FOREST SUCCESSION ON THE WELLDRAINED SOIL IN THE HIGGINS LAKE AREA OF MICHIGAN Thai: for tho Dogma of Ph. D. MICHIGAN STATE COLLEGE Kim Kwong China 1954 This is to certify that the thesis entitled Forest Succession on the WelléDrained Soil in the Higgins Lake Area of Michigan presented by Kim K. Ching has been accepted towards fulfillment of the requirements for Ph. D degree in Forestgx ”95/ 33;); ' ~ Major professor Date November 70, 1951, V‘A‘ . -\- ~i JAN 1 0 32-1 ‘9 i. 555 0,7)t994 , ‘ IA. 3‘ I AUG 05 2001 l \ FOREST SUCCESSION ON THE WELL-DRRINED SOIL IN THE HIGGINS LAKE AREA OF MICHIGAN By Kim Kwong thng A THESIS submitted to the School of Graduate studies of Michigan State Colleqe of agriculture and Applied Science in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Forestry 1951+ ‘4‘, . b04/57 G ,/ 5"." 4““ ii ACKNOl'iIEUCEtLENT The writer is grateful to Dr. T. U. Stevens for his advice pertain— ing to the study and for his assistance in the preparation of the manu- script. The writer wishes, further, to eXpress his appreciation to Dr. C. L. Gilly of the Botany Department of Michigan state college for in- formation, concerning the identification of plant Specimens collected from the field study. Thanks are also due to Drs. Robert Dils, Ivan bneider, A. E. Erickson, for helpful suggestions. Acknowledgment is also due to Dr. T. T. Ku for his valuable assis- tance and discussion in the field as well as in the laboratory. For his invaluable aid in obtaining information about the studied area, especial acknowledgment is due to the Regional Forester, H. V. Borgerson of the conservation Department of the State of Michigan. A.deep appreciation is owed Dr. victor J. Rudolphznriihu W3 B. Drew for helpful criticiSms during the preparation of this manuscript. Lastly, the writer is indebted to his wife, Teenay, who has shared generously in the organization of data and preparation of the manuscript. iii Kim.Kwong Uhing Candidate for the degree of Doctor of Philosophy binal.Examination: November 20,1954. Dissertation: Forest Succession on the hell-Drained Soil in the Higgins Lake Area of hichigan. Outline of Studies: Major subject: Forestry :Minor subject: Botany Biographical Items: Born: August 3, 1918. Honolulu, Hawaii. Undergraduate Studies: National Central University Chungking, china. l939-19k3. Graduate Studies: Michigan State College l9h7-l954. Experience: Research assistant, National Central University, China, 1943-l9h7. Forester, Division of Forestry, State of Ohio, summer of 1953. Photo-interpreter and stare-plotter, Kbrams Aerial Survey Corporation, lensing, Michigan, 1953 to present. member of.Xi Sigma Pi, Society of American.Foresters, The American Society of Photogrammetry. FOREST SUCCESSION ON THE WELLmDRAINED SOIL IN THE HIGGINS LAKE ARDA OF MICHIGAN By Kim Kwong Ching AN’ABSTRACT Submitted to the school of Graduate Studies of Michigan State College of Agriculture and Applied Science in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Forestry Year l95h (.7, it ‘**——mf*m {aw dfi‘”'”‘ ‘fiv - act‘s... -.< a s »-.' .assr-A" .dm£-w Approved {1 t?" ”if. ‘35 _s' t» m-;' r'. " We'h‘ Kim.K. Ghing ABSTRACT The upland forest of Higgins Lake area were studied in an attempt to set the pattern of secondary succession on this vast cut over land. Quantitative data were collected from five different vegetational covers - grassland, oak, aspen, jack pine, and conifer-hardwood types, by the qua-— drat method of sampling. Stands for studying were carefully selected so that (1) they represented natural forests of adequate size; (2) they were free from disturbances in the forms of fire, grazing or excessive cutting; and (3) they were on upland land forms on which run-off water never accum- ulated. In the introduction, the place of forest ecology is briefly outlined. The basic information of original and present forest cover were also dis- cussed. Sections'following were devoted to analysis of the environmental factors listed as climatic, edaphic, and biotic. Comparisons between the composition of different Soil types and profiles were subjected to ana- lySis. Further work dealt with the develognent of plant communities aml their interrelationship with various soil types and the influence of environmental factors on these different vegetational covers. Through statistical studies, the following facts have been established: (1) evaporation increased rather uniformly with increased temperature and presents a more definite eXpression on different forest cover types. (2) the forest soils of most of the studied stands are strongly acid in.the upper portions of the profile, especially the A layer. (3) in the presence of a sufficient supply of available nutrients, pH of soils is of minor importance to growth of plants in the studied stands. (4) the mature content of soil in the field varies according to vegetation types on different dates, but not at the six and eighteen inch depths. (5) the covering vegetation do not influence the soil organic matter content in the six sample plots. (6) soil temperatures are significantly different under various plant comiunities with the grassland type having s definite higher reading. (7) anung all the factors which fall in the biotic group, destructive logging and fires are largely responsible for the present dis- tributien of plant growth. It was the opinion of the writer that the forest successions]. trend on this cutever land will progress in the following manners: the closed canon of an aspen er elk stand will give place to a higher genetic type, the most frequent ones are pines on the sandy upland, whereas on the bet- ter soils, the invading species would be the beech, sugar maple on! yellow birch predominantly. Baler the present prevailing climatic condition, on the more xeric habitat with decreasing soil moisture, a red or White "pine stage may be reached before the arrival of the formation of the north- eastern coniferous climax. On more hydric sites, the conifer-hardwood type is often succeeded by the northeastern deciduous forest climax. vii Table of Contents Page Lj- 3t or Tables 0 O O 0000000000 e O O 0 O O O O O O O O O O O 00000 O 000000 O 00000 O 0000000 i¥~x List of Figures.. ....................... . ...... . .................... :1. flat or Phtes O O 000000 O O O O O O I O O O O O O O O O O O O O O O O I O O O O O O 0 O O O O O O O O O O O O l 0 0x11 1. II. III. IV. V. VII. INTRODUCrl'IONoeeeO eeeeeeeee eeee eeeeeee e eeeeeeeeeeee eeeeeeeeeeeel-3 REVIEW OF LITERATUSES 1. On the Studied Area .......... . ........................... ..h~5 2. Concept of succession.. .............. ......................5-9 DESCRIPTION OF THE STUDIED AREA 1. 2. 3. h. 5. 6. 7. Climate......................... ...... .....................10bll Geology.. ......... .............. ......... ..................11912 Original Vegetation........................................12-1A Present Vegetation.........................................15 Description of Various Types of Forest on the Well-drained Soil.......................................................15-22 History of Economic and Social Changes.....................22-2h Present Land Use...........................................24-25 THE STUDY OF VEGETATIONAL COVERS 1. Terminology.. ...... . .............. ......... ...... ... ......26-29 2. Method of Sampling........ ..... . .............. . ....... .....29-32 3. Observation Grassland Type.................. ......................... ..33-37 Aspen Type.............. ........ ............ ...... .........38-h3 Northern Pin Oak Type........................ ...... ........hh-h9 Jack Pine Type.................. ..... ........;.............50-56 Northern Hardwood-Hemlock Type............ ....... ..........57-6h THE STUDY OF EDAPHIC FACTORS 1. 2. 3. A. 5. 6. Physiographic Features and Description of Soils Under Different Vegetation Types.................................65A7h Topography.................................................75 soil.Reaction........................... ...... .............76-78 Soil.Temperature...........................................78-81 Soilnhoisture..............................................81—85 organic MatteroeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeOeeeee0000085—8‘7 THE STUDY OF CIJMATIC FACTORS l. 2. 3. A. Temperature................................................88-89 Precipitatime....ee.oeeee..................e......e.....ee89-9o Relative Humidity.............. ....... .....................91 Evaporation................................................91r99 Tm STUDY OF BIOTIC FACTORSOOCOOOOIOO.0.00.00.00.00...00......100 2. 