m B H «n F Inga IIIIOIIIIHII'IflmRISIII R d g5° IIIISIIIISIIQSRESEIII m 3 g Q i 199 '9' qwllwl t < gEx gIIISIqMMI m Imfi N m P z 0.3o ”III“I*"”II”II““IIHI a 1.. d m lg§ RIIIRISRRIISIIRRIISI IL. m own wow w m H jag muoI' 'émwlmwmai :1 Q 2 43a 3““!“NSIMIFII“IHI*HI g u N 3g ERSIRRSIRIRIIRISIRSI ... E H m “mono Mm Mb 0 < 50% n O .... .... I ... ... . EE~ bNbOH I30 HNIIOOOIQHO g m 2 «an nasang SQIIRN“I82“ g d E58 88888|§° §° RII§88IBRS é a «II M «s dE dads“ gg saga a no.) 3.335333% +v “3900 3 a kafiss aswoaa3sufiza < 2 gsfis‘“u§ g€55313>a°° s 0’ asoogguwgmdug aaagga I- adnupugdbu 3 a find a n 9% uahmH 83§5550§ °350§°°°3dé <4 oooohéé‘omm 5:95:09 8% .III ‘I. I. a' I! . I C n»- II VI ‘I II a n .0. V-.- I’.’ v.1..- p-..‘ ..L‘ II :6 I... 220 . a.mo mmq ~.m “H m.> mm H.mH ow o.ne OHm mn._._mzun .Oumu >.:mzun dumb. >._._mzmn dumb. >._._mzmn dunk >._._mzmo .Ommm m H n . H mn._._mzmn dum... dum... >._._mzmn :5qu .35. >.:mzmn v m N m._<._.o._. mm.0umm m._<._.o._. mm<._u uN_m .3 Eu»... :5 Bahama. ms 03 n8 8 H598. 38 8.3 .Ho 355m .39 Ham: .NN 8:03 H.HH ..m m .m ..z n .a :99 H.Hgoo .388 nofimnHSH 21 w., SIM swi section 2. Gunnery of tree data based on ten 100 n2 quadrats Iran stand 26, Berrien County, New Buffalo Twp., T. 8 8., R. TABLE XXVI. 5 ° SaIII" ”I I”°°" °' 2 §& ~0HH <5 Io 5%:OOI < _l I— gu 030;. |I>ION H {'0 .stmoo' q o E: OOIIIo'HHIoI IOIOMNOOI 8 '- rL bstmmwomwoomxtmmwbmr-Ixo w t& «zacmasssaawowaqaaa In a N H H H '- Z 3 3a mg°w~xflsmmsawaa°°n~ § 1:. "’ §e\° R888882§888888§8882 IL ‘00! b b Ek" IIIIIoSIIIIH "cSII-II «3 g H mrga Illll""‘IIIIII%‘8‘°I"‘I SR 6 g5° IIIIIRSIIIIII888ISI 3 o ifilll'I'Illqlii'fill * < §®~ 3* 3 a b 22'“ m |_ Z 0.3a m*HI~I~IIrw~~IHII a P I O m fig 83III8I8|IISI88ISII III. ‘2 0.9.. . . ...»:“I 020.1: ~t tk" mmHNImIQ'Noqr-IHNINIAO o _l a N M m u 2 ”an balemiginflgNNMIQL‘F-I E‘ III! N see RSSSISI8|° 2883°|° 02 g E >. o 0.... 0'1 OQN 0“. “i 1.5 20%!“ HNIbNIo mdQININNO 3 m 2 0' fig 888888I88I 88ml3I882 III. gm 6 CI .5“ :33 “5 a35 g.°25§ I§2§§ m g gizaaawgagéggam a I“ O h III-I O O a ggéasgs~§asgwa§a§§§ h Sfiaadg abaa an II-IIIIIIIIIEIIIIEE adsuuoossmas ééamap 222. I- a I-.- \l a d I. \ In.- ..‘6 II I I ...: O I In \- qOOII ... I 0‘.. QO.II t. I . .i . S. . u'o. II)! I- .9. . I . .‘ . . I I . .Ivn \ .. II a II I I I II I. I- I J‘I.‘ ‘.!a Idiot Int bflafimaumjfl qua dtnohubuwanunmmfi ., T. 8 II., a. 3 II., mi SIN auction 10. Sung flmmnme,hupr TBHIHH. hmuaunma Pyrus ioensia Uhmamflum MmsNMa mumadh 0mmmbmdw muwamma Tflnmmfimm hmmafim Hmuam AmrmMm Analanchier hangs». mnawdh Cancun :5 c: QDNOQOQOIAJNNRGNMHGN c: uaaxc>u3 0000000000 o 0.0.. m ¢S\ §0Hm0000NoIHIa0NbIII A < m < J ~ 3~. QQRfifififlde * waH q o 3:: $00NOOOOHOIOI$OSMII| g h m o 9%??9139d§QQ1Qfl1Yfl§fi m E5\ gNomowHHoooqomomoomo _ g N M g; uI m can ggbsmgggHoNRHgHjanH 5 m I m . fig agsasggasaaosaa 0239. m b In In H Inxo so Es? §IIéIIIIéIIII5|§§III & m z Inga “II”IIII"’IIII"I3‘“III SI 6 g‘c\° SIISIIIISIIIIRISRIII L ”r q %9 9 9 9 $99 9 :Eg SIIngNIINImlgwIIII ,I m’ P z 0‘30 3|I°~IHII”I“I“H"‘IIII 3 p... d “a fig 3| °3I3II3|8I838IIII III-I m QNOObHQH «5% b ‘ >0 00000000 . 4 EE\ 35G”NQO¢IMINIZIIIHNH :” u 2 go anboNM3MIHINIni'IHNH g fig 83333838I3I8I8III333 m 19 “H Edfl91 dwede 0 H % IE5~ baInoomH oomONIHIIHI I3 m 2 3o« §I8”$§SIHNR”EIQIIRI 8 I H L‘ S e 00 E5 8§Ias§§° ISHmSRISIIRI 3 35 a I :33 .5: g} 0-5 a 5 f3 - S a H < 8 Eu, .2 +- n- up Io '0 3 id 3 I" I 5% .. o 39.. § 223. I a. u u .— . II 0 lltf a... ‘I .o I"l I to . |§ I n I; -.. 0| lit-l II- .I. o .I V '40 1 .I II IV» '0‘. I I .31 .l a nil I ..l a(. n. l n o v 0.. ...0.- . . O O o 0 I n . I Inc I.I- - I ¢ I :- O o . . I.‘ I..- V324 1!. ..‘l .IJ I... 1.". C . I I"u I‘O. ‘ ~ . t. _ v in 0. ll ‘ I'll. I 1’ '71. b ‘I I..!O {lull \p u I . nl'. .0 Worm-co databuodon 13111001112 quadrats tron stand 28, Jackson County, Watcrloo Np" 'r. 2 s., a. 2 2., w i section 12. TABLE MIL 3 2&8 EI§E§I§§§§|§: : §~ % dfld finaq Q i 0 Li mrowo‘oogglo' 5' I- m p o i19fi9§1§§d9€1_ E "11';& RfliflfioJfibNSQOr-IH 3 E9 Rmfifisflfggggmwm g a. m 0‘ o Ee\ §28882§88§828 E69 3|!§llll§§g:g 3 U) 2 3n "ll“lill°‘:§|ll a gas allanllggnt 3 c a 9. 99 i 9 1 < %5 R'ImilmmISIml # Z 23;. ‘°|l'*ll""‘l°°|"'l as F o a, fig answsmszm (D 3 Eg 33§§§r§313131~ E U) 0 En Sambo'mq'Q'¢' a u 0 ~ 3g SERRSISSISISI ; E E59 Rflfiflfiomoflfio’gé :3 9 . g a gQ’QSfiflfigfiz-[Nnm § gig gseassgaasasa Q Eaééggzfiagg 3 a g 012' 8133"" =- $ Eaagfig "uagia 2 §“§§E§ §§“§§g «£3903: £30) 22A. 1| .tl 71.! 0“. «1‘. a O i ‘ - c I... O I- n I u . . . I C A :o v.) -1 I, ... I'D .II-uh I'd-l. '.tu g...‘ Lennon County, Adrian Twp., '1'. 