3. h. Maneeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeee eeeeeeeeeeeee oeeeeOOIOI-loh Animals..... ...... ......... ...... ..........................10h-105 Insects................... ....... ..........................105-107 Diseases........... ........ ................................107-lO9 V3-13. Page VIII. DISCUJoION OF Fender oUOuneolON IN THn oTUUILD AnflA.........llO-ll9 IX. blLVICULTURAL suoessrlws.... . . . ...... . . . . .120-126 X. UULMRYeeeeeeeeeeeeeeeeeeeeeeeeeeeeeaoeeooeeeeooeooeeeeeeee0127"].30 eeeoeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeseOOOCBB-u3 BIBLIOGRHPHY.......... PMTESOOOOOOOOOIOOOOO... Table 1. 2-50 List of Tables Page Major Changes in Land Use from 192A~1939 in Roscomnon Comty’ Micmgan.0..OI...0..O...OCOOOOOOCOCOIOOOOOOOCOO...0.24 Data of the Vegetation in the Milacre Quadrats of the Grassland Type as Recorded in Percentage of Coverage or Number of Plants by'Height Classes..........................34-37 Summarized Field Data of the Four One-fifth Acre Quadrats or the ‘xsmn Tlrpe by Size ChsseSQOeeeeooooeeeeeeseeeee0000039 Data of the Vegetation in the Milacre Quadrats of the Aspen Type as recorded in Percentage of Coverage or Number of Plants by Height Classes..........................41-43 Summarized Field Data of the Four One-fifth acre quadrats of the Northern Pin Oak Tym by Size Classeseeeeeeeeeeeeeeeeeeeler 12-15. Data of the Vegetation in the Milacre Quadrats of the 16. Northern Pin Oak Type as recorded in Percentages of Coverage or Number of Plants by Height ClasseS..............h7-h9 Summarized Field Data of the Four One-fifth Acre Quadrats of the Jack Pine Type by size Classes.......................51 17-20. Data of the Vegetation in the Milacre Quadrats of the Jack 21. 22-25. Pine Type as recorded in Percentages of Coverage or Number Of Plants b5, HGight Classeseeeeoeeeeeeeeeeeeeeeeeeeeeeeeeeee53’56 Summarized Field Data of the Four One-fifth Acre Quadrats of the Northern Hardwood-Hemlock Type by Size Classes.......60 Data of the Vegetation in the Milacre Quadrats of the Northern Hardwood-Hemlock Type as Recorded in Percentages of Coverage or Number of Plants by Height Classes..............62-6A The Depth,pH values, Content of Organic Matter and Mecha- nical Analysis in Various Horizons, the Soil Types and Cover Tyws in Six Sampling StationsoodOIOOI...OOOOOOOOCOOOCCOOIOOIYA Analysis of Variance of pH value in Different Soil Horizons and umer Diffemnt Cover WSOOOOOOOOO0.0.000000000000000077 Soil Temperature at Six Inches below Surface in Six Different Cover Types on Different Dates..............................79 Analysis of Variance of Soil Temperature....................79 Table Pa 36 30. Percentage of Soil Moisture Content in Different Cover Types .0. 83 “by COlOISIa ham curve.............a...........o...o.... Percentage of Soil Moisture Content in Different Cover Types 31. “By Dam-Bianca's culweOOOOO0.0.000000......OOOIOOOOIQOOOOIQSB 32. Analysis of Variance of Soil Moisture in Different Cover W8, Depths and DelteSOOOOOOOOOOO0......0.0.0.00000000000000085 33. Analysis of Variance of Organic Matter in Different Cover Tms am HorizonSOOO.0..O0..OO...O'CCOOCDOOCOOOOOOO...0......8’7 34. Climatological Data recorded at Higgins Lake State Forest Weather Bureau for the month of June, July, August, Septmber and OCtOber’ 1952.00.00.0000000000'DOOOOOOOOOOOOO0.00.00.030.090 35. Weekly Total of Evaporation in Four Cover Types. Relative Humidity; Air Temperature and Total Precipitation in Higgins lake [ireaOOOOOl00.0000000'OOOOOOOOOO....00...0.000.000.00000009A 36. Analysis of Variance cf Evaporation in Different Cover Types and on Different Dates..................................95 Figure l. 9. 10. list of.Figures General Scheme of the Original Distribution of Forest Species in Relation to the Water Table, Texture of the Soil and Pod- solization in the Glaciated Area of the Podzol Region of the Mk8 SmtesOOOOOOOOOOOOO0.0000.000000...00000000000000.0000...” Map of Lower Michigan Showing Boundary Between the Beech- !aple and Hemlock-White Pine-Northern Hardwoods Regions, and its relation to Tree Ranges and SOilooeoooeooooooooooooon00.0921 Map of Higgins Lake State Forest Showing the Location of Sampling Stations and the Surface Formations...........................32 Phytographs of Important Species in the hepen Type............40 Phytographs of Important species in the Northern Pin Oak Type.h6 Phytographs of Important Species in the Jack Pine Type........52 Phytographs of Important Species in the Northern Hardwood- Hen’llOCk Tymtl0.0.0000.000.00.00.00...000.00.000.00.0.000.000.61- Schematic Presentation of Typical Profiles of Various Soil WSOOOCOOOCOOOOOOOOOOOOOIOOOOOOOOO.00¢OOOOOOOOOOOOOOOOOOOOCO73 The Relation of Soil Temperature to Date in various Cover WSCOOOCOOOOO.00.00.00....0.000.000.00000......l00000000000080 Soil.Moisture of Various Cover Type by Using Coloma Sand Curve Deterlf'ined by Bo‘lyOUCOSOOOO0.000.000.0000OOOOOOOOOOOOOOOOOOOCCBA Graphs Showing Interrelationship among Various Recorded Data on Soil.Moisture, Precipitation, Air Temperature, Relative Humidity and Evaporation on the GraSSland Type covereoooeooooooeoo0.9.996 Graphs Showing Interrelationship among Various Recorded Data on Soil Moisture, Precipitation, Air Temperature, Relative Huhidity and Evaporation on the Aspen Type Cover..............97 Graphs Showing Interrelationship among Various Recorded Data on Soil Moisture, Precipitation, Air Temperature, Relative Humidity and Evaporation on the Jack Pine Type Cover..........98 Graphs Showing Interrelationship among Various Recorded Data on Soil moisture, Precipitation, Air Temperature, Relative Humidity and Evaporation on the Northern Hardwood-Hemlock W Coveroo-oocooQOooooouoo....o..oococo...ooooooooaooooooooo99 Pa ge Plate 1. 2. List of Plates Page An Open second growth pure jack pine stand. A fairly open site is a requisite to the establishment of pine seedlings......138 The stands of oak type occurring on ridges are often of poor quality. Note the thick growth of blueberries on the ground....139 3. h. 5. A typical stand of the aspen type. Some white oak, red oak amrw I'flple aremsqmure...0.00.0.0...OOOOOOOOOOOOOOOOOOOOOOMO A representative stand of northern hardwood-harlock forest. A well-stocked understory of saplings and poles of various species has developed under protection from fire................lhl A typical soil profile of Kalkaska loamy sand under a hardwood stand consists principally of hard maple, beech and elm.........lh2 A dense stand of young jack pine has successfully encroached and developed on this grassland area on a sandy soil............lh3 I. INTRODUCTION ‘With the national forests, Michigan has over 6,000,000 acres of publicly owned forest land, nearly one-third of the total forest area of the state. These public lands are a source of current and future supplies of tinber and at the same time serve a great and increasing public demand for hunting, fishing and other forms of recreation. After a century and a half of settlenent, land clearing, lumbering and forest fires, the area adjacent to Higgins Lake is essentially a cut- over region. The non-timbered character of much of the surrounding ter- ritory-ethe Great Plains to the west, and intensively farmed and indus— trial areas to the south-~created demands for’tinber from Michigan and was a factor in causing early liquidation of the original pine forests. likewise it is a feature deserving sone attention in connection with future timber growth goals. Today the region faces the problem.of supplying as much as possible of its own timber needs without paying excessive cross-country freight, of supporting a very valuable pulp and paper industry and other wood- using plants, of giving employment to settlers in the poorer agricultural districts, and of stimulating an active tourist and recreation industry, all on a foundation of young second-growth timber. Only a very small rewnant of the original virgin forests remains, and relatively small areas of second growth have reached sawbtimber size. Nearly nine-tenths 0f the land Which supports second growth from seedling to pole-8159 trees is in very Poorly stocked stands. In order'to control the destructive agents that threaten to curtail and impair the usefulness of the forests, and promote and maintain con- ditions favorable to multiple use with a minimum of detriment to the forest, one must rely on the application of sound silviculture, which in turn is based upon biological facts and principles. Intelligent manage- ment of our forests cannot be achieved without thorough knowledge of the behavior of tree species and stands. Nature, unguided by man, produces a forest which is in commflete harmony with the soil and the plant and ani- mal life it supports. Such a forest is the climax forest toward which vegetation is always tending. Stable tree associations, characteristic of climax types, are best adjusted to meet the impact of counteracting forces. Such forests are inherently healthy; under good management they are easily maintained in a high state of vigor, thus increasing the growth-rate, their capacity to resist damage from insects, disease, and other destructive agents. However, this is not meant to inmfly that the climax forest type necessarily represents the ideal toward Which management should invariably be directed, for other disturbances such as fire and repeated cutting, may alter the soil conditions to such a deg- ree that the early establishment of Species natural to the site may find difficulties to survive; or certain subclimax species possessing a higher value as far as their lumber is concerned, may justify efforts to main- tain them as dominants in the stand. Nevertheless, compositions charac- teristic of climax associations should be used as guidea for setting up silvicultural objectives. The present paper deals with the general trend of succession in the major forest types on the well-drained soil in the Higgins Lake area and their interrelations with each other and tith certain physical factors of the environment. The plan of the investigation used in this work is an adaptation of the mass collection method used by taxonomists. (1) It is a well known fact that variation in floristic composition is one of the most important characteristics that may be determined in the study of any vegetation, and this may be understood only after a great many examples of the type have been analyzed. It is important that the initial ideas of the type limits be suffi- ciently broad to preclude the dangers of subjective selection of only those stands which fit a preconceived notion as to what a particular com- mtmity should be. The criteria used for the selection of stands for this study were: (1) that they be natural forests of adequate size, not arti- ficially planted; (2) that they be free from disturbances caused by fire, grazing or excessive cutting; (3) that they be on upland land forms on which run-off waters never accumulate. II. REVIEW OF LITERATUAES 1. On the Studied Area The northern part of the Lower Peninsula of Michigan lies in the edge of the deciduous forest province, and is also occupied by the sou~ thern border of the northeastern conifer province. These facts are res- ponsible for the great variety of associations present, and the succes- sional relationship are often much involved in consequence (75). Current opinions and activities toward revegetating exploited and submarginal land in the Lake States (84, 53, 54) justify the eXpenditure of considerable time in improving our knowledge of the behavior of native Species and the successional trends under the stress of prevailing con- ditions. Numerous papers (43,31,67) have appeared during the past few decades dealing with the general ecology and distribution of types of vegetation in.Michigan. Dice (31) placed this region in the Alleghenion biotic province. The thorough investigation of Michigan flora have been well done by Beal (68), Farwell and others (35,67). Vegetation of the sand plains were given special attention in these preliminary studies. It has also been shown that there is great competition between the various associations of the above-mentioned two complexes wherever they come in contact, and that in general the associations of the southern complex tend to diaplace those of the northern. This was early demonstrated by fihitfcrd(102), Gleason (42) and Quick (75) who discussed the general distribution of the associations throughout the lower Peninsula of Michigan. The early days of lumbering and destructive forest fires which had alarmed the country caused many investigations and suggestions as how to utilize this vast area of cut over land (2) (A7) (84) (#6). Other investigators such as Veatch (91) reconstructed the forest cover of’Michigan from soil maps and determined the correlations between soil types and forest growth. Terminology used in this study is essentially as defined by Brann- Blanquet (It), and names of forest cover types were adapted from the publication of Society of American Foresters (81). Reference is also made to Cain (23) Who has discussed the forest climax and its comple~ xities in his work concerning some phytosociological concepts. 2. Concept of Succession In forest areas, the perennial herbs are soon superseded by woody plants, which become dominant; when a cultivated field is permitted to lie fallow, it produces a crOp of annul weeds the first year, numerous perennials the second year, and a community of perennials thereafter. If any disturbance of natural vegetation occurs-~such as cultivation, lumbering or fire, a similar sequence of communities appear'with several changes in the dominant vegetation through the years. This is the general trend of plant succession. However, it was not until.the sevens teenth century that any systematic study of such changes was made, and those studies dealt primarily with the development of peat bogs. Bog studies were continued in the eighteenth century and in addition some attempt was made to apply principles of Ecology to burned and disturbed upland areas. It was then that the term "succession" was first applied to the vegetational changes involved. It was during the years of 1860 (26) that a regional study of vegetation in Central.Europe was made in which succession was recognized as fundamental to all community develop- ment. Between 1890 and 1905, the modern concepts of succession were clarified through the efforts of several workers such as Uowles, clements. The idea about succession is that plant communities are never completely stable. They are characterized by constant changes, sometimes radical and abrupt, sometimes so slow as to be scarcely discernable over a period of years. These changes are not haphazard, for within a climatic area, they are predictable for a given community in a particular habitat. This means, of course, that similar habitats within a climatic area support a sequence of dominants that tend to succeed each other in the same order. Contrasting habitats do not support the same sequence of communities. As a result, any region with several types of habitats will have an equal number of possible successional trends. In the course of studying plant succession, a specific, immediate cause of a particular change of species may not always be Obvious because of the interrelationship of controlling factors. However, two general types of habitat change may cause differences in the community, namely ’ development of the community causes parallel developmental changes of the environment, and physiographic changes can likewise modify the envi- ronment materially. Developmental changes of the environment may result from reactions upon the habitat by the organisms living there, such as the case in accumulation of litter on the forest floor affecting run- off, soil temperature, and the formation of humus. Accumulation of litter, in turn, contributes to soil development, modifies water rela- tions, available nutrients, pH value, aeration, and affects soil orga- nisms; whereas the habitat may also be modified by forces quite apart from the effects of organisms. A flood plain or swamp may become better drained as a stream cuts more deeply into its channel; and silting in off a lake raises the level of mineral soil. Such modifications of the habitat also produce vegetational changes. These two types of habitat changes caused by the development of the community or physiographic origin are commonly in Operation at the same time and their effects cannot always be readily separated. Ecologists of various parts of the world have agreed that the type of plant succession on a bare area where no vegetation has grown before is designated as primary succession. It may be observed on glacial moraine eXposed by recession of ice, or a new island or any similar habitat