6 8., R. 3 12., NW % section 12. TABLE XXIX. Emery of tree data based on ten 100 m2 quadrats fran stand 29, a bbb wa 225’ mgg w6'l‘:o~l666'g§~tl6lll _l .J : g N O 0 O O b- l O 0 0‘0. 0 o. :3: wol'lowlooo'gngo'l' a >_ o OOflniQiqquqflzo<~t~10fi¢q u": : E>\ GHHmO~fMOfiQHON~fCOHOOQ U) "' Z a g a a¢b$<§H5a§>Nfib<~tnN3°g°9 s u. 3: 5105335 “333%5 aa ,_ dud d ggéfisszg ééggfisaézgg 00000;:0 55:08?! ’h :0 cc! 0' .30.. I.“ .l' 'i . O I to". OI]... I... O .. It. 0 r . ..l o ‘I. O O 0 $0 \ O '4... u of. .‘U . no 0 s O 9 I ll I v a . O I 0 . O ’1]. .-ta .O‘.‘ l‘ I "‘ "J 0...." '0’. t.(u / _. . 1 u . '5‘ TI‘ .0 u] 1 U _ .P O 9"- VI». III'O I'D- C.“ '7‘ ' I ..U I... 1 0" .011. I..- o-.. O I‘v... I.... III. Out 30 tcolvo D.l‘ I It] I." . n .. o I... ‘l 1.. Summary of tree data baaad an tan 100 m2 quadrats from stand 30, St. Clair Cmmty, Clyde Tup., T. 7 11., R. 16 3., SW 1» section 4. TABLE III. {lo 9"!"3'1": “2‘33: ‘3 mmkHolmil Hoglflo: 4 < <—l p 5. H¢ b #m qum m o < . on... a 00.. lo a 0 at Noglozlooo 46.-oil g l- o mm in gNQmmqqomamooqg :3 §& 60 Mg} OONoovrooONbMNOH - z 3 g‘: sgaw flfimmmm Ngg 88 "1."1“2°‘2“‘. 8: 0: ::&° walllIIININ§$~7H<| o m P z 0.3:: ”Sliilllfllfl‘ofimllwll 3 I- d mg§881l|l||29h°88|i8l| II. 2 8888 88 888888 8 . 8 4 EB? aw “g I'OO'DOOO 000 '0': 3 m U 2 ”so 8g 88::HHI8HHHHHIHI: g u N §§o 8888:1881888 °8|8I| n w o qqnmebNooNobNmm¢Nm d ES\ @3380No6bg6Hoomoooo s m 2 N30 EEgSHgMHgSHbmHSQQHN a d jg8° 3§88383388388388833 m :3" 8% s» . E H mea &”o .°8883 ozgag Q ofig Eagafifififiiflgfio fl 8 g 00 g g ganja 9688 g o. ogngga tub-pg sgfigfi o m a: 50 050ng gas-I 0 I" 8835883 8838838888 88”§§§2§£$5§§§38§§§ <450000h om wasps 226 . 227 . 0.8 H2. Ham 9 Ham 8 a.“ 3 8.8 one 352 I I H.0 H OH I I I I I I «8 H S I I I 33225 382. I I H.o H 2 I I I I I I I I I «.0 H 8 8638. 3.8.8an m8 m.H m6 N 8 0.«. m 8 I I I I I I I I I «83H: 8685 8.8 0.3 88 S 8 «.8. mm 8 +H.HN o 8 I I I 0.0 8 8 aka 8.286 8.0 m5 H.0 H S «zm H OH I I I I I I I I I 8.1 86.85 I I o.N «N 8 I I I I I I I I I 3” mm 8 88388 388m 0.« «.« H8 HwH 03 I I I “.8 w 8 8:8 8 8 «.8 8H 8H 335?? 858 m8 m.H 0.8 8H 8H m.~ H S I I I o8 HH 8 8.8 8H SH 3828. 85828 «.0 N6 88 H8 8 I I I 0.« H 8 «6 m 8 H.«H t. 8 326.8 38.88 H.«H 2. 8H .HH 8 flw.HH m 8 H.«M m on I I I I I I n3:5 988 «.0 H.0 “8 8H 8 I I I 0.« H 2 8.0 m 8 «.« mH 8 .8 8308888 0.H o6 +H.H,o. HoH 8 m.~ H OH I I I I I I H8 8H 8 85288 88H H.m 8H H.H Hm 8 n.~ H S 88 m 8 88 ~ 8 8m 8 8 SEE .83 05 «.._l._ ORV 0 ok» Hg D $5 05 D g g 0 0§ g a g MEUmn—m as: ._523 .SE $823 8:... n c m _ N 35:: mus“; m._5an SE Emzmn SE Emzua due... . a H w m u 5.2:: 2:33 51:59, 33.: mu; .fi 935.08 $nz-..n «Jim m. .5 SE 823»: 5850 8385. .mm 3.3» 8.6 3.83% «a 03 a3 8 393 33 8.3 .8 g .38" Ema. E Q OOH HHH ooHoom m §S\ 6H6Ibm6IHHN3mwlt 4 < i p mu bNm mom ONwHoN H O at: 6H6ImN6IMHHmonI g '— ° HwONonNbbmcHHHo 3 EE\ oqNOOONHHmHOHNmm m . ' z QHHM bow no a 3 3:: 33 3 3§§~ “mgg § 0 m H g m d o 0000 000 o afiw EE§ Sngbmgbgggggng 5n 0 ~O \DH In fig .25 MIIEHIMI} 35.53:: .: 90 W an 2 34. "ago I”I|‘°“‘||°‘I°‘°SI‘°II R a 6 "am? 55° lanaallalassan _& g7 2 t . finegfilfl,%wggfifll' Q an N w w H NH H ‘ P a: "a. MMMNW‘HMH a “’ I- nc , 5“ °o : m §B\ NISISIHSIINHII SE m "' HHw H '33 j :5 .zagnfslanammu .2 2.5-I u 2 £3 "'30 I"""°I”I"‘II‘.-‘3"'|I“'Il g u . E" g.) °8I3|3||83||8|l 0 mph 8 Q HnmNMMMNHqH ooHN q .g EB\ ggHOOONflmmH 6Hmm a H m H a a a ammwmmga Imam “‘ a}: NO Hr-I M 3 m S 6’9 oo o oo o o m 3 EE‘ §§>H8H853§wI88w8 5; § géé 3.255 .§§ fl g uaflgggg §33H3u 2 3: Egg 1§Hu Emiflsfis 5 m 33538 “a :" €353 P W afiuugfi p a mum HHH a nnH 88”§§5o§ t ggufifi 44500000 0 amp 2w. ,.. w . . .‘ll. ...0 ‘0 3‘. I. .. it I ..0o ..1.‘ ttvfil it... I‘ll! Infl| I IV!- ..| ;‘p . l-‘O _ A-uil..-. I t, ‘II’. It. .1 draft! I?“ am 3'4, oflhnfl. $3 nmsnnm wmuydtnehubuuanunmmfl Ionic County, Boston Twp., T. 6 F., Rm 8‘U., NU 5 0M HNHHHwNoNomw m §§3 NHI'66666N66éfiHH _l < _I h §~. an widwnnnendnfi 9 g t QN ' IOOOOO~¢OH§3NMI g u, I o dfi‘Q‘YiiQQQ‘IfiQfi‘fi‘i‘Y" E 6S~ mgooooqbQONHHMOHo 3 En: £§MH~~na$~anaa~°~ a h. ” Egg sgasaaaaasgaaaass gee Z:II§II::3I3§§33 § 2 J an ”IIIHIIIINI“HS“”I a 1 . 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"“IINNHIIMH*II":II°IH a P o §§2 aa:n2&2:uasat:snltals m u. m momoo c «19000 N o a: ' a: : tfi damma‘éa4alw'a‘l'lg'a 22‘ a . Q 2 3a “fi”°”l3”““l“l”lfill3¥: 3 III N age ”3 :32 RSIRISIIIIRII — a: ll. U) 55° mSOOHomu-IHOOROQ‘H’NONM 3 2 3c: saNamHamgwaamhaaa H "aaH.~:Haa~g aaHmm gavsaséfiéfiggg “=saagé HHsoHossHa 3H éaagHHH 231 . .Io.l \00.‘ ‘0'. -.v‘ ‘0 | "’- a... O "l. -‘\O ... ‘5 .1. 1' d .01... ‘(HIO II‘. in: a . ..cID h H. u. .0 II! 0...... a. I 5 9n) . I.