newly exposed to colonization. A migratory advance postulates the development of individuals in territory not previously occupied and requires an environmental change of sufficient extent in order to permit the growth of the migrants. Each successive group of species to colonize a new area meets with increased competition. Succession results when migration is so complete and is shared by individuals of so many species that the nature of the vegetation is fundamentally changed. Moisture relationships usually control the ability of the pioneering plants which invade the new area. If the habitat is extremely dry it is described as xeric; if wet, hydric; and if intermediate, mesic. The successional trends are similarly referred to as being xerarch, hydrarch or mesarch succession. Whatever the condition of the initial habitat, reaction of vegetation tends to make it more favorable to plants and always results in improved moisture conditions. Thus xeric habitats become more moist and hydric ones become drier as succession pregresses. On the other hand, secondary succession results when a normal suc- cession is disrupted by fire, cultivation, lumbering, Wind throw, or any similar disturbance that destroys the principal species of an established community. Although the first communities that develop again.on this area may not be typical of primary succession, the later stages again are similar. The rate of this vegetational change depends, of course, on the severity of the disturbance. All successional trends lead toward relative meSOp'ny‘tism Within a climatic area and eventually lead to a single community, which is comp posed of the most mesophytic vegetation that the climate can support and Whose moisture relations are average or intermediate, for the region as a whole. This community, determined by the climate, terminates succes- sion and is called the climax community or climax for that climatic area. It is capable of reproducing itself, and since it represents the last stage of succession, it cannot be replaced by other communities so long as the climate remains the same. It is, therefore, a stable community in which the individuals that become overmature and die are replaced by their own progeny, leaving the character of the community unchanged. Just as portions of a single species may be isolated during its migrations, so areas of one type of vegetation may be completely sur- rounded by an advancing flora and left isolated from the main body. These are known as relic colonies. in important cause of such isolation is the failure, up»to the present time, of sufficient environmental change to cause their extinction, or, in ecological terms, it is the relative slowness of the succession in certain habitats unfavorable to most of the advancing species. 10 III. DESCRIPTION OF THE STUDIED AREA 1. Climate All areas of the Higgins lake area have a similar climate with essentially the same distribution of precipitation and temperature dur- ing the year. The main features of the climate are a mean annual temperature of 41.43.10 F., a normal precipitation (including melting snow) of about 27-30 inches, probable annual snowfall of about 60-73.6 inches, low wind movement, low evaporation, low percentage of sunshine and moderately high humidity. The precipitation (rainfall and melted snow) is quite evenly dis- tributed throughout the year, except that the six months from April 1st to October 1st, get a little more than the six months from October let to April lst. There is, of course, some noticeable variation during different years. As much as ll inches below normal distribution has been reported, and as mch as 17 inches above normal. The rainfall generally occurs as low gentle rains or showers; but cloud-bursts, hail storms and tornadoes are rare. In the forest, how- ever, the evidence of past wind and hail storms is shown by the old windfalls and the hail marks on the trunks of the pines. The prevailing Winds are westerly. The summers are characterized by moderate temperature, With a seasonal average from June to August, inclusive, of 63-65.9° F., and a 11 high percentage of sunshine. The pleasant summer compensates for the length and rigor of winter. snow forms a permanent ground cover ordi- narily from November to early in April, freezing to sub-zero temperatures are common. However, the winter has many pleasant days when the tem- perature rises well above freezing. Real spring weather usually starts during the latter part of April or early in May; The growing season, that is the season ordinarily free from killing frost, will average about Ila-122 days. The season extend- ing from the latter part of.hay to the middle of September is the average frost free season, but frosts of sufficient severity to kill tender vege- tation have been known to occur during every month of the year (25). 2. Geology Geologically the lower Peninsula is known as the Michigan Basin, because the rocks are all sedimentary, and the strata lie one upon ano- ther'like a pile of saucers with smaller ones towards the top. The hississippian, which is known as a part of the Carboniferous time was one of abundant vegetation and brackish and shallow waters in the Hichigan Basin. Rocks of hississippian age outcrop in a broad some- what circular band occupying most of the Lower Peninsula, except about a dozen of the central counties. Mississippian rocks consists of sand- stones, limestones, shales, salt«and gypsum, and are perhaps the most important sedimentary rocks in michigan (62) (50). At the time of the glacial period some 15,000 to 50,000 years 380, the great ice-sheet coverediuichigan and extended as far south as the valley of the Ohio River. After several retreats and re-advances, the final retreat of the ice left the state with its former rock.masses planed down, its former valleys filled, and its surface covered With a general mantle of rock-debris and drift with variation in thickness. The soils thus derived are various in character, as well as in origin (61) (Figure 3). The significance of this fact was pointed out by Beal (5,6,7,8) in his publications on Michigan Flora that probably three-fourths of our plant Species are common to all sections, though by no means equally distributed, some being very abundant in one district and rare in ano- ther at no great distance. In most cases such change is due to soil rather than to differences in elevation, temperature or atmospheric moisture. 3. Original Vegetation The entire land area of Northern lower michigan at the time of its first occupation by white men, was covered by a dense forest except for a small inconsequential acreage of bog or marsh and some open land on the drier sand plains. most of this virgin forest was white and red pine, With smaller areas of hardwoods and swamp forests (#3). In its virgin condition, the forest was node up of several types of forest-growth or tree associations that were distributed with close re- gard to the natural differences in soil character and drainage. The following types of forest were represented: (l) The pine forest in which White, red pine or red and jack pine predominated; (2) The hard- 13 wood forests in which hard maple, beech, yellow birch and hemlock were the principal species and elm, basswood, ash and white pine were sub- ordinate species; (3) The mixed deciduous and coniferous forest, in which such species as oak, elm, ash, red maple, aspen, and yellow'birch were intimately associated with white pine, red pine or red and jack pine predominated} (h) The lowland Hardwood Forest in Which elm, black ash, yellow birch, red maple, white birch and balm of Gilead were the principal species, but seldom.without a subordinate amount of white cedar, alsan fir, and tamarack; (S) The coniferous swamp.forest cone sisting of a mixture of white cedar, balsam fir, black spruce and tamarack, but including also limited areas of more nearly pure cedar, spruce, and tamarack (91). There were, of course, innumerable Situations where two or more of these associations merged to occupy soil and drainage conditions of intermediate or transitional character and deveIOped a forest of more diverse composition than outlined above. The following diagram (Figure 1) prepared by'Wilde (97) fully illustrates the general distribution of the original forest types on different geologic formations in the Lake States. 73.: 83.5 .613 motto 3.3 an. as 588. tram 9: t 39¢ 136,645 9:. E Sofieuzomvom use :om 9:. we write... J36... Lows? 2c. 3. c6333. 