-. . n . all: . His: 5.. J. ’t u .1. u . a H {‘01. E Q m: «} section 22. TABLE mvx. Mary of tree data based on ten 100 m2 quadrats frail stand 36, Benin County, Chi-wing Mo, To 7 80, R. 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II 0.« II II II oHoMHQHHsv abuvqquHAHH II II II o.N II II II aann uquwpu mm o.m II m.~m a.mm Q m unwOHuoaw usuaxunh mm w.m II II II II II «HHouchaum msmum II II 93 0. «.HH H H «2.31 888 mm w.~H ~.mH H3 II II II 318 «baa II II N.MH Neq II II II mHHd>b whudo Hm o.m .HéH m6 II II II 3an8 ago II II II 0.« II II II .mm noHnonaHoa< Mme w.bm b.>h m. 81> 81> 81> 81> 28m .8120 monwpuomaH occupHOQEH moQQOHmHQmHm nun 9mm .02 .oz @811: m can m wawaHo m mmmHo N mdeo H uuumo .8 88» no 889 .8“ 8188 801.: 835m .35 Ema. ........... 254. «QHH ous>_chQH ezdanaoo .3 H.m name. mtg. w.MN N N and." nan—Hp mm 0.« 0.HH o.qH II II II anuOHnoaa usaHp NH m.m N.bH <.¢H II II II auvaHd mahudnuam Ne N.HN II II 0.0H N H uansHob muonosc wwm m.oo II n.0H 0.00 b m dandy msouoso mom 0.8 II Ham H60 0 H 81 -1.8.80 mmH H.«m >.mHH m. m.HH H H uanouco mansnm mH m.< «.mH II II II II auabu msnshm II II 3.0 H.0 II II II 82on 8&8 Hm o.m m.o N.HH II II II mHHwaqouHooo meH00 02 imm 0.0m «.mm 0.0m m 0 2.18 950 II II o.MH m.©H II II II asnmnoomm awed II II m.HH b.w II II II Hannah hood 81> 81> 81> 81> 81> 25% .880 ooquaomaH ooquHOQSH monaOHanme Aha and .02 .02 081.13 n 08 110.000.] 8 m 9.18 N 188 H .1040 E .1 H88 .1 88» you 011080 8015 83880 .HHHE Ema 255. mmeH onHabvxmvaH asanpqoo Nm 0.« H.0 0.H o.m N N wnwOHhmaw usaflp II II II 0.00 «.HH m m 8031 18.1180 mq «.HN II II II II II ananHm> 0209090 mbw b.m0H H.«m ¢.¢N a.¢m HN b «Hash msunono oNq m.mw m.bN a.mH b.¢ H H man manhoso II II o.MH H.# II II II mquOOH unhhm mm o.mH b.bN b.0H N.mN m d qupOHom usnshm II II II b.H II II II mnanHmnHP ushomanh II II H.0 N.HN m.mN o m wqalooea manxdnm ..- II .... m.“ 0.3 . .H m_ .0. 28320 II II m.mN m.OH 0.HN 0H N ddHAOHH nuance qu m.#o 0.0b o.mm ¢.mH o m mHHd>o whuwo II II H.0N H.mH II II II .a» hoHaoanoaq cm 0.« 0.0% 0.HN 0.0 N N sunfish Hood oanb ous> osHub mSHw> osHmb mempm .mvwso condpnoaaH oonapuoasH ooquHMHanm Aha umn .oz .02 doustmz w .008 w mmmmmHo m mdeo N mdeo . H mumHo .NN wanna Mo @0099 now voHHaauo movaunH nOHewesnm .NHA mumda IIIIIIIII 256. mde o§Hu>VNownH auanpnoo II. II II 0.H moo H H “GNOHHQEG “HEN—”D 0N N.m b.m 0.0 II II II sawOHnmaw aHHHH II II m.mH H.mH m.@ H H aschHu mwhmwmmwm oNH §.qo II II II II II wannHo> anonmsa boo o.NOH b.m m.N m.@ H H «mash muonoso oHH H.mm II 0.0 II II II 10H“ 8880 HH m.m Q.MH H.HN w.bN m H quponom nsq2hm m o.N II II II II II upmpnocfiuadnw mqumom Nm m.o II II II II II manonpm manm II II 0.HN b.w II II II mannHmnH>.whupmo mm w.v II m.N II II II dOHpu>Hhm wmmhz mm «.0 m.NH m.oN H.Hm w d nQQOHHoaw manqum no m.© II m.< II II II «HHoMquwHw mswwm mm w.NH o.mNH a.mm m.o H H anHAOHM manhoo II II o.w II II II II wwwpo whhmo mHH n.0m «.HH 0.« 0.0H N N mH13 «huge II II II o.N .020 H H 383.3698 050 II II 0.0H 0.0H «.mH m m .0m umHgoachEH II II b.NH H.b II II II ashanoowm Amo< HwH N. ousb msHmb 05Hm> ous> msmpm .mcwsa oonmPHOQsH ooanHOQEH oonuoHMHanw Aha hmn .oz .02 copanmz u on 0 m 81Ho 0 umaHo .HfluumHm .MN 00590 Mo mmohp you uoHHmaQo moofian QOHpaaasm .HH mqmde 257. onH ous>_chnH SSSQHpnoo II II mod Mom II II II dfidOHHme OSEHD II II woNH How II IW ll Efifldpfld mdhfidmmflm 0N N.MH II II II II II manuHmb manhood wmw «.02 II 0.0 o.o~H .HH m «.38 8880 000 n.00 II m.H II II II 13H» 8880 II II 0.HN H.HN 0.0« N m ungonom asuspm II II m.mo «.mm II II II dddHQHmnH>Idhhvmo 3 h. a CH m o 0: .H o 50 o 0 on Q m figfiHOafl mdwfixdhh 5% b.# II II II II II dHHOhHundAm mnMNm II II II 0.0 II II II .mm unwoupuho II II m.< m.bH II II II deAOHM nuance NR H.HH. H60 H.0 II II II 318 810 II I...- 0 . .HH 0 o m H II II .II Hmfiguqmflmfiq II II II m.¢ II II II enhanced» 9004 II II m.< H.m II II II sunfish 9004 05Hm> 05Hw> mSHw> madmb ost> mswvm .mvaso megaphomsH monprOQEH oonwOHMquHm aha Aha .oz .02 08.112 quqauqluumuuflmllll m mmeo N mmeo Inna mmmHu :‘N .0595 M0 away. «HON ”0:980 @0039“ gfifidgm .HNH flan. 258. mme 0HHHHS xownH awanpnoo II II II m.m o.m H H manna msaHD II II II H.«H o.m H H unonHmau mfiEHD II II II 0.H II II II unwOHhmew wHHHa 0H m.o II 0.0N II II II auanHu mmhudmmum 8H 0.8 II II Ham 0 H 83:1: 8280 COOH a.mmH II 0.« b.NH m H «Hash usunmsa No m.wH II II II II II dan nachosa bHH m.mm II 0.0N 0.0% mm m «Gauchom manfihm II II II m.m II II II 25H>w manshm II II II m.m II II II :qaoHHmsu magnum II II II 0.H II II II ananHmqu uhupmo II II H.Nw H.Hm «.mN 0H 5 QGwOHHme nsndxdnh II II II 0.H II II II .am mswocvono II II II 0.H II II II upubo dunno I II II in m.m N H 03.26 «So II II 0.30 mdN «.mH NH 0. .8 820818 II II II II II II II abnmnooum hood NmN ©.mm II 0.00 o.