5 homes}. wmuyou we intacttfi .eEmIO mi. wo wfiosom .323 ._ oysnfi :33 Lamonnmidam team 3303 3 «no: 33663.0 Yam cawLomumteam £836 $383 :93.6 Eta tweets: 2.38: 30: «1:35:39 . JusZ «3°th 326w. acadeOZIclmwwr: 1 ' . e c s . n...” . .. . . . 1. .o V Ill-I" ‘ , . . . ,_. . . . o. s e u s I. c... O I . \I So, 3.3.153 ll I! Is c, _.w .cOo‘f I 0,, 0. . 4 . L. em. :3 a2} 32% .35 use 333 JR“ wen 1%? sets.ch s 3: . . .9: m w diagonhmew. - .8:th 323 bu serum .m 5....mrc F2552 x376”. 3 (In Edflndm Neil ”3&3 , . . . cindtf»$.43 $32.33 did: harm 15 A. Present Vegetation Much of the rolling to hilly part of the former pine forest land has been converted to desolate stump land, covered with a dense growth of brush, briers and grass, or has grown up to aspen.scrub oak and red maple with very little natural reproduction of the original dominant species. The level pine plains now support a patchy stand of jack pine with sweet fern, low blueberry, bracken fern and grasses in the open- ingS. The hardwood forest lands where not cleared for farmland, are now partly occupied by stands of second growth hardwood of varying age, den- sity, and quality, and partly by the growth of briars, grasses and tangled reproduction that has covered the burned over slashings. The Open bogs have a low shrubby growth of leather leaf, Labrador tea, and blueberry with sphagnum moss and cotton grass. The marsh growth con- sists of a variety of sedges with reeds, cattails and blue-joint. The swamps have suffered less change in the character of their forest growth than the uplands. Poplar, alder and willow have encroached on the severely burned cut-over swamps but elsewhere the present growth contains a generous reproduction of the original species. 5. Description of Various Types of Forest on the WelleDrained Soil In? White pine. This type of forest is of little importance in this study because the natural growth total acreage is small. The original pine stands of this type have nearly disappeared due to destructive logging practices and subsequent fires. Today only one area of virgin white pine remains near the region, the Hartwick Pines, located near 16 Grayling in sections 15, T. 27 N.R. 3W. The white pine, according to other reports, very seldom is found in pure stands, but often mixed with a small percentage of red pine (58). Mixture with other species is also quite common so that on the sandy sites the white pine stands uSually contain.more or less red pine, oak and aspen as the principal associated species, while on the more loamy sites the associated species may con- sist of hemlock, maple, beech and yellow birch. So far the white pine has not been able to reproduce itself to any general extent on the area formerly occupied by the virgin white pine stands. The redgpine type. This type, like the white pine, was one of the most valuable produced in the virgin forests and likewise was eXploited at an early date. Today, its importance in the region is small because of its limited acreage. The red pine, like the white pine, has also been unable to reproduce itself extensively because of repeated fires that followed the destructive logging methods (102). The red pine type is generally limited to the so-called "poor site", i.e. the drier and sandier types of soil. This is largely'due to the fact that red pine cannot compete with the hardwoods for continuance as a type on the bet- ter classes of land which favor hardwood reproduction. Jack pine type. It occurs both in pure stands and in association With other species. It is found mixed with white and red pine on their preferred sites and is commonly found grOWing with oak and aspen on the drier white and red pine sites. Repeated burning on red pine and sandy white pine lands usually results in a stand of pure Jack pine or jack pine and poplar. The type is rather wide spread and has persisted in l7 spite of the abuse from cutting and fires. Jack pine is considered by many as a "weed tree", since it so commonly takes possession of old abandoned fields and heavily burned lands in this part of the state. It is the principal forest type on the admittedly poor, loose, dry, strongly acid, deep sandy soils. This species starts to seed early in life and usually reproduces itself well under ordinary conditions since dense stands commonly cone in on the pine areas that have been recently burned over. The chief defect of this type as it exists today is the fact that the stands occur in.densely stocked clumps with open Spaces and poorly stocked areas in between. The present utilization for jack pine is principally for box.boards, lath and some rough lunber, though -in other localities it is also being used for pulp. Northernppin oak type. This is the most widely spread species of the region. Formerly, because of the prevailing poor site condition, the oak stands were generally considered of little future value for commer- cial production. They were commonly understocked, short-holed as a rule, and frequently showed much fungus and fire damage. At present, under the protection from fire, in general the diayeter growth of these stands has been fair and the height growth has been satisfactory. To-day a large proportion of this type consists of open ragged stands of straight bole trees and clumps of Sprouts of other species. Red oak, white oak, Hill's oak are commonly represented in the oak StandS. The common associated species are aspen, red maple and frequently JaCk pine. This mixture produces open stands Where all individuals have plenty of room to grow. Occasionally scattered red pine- a remnant of 18 the previous pine types- is also included. The oak-type reproduces easily from seed and sprout and has gained ascendancy over land where previously only individuals existed in the virgin stand. In fact, there is little indication that a pure oak type existed alone in the virgin forest. The type owes its existence largely to the fact that the oak was able to replace white and red pine after the pine was cut. It is the most common type on the rolling to hilly sandy and sandy loam.soils. This type is apparently there to stay for a long time since it occupies areas on which red pine, together with sore white pine formerly grew, and there are only a few scattered pine seed trees left. To all appear- ances, the red pine is not restocking these areas. Oak will likely remain the dominant type in these areas unless sore other species are artificially'planted. It has a rather wide and general distribution over the region as an associate in corbination with Jack pine, poplar and red maple. With fire protection, tie material can be produced but Where fires are allowed to reburn the area at frequent intervals, the oak is reduced to a clump growth. Northern hardwood hemlock type. The principal associated Species of this upland hardwood type are sugar maple, beech, elm, basswood, and yellow birch With an occasional hemlock, White pine and black cherry. The basswood and elm.are usually absent on the lighter, sandier hardwood soil.. According to the record of Land Economic Survey of Michigan, there is a considerable difference in the composition of the virgin and second growth stands. The virgin stands usually contain all of the Species mentioned above while the second growth stands are predominantly 19 maple (58). This condition no doubt is traceable to the maple's ability to reproduce freely from seed. Not too large an area is now occupied by the hardwood type and nearly all of it has been cut over. At the present time, the bulk of the hardwoods is limited to the loamy sand soils because the heavier loam soils that it once covered are now mostly cleared for farms. The condition of the present day hardwood type varies considerably. The virgin stands which remain are generally well stocked, but the second growth ranges from.excellent1y stocked to very poor. The variable condi- tion in the stocking of the second growth stands is due very largely to the occurrence or noneoccurrence of fire following the logging operation. The capacity of hardwoods to reproduce more prolifically than pine types has enabled them.to withstand to better advantage the destructive effect of logging and fires. The most profitable use of most of the timber produced in this type is for saw logs. Chemical wood is in denand in some sections for the small inferior material and the sound portion of large "cull" trees. Aspgn'tgpg. 0n the basis of the area occupied, the aspen type is third in importance to the oak and jack pine types. The type is comp posed of trembling aspen, large-toothed aspen and Balm of Gilead. Trembl- ing aspen is the major species and is found to some degree on all sites occupied by the type. Large toothed sepen is found to make up a signi- ficant portion of the stand on the dry sandy hills and on areas where the type is in association with the oaks. Balm of Gilead is the least imp portant species of the aspen type. It is found associated with the_treflbl- 20 ing aspen only on the more moist upland sites and the better drained swamp sites. Variations from the pure aspen stands, however, are com- mon. These variations include mixtures of the two aspens and the oaks, red maple, white birch and conifers. The condition of the type from a commercial standpoint is not very promising. like the oak type it reproduces easily from seed and sprouts, and is chiefly a replacement type after fires. The stocking is generally fair to good, but on the poorer sites the trees often rennin small and die at early age. Fungus and insect damage is rather common (102). The type on the more favorable sites does have some value, however, for pulpwood, excelsior and box boards. Probably its chief value at present is in providing game cover, in erosion prevention, and as a nurse crOp for more valuable SpeCies. The aSpen type occurs on nearly every class of soil and site from denuded hardwood and pine uplands to swamp landS. On most of the soils, it exists as a temporary cover, and is quite easily replaced by other Species in time (51). Usually on the hardwood sites, the area will eventually revert to the original growth if fires are kept othBS). Repeatedly, hardwood burned over areas generally restock with the aspen type because of its ability to reproduce from sprouts. However, the best stands of aspen are produced along stream'bottoms and on lowlands which are less frequently invaded by fire . Agpen and paper birch type. It occurs in small areas scattered throughout the region. This type is usually found on the moist sand, sandy loams and loams, although it may occur on practically all soils legend: .. ._..._— spruce +W Red Pine ...... Jack Pine ‘ ———-——- White Pine -———----- Oak-Hickery -———- N .Boundary of Beech-Maple Podzol ELI: Transition Soil Brown Forest Soil — Higgins Lake State Forest K. . . . . . .Kalkaska County 0. . . . . . .Crawford county M. . . . . . .nissaukee County R. . . . . . .Roscommon County Figure 2. lap of Lower Michigan showing Boundary Between the Beech- Kaple and Hemlock-White Pine-Northern Hardwoods Regions, and its Relation to tree ranges and Soil. (After Braun, 1950 and Veatch, 1928) 22 from the drier sands to mucks. It reaches its best development on the heavier soils. It occurs on nearly the same range of sites as the aspen type, with the exception that unite birch drOps out of the association on the drier, sandier soils. The poplar and white birch type makes a good cover or nurse crop for planted red or white pine if constant atten- tion is given to the removal of competing vegetation (75). The majority of the natural seeded young red and white pine is found growing under an over story of this type. Given fire protection, poplar and white birch reach mechantable size in a comparatively short period. It has further value for game cover. The present utilization of poplar is chiefly for excelsior bolts and to a great extent for pulp. The large size of white birch has an extensive use in novelty manufacturing. 6. History of Economic and Social Changes In lBhO the northern part of the southern peninsula of hichigan was divided into counties. At that tine the area now known as Crawford County was called ”Shawona", the name of a famous Chippewa chief. Later in 18h3, the name was changed to Crawford. In 1871 and 1875, Kalkaska and Roscommon counties were organized respectively. Settlement began in 1872. In the early days of this area's development, the luster in- dustry unquestionably brought the largest number of people into the counties. At the same time agriculture was also gaining a foothold so that the population on farms was being increased. Due to the fact that neW'lunber camps and the continued inflow of mill workers and woodsmen created a good.narket for farm produce, the agricultural population also increased rapidly. 23 After the lu bering operations started, the farmers did not depend entirely on agriculture for a livelihood. These pioneer families usually made their homes on the farm.for the entire year but the older boys and men worked in the logging camps during the winter and customarily Spent only the summer months at home clearing land and working on the farm. Hunting and trapping, cutting ties and henloek bark, and working in the wood-using industries were other sources of inco e for the pioneer fann settlers. The logging operations of the forwer decade had left freat areas of ad cut-over pine and some harduood land. Much of this had been burned over by forest fire. Farm settlement was not then being attracted to these cutover land. For larre area of this for or timber land, no promisingly profitable use seemed evident and tax.payments were being defaulted on such land by their owners. Around 1900, when most of the virgin tixher had been out, and the value of the lakes for resort and recrea ion purposes had not yet been generally realized, the county fund of all the counties which conprises the Richigan "pinery" were in a bad condition. Of course, the hard- pressed financial conditions were being felt more keenly by those counties which had the largest area of cutover pine lands. Beginning in 1902, various intensive real estate selling operations were carried on, various holdings fiere disposed of to land seekers and settlers from southern'flichigdn and neighboring states for poultry farms and general farms and ranches. host of these settlers have since left their farms and their lands have reverted to the state for delinquent taxes. 2A During all these years, federal and state land laws encouraged settlement, and numerous grants were made to open up the country and get land on the tax roll. However in northern Michigan, the tax base was shrinking and tax delinquency mounted. Here farms have not replaced forests to the extent as expected; in fact, by 1920 that part of the state had fewer farms than in 1910. There were large and increasing areas of cutover, burned-over, idle lands. The following table shows major changes in land use from l924ul939 in Roscommon county (2). Table 1. Major changes in land use from 1924-1939 in Roscommon County; Michigan. Type of land use 192L 1939 net change in area 4 - Forest 319,167 320,692 1525 - Farm. 15,529 11,939 - 3,590 Abandoned 3,865 5,305 1440 -- Other 35,hh3 36,068 625 - 7. Present Land Use The most recent development in this area has centered on resort and recreational property. During the years when travel was still mainly confined to the railroad and horse-drawn vehicles, access to lakes of the area was considered difficult so that their development languished. With the advent of improved roads and motor tranSportation, however, various lakes and surrounding areas began to attract more summer visitors each Year as hunting, fishing and outing grounds. The data which were obtained by Andrews and Uromley (2) in studying trends in land use in northern Michigan, show that shift from one land 25 use to another are constantly taking place; nevertheless, many settlers, particularly since the depression, are still trying to make a living on soils too poor for profitable agriculture. Their report reveals that the recent measures of fire protection have so greatly improved forest conditions that it is possible to expect the region to be able to produce considerable quantities of merchantable forest products in the near fu- ture. The data also indicate that recreation has increased so enormously as to constitute one of the.major forms of land use in the region. Con- sequently, little or no tax delinquency occurs in most of the area today; As far as timber resources are concerned, this region is by no means a worthless area as it is still generally regarded, but contains exten- sive areas of conifers and hardwood lands, and the latter, especially is valuable for farming purposes. Whether or not this conversion of forest land into farm may be less profitable to the state at large than the con- tinued production of timber will not affect the case. For when there are lands of such value for farming, they will be purchased and held for this purpose and the interest of the state will in the end have to be adapted to those of the individual, provided the land is being utilized properly. The public agencies, both through actual land ownership and through cooperation with private owners, have an important part to play in the future development. Adjustment in land use and improved resource management are still needed. 