#w MMH OH Hannah awed 81> 81: 81: 81: 81: 8.3 .880 monprOQEH moquHOQaH oOQdOHmHanm 9mm ugh .oz .02 08:38 J3 m m8Ho N 18H: 140.3% .mN camp» no amen» pom voHHmeoo mmochH nOprsazm .HHMu MHmHB 259. NmmH 81: «85. 8:888 II. II ®oN QoN II II I... dfldOHhOSG @555 0e 0.« m.bH 0.« 0.0H N N unwOHhoam “HHHB 2 .3 2H 0.0 II II II 83:. 88.8 #0 0.Hm II II II II II mnHHSHo> 0509050 mph 0.mHH N.0 0.« 0.0H N N «Hank «anyone 00¢ N.Hw 0.H II II II II «9H6 machoso II II «.0 0.0H 0.0« m H quponom magnum mm Q. 0.0% 0.«H w.0H N N andHaHth>_ahhpm0 II II m.m 0.0« 0.mm b m aquHhmaw manNuhh II II N.0H II II II II wHHOMandum mnmdh II II III 0.m II II II .00 mswccvcho 0% 0.0 H.ww N.HN II II II acHHOHM manhoo om ©.m II II II II II upwbb ohhdo 8 0.8 0. HR 0.0H N N 8.18 988 II II II N.m II II II mHanouHcHoo mhhwo II II 0.« m.0 II II II wannHHono unnHahdo II II 8..» 0.0H II II II .8 880818 mq m.< H.No N.N< 0.0H N N anaonowm 9004 00 0.0 0.0H 0.0 0.0H N N sunfish 9004 05Hw> osHm> 05Hw> osz> oSHm> namvm .muwsa megaphoaaH oomwpnomsH wocmoHMHanm awn nan .oz .02 08883 n 0mg m wowwuHo m mmeo N mmeo H uuflH0 .oN undpm no momhv you GOHHQEUO QGOHUQH QOHpmsafim .HHHNH mumda 260. mooH 05H0>.N000H esanpnoo II II 0.H II II II II 09nn9 maaHD II II m.@ H.HH MoNH N N wHHwOHhmfiw mfifiHD II II 0.m II II II II 0n00H9080 0HHHB mH H.NN II II N.o H H 0ansH0> 0509050 0mm 0.00 II m.b H.mm w H 09909 0009050 mN 09H II II II II II 90H00Hp 0009000 0HN 0.0H N.NH 0.0 0.0H m N 0QH0 0009000 II II II m.H II II II 0Hun00H 009hm II II II 0.H II II II . .00 mans9m mm H.0H m.HH 0.0H m.NH N N 0qHPO900 00:09m 0H 0.H H.NH- m.H N.o H H 25H>0 msqs9m HH m.b II II II II II 0909000ch09w mstaom II II m.NH 0N II II II 08H§8H> 88.8 0N 0.H n.0m H.mm 0.0N m H 0000H9020 manN09h II II N.NH H.HN II II II .00 1:000:080 NH m.o 0.0 m.H II II II 090>0 0h900 ONH 0.0H H.0N 0.0H 0.0H m N mHH0:o 09:00 II II H.mN II 0.0 N H 0§H5H0H00 055.0900 II II H.0H N.0m a.mN 0 m .00 :0HgoquHoaq NHm N.Nb 0.Hw H.0N 0.Hm 0 m a59ps9 9004 05H0> 05H0> 05H0> 09H0> 05H0> 09000 .00000 88888 88888 00:8H9H8H0 :8 :8 .oz .8 8801: II n _08I_H._mo_mm_0H._o.. m 880 N 088 H 088 .bN 00000 90 0009p 90% 00HH0500 000H00H 00Hp0aasm .>HNH mqmda 261. 0SH0> x0qu aubchnoo 0HOH II II II N.m II II II 0000H90a0 azaHD 0H m.m m.NH 0.N II II II 0000H9050 0HHH0 II II II N .H II II II 56.5.? 00980000 mm0 m.mHH H.mH 0.0H 0.NHH HH OH 09959 0009000 Nmm 0.00 II 0.N II II II 0QH0 0009000 0H H.m H.0N 0.mN H.0H m N 00Hpo900 00::9m Hm N.m 0.0N 0.Nm 0.0 H H 0:00H9080 mnuHN09h II II II H.N N.NH m H .00 0:000:090 II II wobN bofiN II II II GGHHOHM QDQHOO 00H H.0H H.mN N.0N N.0H N H ..-H18 898 II II «.mN 0.0H II II II .00 90H0000H00¢ II II 0.m N.m II II II 3090:0000 9004 0MN 0.Hm 0.0mH H.0H 0.0 H H 559909 9004 00H00 00H0> 00H0> 05H00 00H00. 05000 .00000 0on0p9oasH 0000v9omaH 00:00HMHaMHm 900 9mm .02 .02 00803: n 000 H m00m0H0 0 000Ho N 000H0 H wmmHo .wN 09900.0 no 0009p 90.“ 003980 000.205 00Hp0§m .33 mama. 262. Hm0H 00H0>.N000H eaqupcoo II II 0.HH H.NH 0.0 H H 09909 0850 II II II H.H II II II 0000H9020 0:5H0 II II II 0.H II II II 0H0q000000 00:00 H0 0.0 N.0 H.0 II II II 0000H9000 0HHHa II II II H.m II II II 0:0H9H0 009000000 mm m.©.n II H.H II II II dfipfiflwb 9.59650 m8 H.00H II II 0.HH 0 H 889 8880 HON H.wm II m.H II II II 0nH0 0009000 II II II 0.H II II II 0H0000H 00900 0H m.H II H.0 m.mm m H 00H»0900 000090 on 0.0 0.00 H.mN 0.00 0 m 088.909? 8980 On m.w 0.0 0.HN II II II 0:00H9050 000Hx090 II II II 0.H II II II .00 00M009090 0H 0.0 n.0HH N.HH 0.0H m N 00H90Hu 000900 HwH 0.0N o.NH 0.0 0.0H N H 3:8 8.8 N0 H.HN 0.0H 0.2 0.0 H H 81.5 8.10 II II II H.m II II II 359003900 00900 II II II 0.HH II II II 000H0HH0900 000H0900 II II II N.0 II II II .00 90Hnoq0H054 0m 0.m w.mH H.N II II II 009000000 9004 HON 0.00 H.mm m.NH m.mH 0H m 009009 9004 03.00 03.00 03.00 030.0 00HI0> 0 Emvm . 00000 0000p9o0aH 00:0»900BH 00000HMH0MH0 900 900 .02 .oz 0000mH03 44.08.0041 0 088 N 088 H8080 .0N 00000 mo 0009p 90% 00HH0000 000H00H 00Hp0sabm .H0Mu mqm4a 263. 005 0090» 0800: 5003000 II II II 0.H II II II 09909 000HD II II II 0.H II II II 0000H9000 00EHD Hm H.0 o.m N.m II II II 0900000000 0w009 .... 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I I «538% flag I I I NJ I I .... 53p? mmfimfim .N 0.: I I I I I 3333 2538 NS 3.3 I mam m.$ ma 5. 88a mango NN .3 I I Q: m N 21 magma I I I «.3 m.NN N N 3323 83 SN don Nam: do.» N.NH m N «fiafimhp «bane H.N. m6 N.$ 9.3 9% 3 o 2822a Bfifié NN m3 N.NN H.NN I I I «3qu 3880 3H 9% I I m. m N 318 ago NH 5% 1a N.NH I I I .% 332384 on 0.m II ¢.wm N.0 H H abhwnoodm Hood RH N.NH mJN N.NH m6.” e N Eng .82 81> 03.3 2.3» 2.3? 8?» 33m .380 oonmphomaH monprOQEH oonwoNMHamHm ppm has .02 .02 @3532." m 2% N .3330 m 268 N «38 Ijqflo :8 ~53» Mo amok... no.“ voflwmgo moowwn.“ «sauna—pm .HHHE Ema. 265. wobH ode> xmwcH asqupaoo II II II m.q II II II andoHAmaa msaHp II II II b.m II II II auchHm magnummam 0H ~.m II II II II II uanaHmh usohoso onH «.mbH m.m b.0H a.mm m< 0H candy manhono om o.mH o.NH H.m m.< H H wnHu msvamzo II II o.mm m.<~ 0.0« m m unaponom magnum II II II m.H< m.mH m w unwOHnmau manMunm mm 0.« H.©o o.Nm m.mm MH m «UHAOHM mannoo m3” QS 9% 93 EON o e 315 938 I I mAN «.2 I I I .% $305035 mmm b.¢¢ b.mb b.mm a.mH m H Banana Hood dew> 05Hw> osz> mSHmb onHa>. msmpm .mwado ooqapuomaH ooquhogaH ooqaoHanmHm nan Ann .02 .oz uflnmfimz I .I.::njmqqulwmmmwwmllll m mmeo N mmcHo Almanac .Nm vamp» mo amok» you coHHmauo nonfiunH mafipwaasm .NHNH mumds 266. NHwH 05Hw> xmch asqupsoo II II II «.0 o.m 4 H nanny nuaHD II II II b.MH m.HH o m unwownoaw ansHp mbN 0.0m N.bN w.b «.0N on b Newcahmaa wwHwa «Hq o.mo II w.# o.NH o m waysh unonmso mom o.Nb II II II II II aan usunmso mm 0.0H Q.NH b.m «.NH w m anflponom msnshm m3 oéN can: «Ion N.0H N.N q afififigp “bane <0 o.mH II w.m¢ m.vv hm 0H anGOHnoau maqfinaym I I I «d mé N N .3 $0330 on 0.5 H.0H m.w 0.HH 0H Q aoHAOHm asunoo mm N.NH I I: N.N H a 3.28 «has 03 «.mm m. N.N N.N H H 318 N630 II II II m.H II II II mquQHHonwo uznwmmwo mm m.@ «.mq N.N m.oa Ha m .a» umfiaoquaoaq HMH H.MH 0.0 b.NN m.v N N asnmaoomm 9004 mb 0.0H H.mN m.ON m.om 5% w aznnsh hm04 musp mus> 09Hw> osHm> ousbv mswpm .mvwso ooqdpAomaH mommpnomeH ooquHmH:MHm Aha aha .02 .oz 32.30: m can v mommeo m mmmHo N mmaHo H mmmHo .mm vnwpm mo mmmnp mom umHHasbo «mafiqu nOHpmasum .NMH mum_xmvnH abnannoo II II II >.N II II II unwOHuoaw usaHp we m.m II o.w «.0 H H anwOHhmaa wHHHa mH o.N II II II II II anpsHmb msoumso owe b.mb II II II II II wanna manumso mam 0.HHH II II II II II mnHm muonoso wN o.w II N.NH II II II uanonou magnum 8 ms N.Nm N.N .3 H H afififimhp 953 mm N.N II II II II II «NNNNNNHsp noNNNINONNNq mN m.m II H.0m 0.0N < m quOHnoam manHNdAh HOH «.0H H.5v «.HH II II II dHHoqunwnw mnmdm 5H m.m «.HH b.mN II II II wcHHOHm mannoo NH o.N II N.N II II II mNHapo ayou we m.b II II II II II mHEHOMHwhoo dunno II II II N.N m.mm o m «nuanaaoamo osnflguuo II II II o.m II II II .Qm hmHnonuHmad QmH «.mH o.qu >.Hv H.NHH hm m asnwnoowm hoo< 5mm 0.Hm 0.4H o.m N.QH m N ashnnh uwoq 05Hm> oaHm> Nada» mus> msz> mamvm .mcwso megaphoaeH moquuomaH ooqNOHmHQMHm ham hmn .02 .oz uopgmfios w chIw,nmnmeo m mmeo N mmeo H mmeo git; iii. 1% 989» no womb. you coHHEBo $3qu nogwgm .HNMH mama 268. $3 3H,; x85 3:528 Hm he I N.N. N.NH m N «33 35H: II II II N.0 II II II aquoHnoau usaHD omN m.wN w.Nm H.>H II II II anOHnoam uHHHa II II II N.o II II II aaanHu nauuunoam Ob 0.Hm II II II II II uan=H¢> msbnoua Heb 0.50H II o.m H.Nm 0H 0 unnbh muonoud II II II II m.m H H unHa «sonosa mH H.H N.N. N.0H mém “H m 2:398 355 II II II m.m 0.0H N N updpGachnuuw nsHsnom .HH. N.N HEN AHHN 0.3 H H 2353.? «bio wH m.m II II @.0 N H unan uanNuhh m5 m.NH m.o 0.0N H.mm mH H NQNOHuoau nannunh 3 ma m1: o.N ma H H «30.3.5.5 Ewan II II w.b o.N II II II .9» unmoupuuo mH H.m H.wm N.NH m.w m H 3HHOHH 35.30 HN 0.5 I EH II I I 336 «has mp o.N m.m N.H II II II nHanouHcaoo dunno I I NdN 0.HH I II I «fiHfiHofio 3593 II II N.w o.N II II II .mn ncHaonuHoaq ooN 0.0N b.mo N.mN H.5H H m enhanced» hood mHH n.0H bém H.MH H.mH m N 35.53 .304 3H; 3H,; N33 IoImHap and; an .mHflHa megaphonaH megaphogsH 00:60HMHQMHm nan nun .oz .02 833°: Idldalj; o ulHoiII m SuHo N 238 Hfluuwfl E .mm unapn no mocha you voHHnauo noOHqu naHvuaasm .HHNMH mamas 269. oomH msHm>_xquH auanpnoo HH .3 .... I .... .... II 338% gas II II II H.N II II II dandy ansHD II II II w.m II II II .283an wHEHD II II II b.H II II II unonnoau wHHHe II II II m.m II II II anHd mmhnommmm ONOH o.NoH II n.0H H.mm b N Nanak «soumgo oHH N.NN II II N.N H H «pHa maouosc Hm N.N II N.NN H.Ho HH H “NHHoaom magnum II II II w.m II I. II «HBHGHNHHP dhpmo NHH H.MN II II II II II «NIHHNHHsp conunIHOHNHH MN 3 N.N 0.3 0.3 w 0 «$0203 $33: NNN N.NN N.NNH H.0N II II II NHHoNHuqaum mamam II II N.N N.HH II II II moHuoHH mannoo NN H.H 0.HH H.m N.N H H samba shade II II N.N N.NH II II II aannHHonuo usNHaaao 8H N.0H 0.2. 