26 IV. THE O‘TUJl’ OF ‘Ji‘JCuianl‘IUNAL CU EMS l. Terminology A. Freguency. It has to do with homogeneity, the uniformity of dis- tribution of species throughout a community. It is the ratio of nunber of quadrats containing a given Species to number of quadrats surveyed, eXpressed as a percentage. The relative homogeneity of stands and come munities may be compared graphically by frequency diagrams, provided quadrats of the same size be employed. These diagrams show percentage of species belonging to each frequency class. The five frequency classes here used are as follows: Class A -- species in 0-20 percent of quadrats. Class B - species in 20—h0 percent of quadrats. Class C - Species in 40-60 percent of quadrats. Class D - Species in 60-80 percent of quadrats. Ulass E - species in 80-100 percent of quadrats. Quadrat frequency (Fq) is expressed as the percentage of quadrats in which (1) a given size class of a given species occur i.e. frequency per Size class per species Fan of species A g No. of quadrats in which size class n of‘A occur 1 100 Total number of quadrats examined (2) a given species occurs i.e. frequency per Species. F s of Species A 3 Nb. of Quadrats in which species A occur'x 100 q Total number of quadrats examined 27 (3) relative frequency'(Fr) -- a relative expression (as a percen— tage) of the frequency for a given Species as obtained by Fq and based on the total of the frequency values for all species Fr of Species A 3 Frequency of species A x.lOO Sum of frequency values for all Species B. Density. It is the number of individuals on a unit area basis. The nudber of individuals recorded from.the actual area surveyed were reduced to an acre or quadrat basis. (1) D of species A . Total no. of individuals of species A counted Total no. of quadrats examined. (2) Relative density (Dr) - a relative index.of plentifulness eXpressed as the percentage representation of: (a) Each species, based on the total number of individuals counted i.e. relative density per Species. Dr of species A . Total no. of individuals of species A counted.x 100 Total no. of individuals of all Species (b) Each size class of a species, based on the total no. of indivi- duals in that size class i.e. relative density per Size class per species Drn Of Species A 3 Total no. of size class n stems of species 1 100 Total no. of size class n stems of all Species (c) Each size class, based on the total number of individuals counted in all quadrats. D rc of size 2 a Total no. of individuals of size class 2 counted 1 100 Total no. of individuals of all size class C. Abundance. This term.is an appreciation of the relative number of individuals of each species entering into the constitution of the plant Population of the territory under study. A scale of five degree of abundance is suggested by draun-Blanquet (14). 28 Al. -— species very rare in the community. A2. -- species rare in the community. A3. - species not very abundant in the community. Ah. -— Species abundant in the community, and A5. - species very abundant in the community. The value of abundance estimates depends on the extent of area, richness and variability of the flora, and the ability and experience of the estimator. Abundance is also expressed in a more concrete way in this study in the folloWing manner for the tree Species. Abundance (A) - the average number of individuals of a tree species per quadrat considering only the quadrats in which the tree Species is represented A of Species 5 - Total no. of individuals of Species B counted No. of quadrats in which Species 5 occurs But 1) . Total individuals Total quadrats and F = Quadrats of occurrence x 100 q Total quadrats SOA-w q D. Cover. It deals With the surface covered by individuals of the herb and ground layer vegetation. The five-point scale of brauanlanquet was further subdivided. Class 1 was divided into two classes: class 0 representing Species with less than 1 percent coverage, and class 1, representing Species covering 1 to 5 percent of the surface. This proved very satisfactory Since such a large number of Species occurred in each 29 community with coverage less than 1 percent. The six point scale as used in this study follows: Class 0- species covering less than 1 percent of the ground surface. Class 1p- species covering 1 to 5 percent of the ground surface. Glass 2- species covering 5 to 25 percent of the ground surface. Class 3—- species covering 25 to 50 percent of the ground surface. Class h—- Species covering 50 to 75 percent of the ground surface. Class 5- species covering 75 to 100 percent of the ground surface. E. Easel area. It is a concept of foresters and is the total cross- sectional area in square feet of the stews of a tree species based on diameters at breast height. This is a convenient way to show physiolo— gical dominance of tree Species, since there is probably a close corre- lation'betwecn basal area and the surface or voluhe of tree crowns. Relative basal area (Br)- a relative index of dominance eXpressed as the percentage of basal area occupied by a given Species. Br of species A . Total 1:5.A. occupied by individuals of species 5 .x. JQQ Total B.A. occupied by all Species 2. Method of sampling Quantitative studies of plant communities yield valuable data rela- tive to the number, size and diapers 9a of community elements. The value of such data accrues in large part from.the statistically valid means they provide for comparison of plant communities in either Space or time, or in the concrete or abstract sense. Also, as a permanent record, they are subject to reconsideration by other investigators, who may reinter- pret them.in.the light of additional experience or information. 30 Usually the numbers of an entire plant cornunity cannot be counted or.measured, and even if this were done, the information would be no more useful or Significant than an adequate set of data acquired by proper sampling. However, prior to sampling, observation and reconnaissance of the community are of extreme importance in determining where, how, and what to Sample. The community should have been observed repeatedly in different parts of its range and more particularly under the varying local conditions where it exists. Finally, the Specific stand should be observed thoroughly to determine its Obvious variations, its extent, this together with the size of individual plants, the strata present, and the purposes for which the sampling is to be done, one may plan his procedure in terns of the desired results, the necessary degree of accuracy, and the time available for doing the work. This is the exact procedure this writer has carried out in this study. In this ecological study, the list—count quadrat method has been used. This method was subjected to some modification.depending‘upmn circumstances. For trees, the individual.diameters were also recorded and segregated into different size classes according to Weaver and Clements (9b) as the following: hedium.reproduction size class 2; more than 6.0 ft. high and up to 0.9 inch D.B.H. Large reproduction size class 3; 1.0 in. to 3.5 in. D.B.H. Small trees size class A; 3.6 in. to 9.5 in. D.B.H. Large trees size class 5; 9.6 in. or more D.B.H. 31 The basal areas (indicative of dominance) for tree Species were also computed. Increment borings also were taken from representative trees of various species to determine their approximate ages. The cover of ground vegetation were recorded by height class designated as: Class 1. Plants up to 0.9 foot tall. Class 2. Plants from 1 to 3 feet tall. Class 3. Plants over 3 feet and up to 6 feet tall. In this type of study, as one soon learns, the major concern is to get adequate data with a minimum of effort. With this object in mind, 16 milacre quadrats were used on grassland type sampling while 16 one-fifth acre square plots, with 93 feet on each side for sampling tree species were employed for the four different forest cover types. The location of such quadrats were randomly placed within the realm of the stands where the border effect of streams or ponds or highway would'be minimized. And smaller quadrats (one-tenth of an acre and one thousandth of an acre) for sampling subordinate strata were to be "nested" in the largest quadrat. All field data were recorded on "Field-data Quadrat" sheets, one such sheet being required for each quadrat. 