2N I II I. £5508 .82 omH m.mH 0.HH H.NH N.0H H H aspnun HooH SH; SH; SH; SH; 033 mfipm 538 mommfioafin 0032095” ooquHMHanm 9.3 Arab . oz . oz copsmHoz m can H NQNNNHO m mmaHo N mmaHo H uNNHu .om unapm mo mNINP NON uoHHaaoo umoHunH qupusssm .HHHHNH mHmHa 270. meH 05Hw> menH assachoo II II II ¢.N II II II «GNOHHmaw “HHHH II II II H.N II II II avanHw mdhhdmmwm ow 0.0H II II II II II quanob machosc ooOH o.b© II o.m w.©H N N wansh machosc .Vm 5.3” II II II II I an? 98.33 II II ¢.©H H.«N 0.0H m N waHvohmm mandhm Hm H.0H v.3 m.$ v.8 HH m ufiHfimfip ubpmo Nb o.NH H.NOH m.bm w.¢m m m aquHhmsw manxwhm II II N.mH m.wH II II II mcHHOHm manhoo 3 H.0 I II I II I News «Eco EH 9%. H.Nm H.N H.N H H 3.38 2.30 II I o.N N.0H I I I é... pmHfifiHofi II II II m.OH <.m H H enhwnoowm hood mp N.N 3 N.NH H.HN m m as?» .32 mus> msz> wdeb oaHm> 05Hw> msmpw .mcwso ooquAOQSH momwvthEH monwOHMHanm Aha Aha .oz .02 @3332,» . m wad v mmmmeo m mmeo N mmmHo H mWwHo .Hm_©na qN mcnwpm Mo amen» pom cmHHmsoo mmofiunH GOprsEsm .>HNMH mnm II H H I II .I. .H H H ..I II I I . H 3 H H .2. II u m m H. H HI ...» H .w .2.» 3:328 33> II II II II II II II II II II II II II II II II II II .2 I. II II II II II II II II II II II mHH¢>Hum0m 59:» II b < OH c II II II II n II u II II N 6 II II II I II u II n II II II b m II II aschuoooh eschdnH> II II H II II x II I- II II II II II II II II H II II II II II II II II H II H II II II LnHHouHadhc EfiQHSDH> I II I I H II II II I II H .3 H II II II II I- II H II H II II H -I ,I. II H II II omficoH 5255..., II II II II II II II II II II II II II II II II II II II II H II II II II II II II II II II nochnHmmoo nsnthH> b o 3 OH I m H H m H. II H II 9H .w m m H m II I. II I H II II n m m I II :3 H328- 5&3; v9 cx h\ I“ ~t m m m 0 0 MI 4 m N H 0N mm 4m mm N.N HNCNCH cH ..IH 0H «H.H mHhH HH 0H0 m 93.5 Hm On 0 RI ‘n‘I .cvneac HHu c. cc;wHH£cHae m~ers2r .5 OH x or cow no n.0n :H cccaco:g caveman pagan uo accccguzooc uo hopazu Amachaccov .beuu mumdfi . I I: m . .. . , H ., . TI Idvm II, IIhrIQI-wifluIIflHHIJ; I . .I I . :.m«.§IIIII.IIuwI: I I.: VIII LHII:.IIIIIIILIM.H..I.IMH; . I- I . I . .. I . I r , I .II (1' rI:Ill .rIIhIJI. II I I . I I I f I II.IIII II TABLE W11. III-bar of men of herb upocin noordod in not. of ten 10 x 10 n. quadrat; 03%.!!th In Ill ltlndn. Stand 91011121314151!)17181923322824252627a2930313233343536 678 5 I“IIIII l”I"l"I IHIIIII I"I|I"I I"I“l"‘l l"I‘°l"'I Ill"I|I I“l"I"‘I I"I"‘I"‘| IIIIIII I"I"'l"'| éi: 5 s§§§ 3a 3 Elgafgi 5 E??? 25323:“ 2. 2355 H X_.__--_----—--_.. X_..-- II“l IV I II“ “R“ II” II” KIM II“ H:M’\ I ll” (“‘1‘ II” II" II” I I I I I I I I I 'I"! I I I I I I I l I I l I I l""I I 'l"‘I I II"‘I I ”IQI I I II"IIIIII II"‘|I"“‘|I IIHIIIIHI I""II"III I|"‘I""Ill Il"‘l|"lll I I I|||“"“” |I"‘I||ll Antennas-1a plantaginifolia Agroatia acabra II“I“‘II II"'IIII lIIIllI IIIIIII ll‘°ll"I II”II"I II"II"‘| II"I""I I"I I"I IIIIIII IIHIIII I"I Ill I"I I|"‘l IIHH 'IMH ||"'I .|"I III-ll III III-I II II“ Il"| >‘III I"I (‘1' IH I I I I I"II I I I III |IlI"‘I IINIII I IN 'HH below; pun-wrote.” Analog-Jinn tuberou- Arabis miaaourienais Amalia nudicaulis Aralia ramoaa Mel-pin oxnltau Arabia laovigata Aria- up. l"'II"‘Il I"I"‘“II IIIIIII I‘l"”‘lI ln—I‘OQI I‘°"‘I'°Il I‘°|I“Il 'l‘HIF‘N' l"‘||‘°"‘I |“II"‘II l”l"l"‘I “NIIO‘QI I"Il°‘lI lonloml I"!I"|I I“I|l"‘l IIIINII IIII‘II I'*II"‘II :Hm:~t~ol I“IIIII :NK:MH: I‘IISII 'Nw'bd: :III°II I“°l“°l IHIIIII IS"I“”I IH”IS“I INII"|I IHII”“| l“Il“I” IHIIIII I"II""I Astor uglttfloliul Ante:- Up. II“ |I'° II II II II II :N II II II II II MN II II I I Mun-inn run-rum Bani-1a tuctaria IIII |"I" IHII IIII l"‘ll I"|| I"‘II I"'II IIII I"‘Il l"‘I" I‘ll IIII I"'Il P‘Il I"'II l“|l I"II I"ll IIII l"'II I"II I"|I l"ll lIlI IIII Illl Il"l l"I| I"II I"""I I‘II IIII II"I I"""l II II II II II II I"II P‘II IIII""II (halt. atriplicitolh C-pululn inn-1m "rho cymbal 'X' indicato- wounoo 1:: Im- (mum)Mbuotomoudhorblpociunmdodmmdmhxml.quldntluhh1w1ndllm. um m1. Stand 91011121314151.617121920212223242526?]82930313233343536 5678 3 4 IIIII"I“"'I "IIIIII“°“ IIIIIII"I" IIIIIII‘°‘I I‘II"‘III“‘I' ‘°|III|I“'II IIIII"I“’II “Ill“ll‘II ‘°|I“'III"II IIIIIIINMI “III|”I“II |"‘Il"II"""I NIIIIIIIII IIIIIII""I "III"I|”II “II"IIII"I IIIIINIHII IIIIIII‘II *lIIIIl”"I "IIIIII‘II III"““II""‘I IIIIIIIIII "I“IIII"II IIIIIIIHII "IIIIIIHII IIII"I"‘III Ill"'|II"“°I IIIlIIIIII ”II"“II"'II III"III""'I IIIIIIIIII “II“III"”I IIIl"I|I"'l |II"IIII‘°I ll.|Il||III ll"IIIIlII . E §§35 gqu . uua’ 2 a 5:3: 838:3 naguzvagsg giggigaEE: ggsgiééié’é >>>>>>>> II '0! I H I I I l I I l I I I I I l I I l l l I I | I I I I II II "‘I 21:1: sure; 'l'ho lylbol '1' indium me. in land. Stand 91511121314151!)1718193)212223242526NZBZ930313233343536 7 8 6 III-bur of common or hub spun. ”corded in not. at an 10 x 10 I. quanta establish“ in .11 numb. TABLE mu. 274.. I"|IIIIII'INIIIIIIIIIIII”I“IIII”II“I|II"II|I“IIIl I"l"l“lIIII*IIIIl|III||I‘llIIII‘I“*IIII°II“III“I“ I‘llIIIIIII“IIIIIIIIIIII|IIIIIIIIIIIIIIIIIIIII":| I“III“III“I“I”IIII“IIIIIIIIlIII‘I”“IIIIIII“IIIIII I“I*I”|IlII“IIIIIIIIIIII°II“III””l°"IIIIIIIIII"II I"I°l“lIIII"IIIIII”IIIll~ll”llI“”l°”III“IIIIII“II IIIHIIIII“I"IIIIIIII“III”Il“|Il°“l”llII"ll*IIl“II l”l”I“lIIII“II|”ll“lIIIl”l“"l|I°II°IIII*II“III”II I"I”l"IlI”I“IIHIII*II”II“II”III°“I”"IIIIIIIIIIIII IIIIIIIHIIImlmlIHIINIIII“”l“III”lI°”III”I"I|:I”II I*|”|”IIIIIHIIIIIIIIIIMIW"”"III“II“IIII”II“III“I” II”II|III“I“II”“III"llIIIIIIIII”I"|”IIIIII"III*II I"IIIIIIIII“III“IIIIIIII”!IIII"”II°*III”IIIIIIIII I”l“I”IIlII“IIllIIIIIIII“”I"III*II’lIII“|I”IIl“Il IIIHI“IIIIISI””III“II“II IIIIII°”I°”III"II“"II“II IIIII“:IIII“:I“!Il”"”lIIIII“|lI"II‘IIIlIII“III"II IllII”!IIIIHIIIIIIIl“"I|”II*lII“III“III“II|"II"II I"IIIIIIIIIHINIIIIIIIIIIIII”IIIIIINIIIIIIIIIIIHII IIIII"IIIII"IIIIIIIIIIIIWII“ IIIII“II|I“IIII||”I| IIIIIIIIIII“IIIIIII”IIII”IIHIII“II“IIIIHIIIIIIIII IIIII"IIIII”IINIIIHIINIIIII*|II”“I””IIIIIIII||"I| IIIIIHIIIII”I“IIIIIIIIIII‘lllII"lIIIIIIIIIIIII“II "IIIII°IIII"“I“IIIIIIIl:”Il“III““I”“IIIIIIIIII“|I lIIII"IIIII”!lII“II”**II“II”III*IIS IIIIIIIIII”II *I"II”IIIII”II “III”!IIIIIIIIII“"I“”IIIIIIIIIIIII IIIII*II:II“III“IIIIIIII"II”“IIIII“:II"IIIIIIIIII ”IIIIIIIINIOI“II”I|“IIIIHINIIll”°l“°llllllll”|lll IIIIIIIIIII“IIIIIIIIIIIIIIIIIII”III|I|III|IIIII”I "IIII”III”I“II“"“I°“IIII” IIIIIS“I“"IIIIII“III“II IIIII“IIIII“II"”II““IIII‘II”III"”IS*IIIIIIIIII”II "IIIIIIIIIIHIIIIHIIHIIIIIIII”!I"II“IIIIIIIIIII”“I II“II“II*I*”II"IIII|IIII”lI”I”I”"I°|I""IIIIIII*II IIIIIIIIIII“I““III”IIII“IIIIIIIHII“*IIIIIIIIIIIII 3 IIIIllIIIEI“II“III°IIHIIIII“IIINII“I”IIIII"IIII"I . IIIIJNIIIIIIINHHNIIIIIIII":IIIIHIIIIJIIIII“III““I 5 Illl"lll*ll”llllII|"IIIII“I“”II”II““IIII”|III""II I E a I a a 5 gig igéa§s§§.§. 33:53 ”am. 3552; 3 as . §.sg=«§5§. gag. : a“: Ema It: 3 5‘ aa 9 EggsggfigzxfifigggiaaI§§éaéai.§§§a§:§§§ 32 Egg a?.3i g “amnga- 3m IIIIEfi-wéz‘iagwg s 5” a e 3.353883252 ggggg v..:33§iiaa°333~-9§§ 33 a 5 «gagamgsgafi- gnaw aaaassgsssgaogggjggg . §.. 3555 5 a 33555< £52:3::::::: :aa‘a£ (continua) hwdmmflhcbamiunwu uuxmmxmu.muuuuwnmaum. rm mm. Stand 910111231415161718192021222324252627129303132333435” 6 7 Voranicaatrun virginiana Vicia caroliniana Viola madmia Uvularia “sailifolia Viola padata Uvularia grandiflm-a Verbena micitolia Viola pubaaoem Viola aororia Viola atriata 'I'ho mbol '1' indicate. mum. in stand. labor of 0mm of barb spacio- rooordod in not. of Ian 10 x 10 I. qmdrau utabliahod in all atandl. (min-d) mm 1371!. Stand 15 16 1'7 1? 192021227324252617282930313233343536 12 1" 1!. 0101! I‘lIIIIIININIHIIIIII"III”I|IIIIIIIIIIIIIIIII“IIII I“III”III”II°IIII“I“““°I”I"I‘“IIII“II*IIIIIIIIIII lllIIlIIIIII”IIIIIIII”II“IIIIIllllIIl“l|IlIIIIIII IIIII”I”I“IISII”I”II”°IIII"II”IIll“|IllIIlIlIIIII I“III”I”IIIISIIIIIIIII*III”IIIIIII“II*IIIII”Illll I*III°I“IIII*IIIIIIII”“IIl"llI“III”II“IIIII“I““II I‘lIIIIIIIIIbIIIIIII""I“I““ IIIIII"|I"IIII“IIIIII IIII"IIIIIII°"III|"l""°IlIIII“lII“”II‘llIIINIIIII I"III"III|II°“IIIIIIII*III"IIIIIII“II*IIIII“IIIII IIII“””~”“IIW*"IIIIIHIIIIIIII”III|”II“"”III”IIIII IHIMIMIIIIIIHIIIIIIII”I“II”II”lII"lII“IIllIIIlIII II”I”I|II”IISIIII“III°IIII“lI“lII“”II””IIIIIIIIII l“lII“I"IIIIIIIIIIIII“IIII“IIIIIII“"IHIIIII*III"I |“III”I"IIIIFII|IIllI““IIIHIIIIIII°II”IIIIIII”III IIIIINIIIIIISIIIIHIIIHNIIINIIIIIIIS I”IlI“l*IIIII IIII”IIIIIII”IIII|IIlelIIIIIIIIII"II““II"I"II“II IIIIIIIIIIII"IIIIIIIINNIII"lIl|II”“II“l"IIIIIIIII IIIIIIIIIIINFIIIIHII|“IlIIIIIII”II“II”IIIIIIIII“I IIIII"IIIIII° IIIIIIIIIIII”III|III“IIS“IIII“III“I IHIIIIIIIII”W“II|”IIIIIIIIIIIII"II“II°“II"I“IIIII IIIII"IIIIII“:III”III””III*IIIIIII” I“III“IHIII“I IIIIIHIIIIIflmIlIIIIIIVIIleyIIIII “II“IIIIIHIIIII IIIIIIIIIIIIoIIIIIII*”"IIIIII”IIII°IIS"IIII“IIIII I”III" IIIII“:III”“II*IIIIIII“IIII“II°IIIIIIIIIII IIIIIIIIIIIISIIIIIIII"IIII"IIIIIII"II3”II”I|IIIII IIIIIIIIIIIISIIIIIIIIIIIIIIIIIIIII“II”IIIII“IIIII IllI”“IIII”I°“”I“”IIIHINIIIIIIIIIIHIIHINIIIIIIINI IIIIIIIIIIII*|IIIIIII”IIII*IIIIIIIIIISIIIIIIIIIII IIII““IIIIIISIIII“III“IIII”I”IIIII“II“III“I"III“I IIIIII|IIII|°IIIIIIIIHIIII“I*IIIII“II°”IIIIHII“II IIIIIIIIIIIleIIIIIII*IIIIIIIIIIIIIII“III“I“IIIII IIIIIIIIIIIIFIIIIIIIISIIIIIII;IIIII||“IIIII"IIIII "I”IIIIIIIIH°IIIIIIIIIIIIIIIIIIIIIII“°II“II“III"I IIIIIIIIIIII‘IIIII"I|”IIIIIIIIIIII*II“III”IIIIIII IIIINIIIIII“°;III”IIlmllIIII”III*I”II“"IIIIIIIII“ IIIIIIIIIIIISIIIIIIII”II|I“"|IIIII“II2"II”III“III a . a 5 E 53 3% g» 9% a? 5 3. 3 - § 83 a 3:3: 53:35: °2 s 2 33; I: Iiaggé .3 @353??? Isaaasafaf fisggéégéggaz.“iz 5% 5 EE agbgggsggdséii§§i.jg:§§ 1§38§§5553§§§§E§Ifig I 3 §€25§sga§§sssssgggtusgg ‘ *'° ‘5: =5 : 3§2g§§§§§§§§§§s 3§§§3§ IIIIIIIEIEIIIIIIEIEEEEIIIEIEEsagggggggggggéfiggsfis 0 0 00 0000000000 00000 D -II 275. “Tho mbol '1' indicato- mum. in “and. Stand 9101112814151617121920212223242526272883031323343536 8 6 ’I (continual) labor a! ocean-ences of herb apacias recorded in acts of tan 10 x 10 n. quadrats established in all stands. TABLE mvn. I:I“l1““:II”“III“IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII “I“:"“IISI°I2”III“:llII*IIIIIIIIIIIINIIIIIIIIIIIII “IIIIIII*III"”IIIIllIIIIIIIIIIIIIIIIIIIIIIIIIlIlll ”III‘”II”III**llISI“IIII”IIIHIIIIIIIBIIIIIIIIIIIII IIIIIHI°“IIII*IIIIIIIIII“I 2”!IIIIII|II”II|I|IIIII IIII“II”*III““IIIIIIIIII"IIS”IIIIIII‘lIIIIIIIIIIII ”III“II“"III*“IIIIIIIIIIIII”IIIIIIII*IIIIIIIIIIII° ll“I"l”“III“°III°IIIIIHHIIIIIIIIll"“ll|lllIIllll| IIII“IIW°III”*III||IIIII°I SNIIIIIIIIIH”IIIIIIIIII “I:I°~20MI:I~°:II~~“:::~<::II:I::a::<~:|:lzasllll" ”*"l“l|"““lilwllllIIIIIIIII*”IIIIIIIIIIIIIIIIIIIII IIII°IIH°I“IS°“II°IIIII"“IIIIIIIIIII’IIIIIIIIIIIII IIII”I“””IIIINIIIIIIIIII”IIS"IIII"II*IIIIIIIIIIIII "III”“”I“lII“°"II":IIIII”IIIIIIIIIII“IIIIIIIIIIIII Illl““l°°l:I"“”“"|III”II”IISIIIIIIII”II“III“IIIIII 'II”|““I”*IIIl”ll“°lI:“II”II“!II|IIII"I“II"““IIIIII IIIII“I°““III“IlIIIIIIII”IIIIIIIIIII"||IIIIIIIIIII :2::~:~~~::: “IIl‘IIIIIIIIIII“IIIIIII“I”IIIIIIIIII IIIII"II“III:* I:III:I:I”:l"IIIIIIIIIIIIIIIIIIIII" IIIIIHII°I”I”*IIIIIIIIIIHIIHIIIIIIIIIIIIIII”IIIIII IIII‘II“°"III“III"IIIII|“II°Illlllll"ll“l”l”llllll "III“III°"II““III"IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII IIII““I“SIII““III*IIIIII“II”” IIIIIIIIIIIIIIIIIIII “Ill“!"l’lflII”:Il“|IIIII”IIIIIllllllll|"llllllllll llll””l°”IllI°”Ill|lIlIIVIHNIIIIll”l“|l”|lll”l”lll II:~:~:I*:::Ibl::III:II:~::*IIII~~IIIII*II~IHIII~I lllII"”°”lIIIS IINI“IIII“IIIHHIIIIIIIIIIIIIIIIIII“ II"I||II”IIII“ III:lI“II“IIIIIIIIIIIIIIIIIIIIIIIII Ill|“"I*SIIlIS““I“IIIIII’lISIIIII"IIIIIIIII"IIIIII III””““I°III””IIIHI"“III“"l*|lllIIIIII“IIII“IIIIII III“"”I"°III*”Ill”lll"II”|I”ll“lIIIIIII“|IIIIIIIII IIII““““SIII“2III”II|”II"ll°”lllI“|II||”I|IIIIIIII IIII“III““I“I*I"I”III|“|“II°IIl“l|ll"l“"lll"l"llll III“IIII“I"II”IIIIIllIll“!IIIIIIIIII”II“I|IIIIIIII III““II"°“III“III°I"I”IINIIIllllIlll”l”lll“l|lIlIl lIIaHIIIIll“"“I|'“|II“II”|III|IIIIII”III“IIIIII"II 5 ° :3 fig 35 ' a a . E a n a a “s u§§V§ o a“: “ a - . agga.ao.§53 g§.Ia.g§g§§%a§2§§“§§II_§ aag'gggg 3 3 5 Egaysggééég.sgagsaoywaawtgagéaafie i§§§5%:52.§§ég Egga‘gsgfi53%533géégéifi'3ééé335222525358} 5:31:33 » £3555§§§5§”§ E a 35333335382£°°'33 E-E §_a g aéégsgaaga33353353Eag§§§§§33§5§§a§§£§§§ai IIIIIIE 0m systol 'X' indicates m in stand. Stand 9101.112131‘1516171819202122232425$278830313233343536 (continued) III-ho!- of oemmncas of barb spacias racoMad in sate of m 10 x 10 n. quadrats n-tahlishod In all stands. TABLE ann. :H:I:~OHIIF\IIHHIII~II I III IIIIIIIIIIIIIIIIIIIIII“I""I"‘III N"‘IIIIIIII"‘|I"‘II"“"’IIIIIIIIIII“‘I"‘IIIIII"I“’II"”"I‘”II IIII""‘IIIIIII"‘III"IIIIIIIIIIIII”IIIIII”II“IIIIII" NI"IIIIIIII"‘IIIII“IIIIII""'IIl“‘l"‘l"‘I"'II"""IIII""“II"‘ IIIIIIIIIIIIIIIII"IIIIIIHIIIIIIIIIIIIIII°‘IIII"III I“III|IIIIIIIIIIINIIIIIIIIIIIIIIIII”II|"“"III"I"‘II IIIII‘“II’"“°II"III"‘|IIIIIIIII"IIIIII“'IIIII‘°"III“I'° IIII|I|IIIIIIHIIISIIIIII"IIIIIIIIIIIIIIII°°II"‘"II“ IIII"'IIIII|“I"III"IIIIIIIIIIIIIIIIIHIIIISIIIHIHII "IIIIIIIIIIIIIII"”"IIIIII"”‘II"IIIII“‘I"‘I'*IIIII°°°‘III IIIIIIIIIIIII"lIII‘“|III-l"‘|IIIIIII"I”III|I°"“|IIII” "lIIIIIIIIIl""‘"'I“IIIIII"“"III"'I"‘I"IIII"“"4IIIIS°°”II NIIIIIIIIII"'III"I"‘|IIIIIIII“‘IIIIII"IIII|"I"‘IIIIII I"IIII"II"II""III"IIIIII"lII"‘IIIIIIIIII“’IIII”III“ IIII"'IIIIII“IIIIIHIIIIII"II"‘"IIIIII"‘III""III“'I"II "IIIIIIIIII"IIIII”"'III“I‘“‘I"‘IIll""IIIIII"’IIII"IIl"‘ IIII"|IIIIIIIIIII“IIIIIIIIIIIII“"IIIII"‘I°‘IIIIIII° MIIIIIIIIIIII"III"‘IIIIIIIIIIIII III"IIIII"‘IIIIII" III"IIIIISII"IIII l':|IF\ININIIIMIIIIIHIQIIIIIIIIIII:MH::H=I:‘IHIIIN ”IIINN::I:I“III:IM\IIlIll><|llIIrINfI‘:~'l-I::‘HNIII 'I:ll|:::'lll> 3.. H . 3 .I .. . E 3:55 .1 v flaw-‘35.. :3“... algae“ =33? 5-5? igézés 3§ v38§~ = 2%; 3 §% s sfiasgisfié an; ~* “gum mamgzwgzswgkaw ...,_E.oo.. o 5: a3 azbzofl-a Mada 0‘8 338 II g0. 009 .80 “suga‘gg I-aasgggggssfiasseiagaa=~s駧§cia§a§§ea§%»§:.g§I-a 502: v.4 dldfldg’ga £5333 oooooogooog‘gE-E 5:} a aéaagigggaigfigfifigIsaigggggIEE§§§§§§§§I§I§§§55§§§§ 2955 Eggggfimg3333355335833gfifigfifififigfifigéfiafiifiéiacz “Tho sylbol '1' indicatas panama in stand. _- -.‘NL "IIIIIIIIIIIIIIIIIs