32 4. «$2.33, hgtcsém .vo £03.69 A¢n¢_ £33m... >m+m$mxorsfiyou 313m 3: uct 04 wf mfsaxw uuagou 33m 9—3 9132.. yo as: .m 92.9“. wco,.+.£m mfisdm wwde meSoz _ M @ msuoiw \V\.I ”ESQ .2... Dummyfiocsom 55:38. IIIII 056 we? 5 E x + o B mwiducsom 3:300 l.l “VINQWJ 33 3. Observation Grassland type. The type of upland grass occupies those open burned wild lands on which the established cover includes the genera Andropggon, Pba, Danthonia,lggrosti§, Schizachne, gromus and Koeleria. One of the characteristic plants of this area is the sweet fern (Comptonia peregrine) located chiefly on the drier sandy pine plain where no type of forest cover has been established. The only true fern species that is really abundant and becomes a component of the ground cover is the common. brake, which forms large patches in the light soil of the plains. W. J. Beal (8) made statistical studies showing that 95 percent of the plants common in the "Jack pine" plains are perennials with deep roots or rootstocks adapted to severe conditions of drought or of surfaceéburn- ing. Nearly half of them are included in the four plant groups, namely, the Compositae, the Gramineae, the Rosaceae and the Ericaceae. In the spring, wild flowers such as one of the bluets (Egggtggig loggifolia), sand violet (Viola adunca), pink milkwort (W pglygama) are certain to attract the attention of the visitor to this reBion, and during the summer or early fall other species lend their color to the landscape, such as blazing star QL§§££i§_fl$gE!1§E§$1), Wild sunflower (Helianthus occidentalis), a golden rod (§gli§ggg,3pp.). In August especially) cudweed(Gnaphalium.Macgggéé) may be seen whiten- ing here and there. 34 Table 2. Data of the Vegetation in the milacre quadrats of the grassland type as recorded in percentage of coverage or number of plants by height classes. “‘ Station 1 Qiptribution of Coverage Classes for the m Location: SW£,Sec.18, Quadrat fi T26N, Rhw. ___ 1 2 3 g Height class in Feet Elnnistin_List. Obl 1:3 3-6 0-1 1-3 3-6 0-1 153 3-6 0-1 123 3-6 A ostis scabra 3 Arabia glabra - Arctostaphylos Uva- LII-Si - Artemisia caudata 1 Aster laevis O Browne Kalmii 1 Campanula rotundifolia - garex tonsa 3 Cirsium Hillii - Com onia erecrina 1 Convolvulus s ithamaeus 1 anthonia spicata 0 Erigeron annuus__ 1 Erigeron racemosa O Fraggria tirginiana 1 Gna alium Hacounii - flieracium aurantiacum 2 Hieracium venos_yx_n O Houstonia lo ifolia 1 Lgctuca canadensis O 0 0 O 1 1 1 2 1 1 0 O 0 O O IIIIIIII III-‘II IIIIIIIIIIII III-I'll IIIIIII-‘II I Lgatris hieuwlandii Lithosggrmum arvense salis sub labrata Po ala ama 29a compressa Prunus 22211; Prunus vir iniané Fteridium E'uilinum Rmbus idaggg anex‘noetosella §§1ixlnmfill$ Schizachne'“‘ asc gg Senecio Balsamitae Solida o canadensis Solidago Juncea Solids o racemosa Viola adunca ‘ II-‘Ol I IOI-‘I I IHIHHHWONOHHI I IN IIIIII-‘OHOIHI-‘I IOI—‘NIUIHHHIOW III—‘I IIIII 'ONIIOIIIIIIOIOIOIOHONINIIHININ IIIII IIIII-‘I I IIIII-‘I IIIIIIIIII-‘IIIIIIIIIIIIIIIIIIIIIIIIIII I IIIII HHOHOIIHHHI oooounHHmnn OHHHI * number of plantS. Table 3. Data of the vs the grassland type as recorded 1 coverage or number of plants by n percentage of height classes. 35 Station 2 location: SE£,Sec.ZL, T25N, R6W. Floristic List A ost' scabra Andro on sco rius Arabia glabra Artemisia caudata Aster laevis Cirsuum Hillii Comptonia peregrina Convolvulus s ithamaeus Danthonia s icata Eri eron annuus Erigeron ragga Fragaria'virginiana Gna lium.Macounii Hieracium aurantiacum. Hieracium venosum Koeleria cristata Iactuca canadensis Liatris Nieuwlandii Ph salis sub labrata Poe angustifolia Pb ala ama Prunus pgnsylvanica Prunus 1a Solids o canadensis Solidavo 'uncea Solida o racemosa Viola adunca fteridium a uilinum Rubus idaeus ex cetosella Salix humilis _—.‘ Distribution of Coverage Classes for the Type _‘¥ 0-1 1-3 3-6 0- IINHNHHIOIH IHHIIOOOIIIOIIOHIOII 1 I I I I I \DI I I I * number of Plants. Quadrat 2 3 Height Class in Feet IIIHNIHIIHO OIIOIOHII IHWIIOHOII IINIIIIII IOHIOOHIIIOIOHIIIOOHIIHINIOHIII IIIIIHIIIIHI IIIIIIIHIgIIIIIIIII IIHI IIHHIOOII§OOIIIIIIIHHOHHNII 4 IINIIIIIIII HIINII W __ 1 1-3 3-6 0-1 1-3 3-6 0-1 1-3 3-6___ 36 Table A. Data of the vegetation in the milacre quadrats of the grassland type as recorded in percentage of coverage or number of plants by height classes Station 3 Distribution of Coverage Classes for the Typew Location mi, SecJJI, Quadrat T26N, raw. 1 M 2 1 I. , Height Class in Feet Floristic List 0-1 1e3 3-6 0-1 1:3 3-6 0-1 1:3 3-6 0-1 1&3 3:9 Aggostis scabra Andropogon Gerardi_ Anemone cylindric§_ Antennaria neglecta Arctostapnylos Uva-ursi Artemgpia gaudata Asclepcias syriaca Aster laevis Campanula rotundifolia Comptonii Eeregrina Erigeron annuus Erigerggystrigosus Gnaphalium Macounii flglianthus'occidentalis Hieracium aurantiacumw—' Liatrisffiieuwlandii mmchia guadriflora Qenothera bienqig Oenothera_psrviflora Egg angustifolia ‘fi_ Polygala pglygama Prunus 23mila prus idaeus Hume: Acetosella éalééésa i.___uncea I I II—‘I-‘III—‘I IOI-‘OWI I I h)! I h‘C}! I I ADA) I I t I I Imoo§HIIIuo I II-‘II I IIOI I I I I I I I I I I I IF4 I I I I I I I I IOIOIIOINI I I IOIII II II I-‘I II II II II II I I I NOI—‘OOOOI I-‘OI OWOI I I I I I I I H H IIIOI I FJA>I I I H I I I I I I I I I—' I O * number of plants 37 Table 5. Data of the vegetation in the milacre quadrats of the grassland type as recorded in percentage of coverage or number of plants by height classes Station A Distribution of Coverage Classes for the Type Location NW%, Sec.10, Quadrat T26N, R3W. 1 2 v 3 A Height Class in feet Floristic List 19:1:1-3 346 0-1 123 3-6 0-1 1-3 3-6 0-1 113 3-6 Agrostis scabza Andropoggn'derardi Andropogpn scoparius 3 ~ - 3 - - 3 - - A ~ - Danthonia Spicata Schizachne_purpurascens Antennaria neglecta Artemisia_caudata Aster laevis Campenula rotundifolia Uomptonia peregrine Erigeron EEQEEE. Erigeron strigosus, gregaria gigginiana Gnaphalium Macounii flieracium venosum: Houstonia longifolia légtris Nieuwlandii Rubus idaeus Solidarro juncea Solidago nemoralig Solidagg'racemgsa I I I-‘ I I I I I I I I O I I I I I I OI I I OI I I IIHONII—‘I—‘O I I ILIJOI I I IMI I I lI—‘I I—' I I I OOI—‘HI I I OII-‘I-‘OI l I I I INIOHI-‘OI I I I I II-‘I-‘II-‘II I I I I I I OOI I I I I I I I I-4 I I 38 Aspen.type. In the thick pine country, where the lumberman's ax has let in the sunlight, different plants sprang up freely. The most fre- quent ones are pin-cherry and blackberry with two dominant Species of aspen, namely, Populus tremuloides and‘g. grandidentata, and they are also often supplied with red maple and red oak. Of the latter two spe— cies mentioned, red maple is most widely distributed in the ground cover, agreeing with its importance in the large-stem classes of the mature stand. The most abundant and common ground cover is Vaccinium.angusti- ggligm with 23233 spp. a close competitor. The bracken fern, Pteridium aguilinum which is abundant and charac- teristic of the aspen and jack pine forest, should be considered as a subdominant, since it may act as dominant both in invading new areas and in remaining as relics if the aspens is replaced, which is true in the case of station 17 in the conifer hardwood type forest. This particular species occurs in great abundance, often composing 50 percent of covering of the ground vegetation. It also has rhizomes about 8 to 10 inches below the surface of the ground, out of reach of fire. Their leaves spread out at a height of 10-20 inches in the sun, or in a shaded area as high as A feet. They cover the ground with a moderate shade, which, howa ever, does not preclude the development of various secondary plants such as blueberries. Although the shading effects of the wide fronds has been pointed out, yet the dominance of this plant shown in Table 8 can- not be taken as an average figure for all times, for at different periods during the year, this plant varies widely in its spread or coverage in the ordinary sense. 39 Tahlo Q, Summarized Field Data of the Four l/S icre ,uadrats of the aspen Type by size Classes . . . ‘y . , '11 \JL- 1 .m u.) ll ' T OT‘ALLS i p _ . fl , ._L S . Freq. Dens. ‘Abun— ; Basal Area ' Freq. Dans. Freq. Jens. rreq.: Jens. m I, ~ 1 . ,4 r , . ,,I . »; 2 , ‘ ' . ., ; V, w ‘ 1% 161. { /JCT6 39, p Le. x 10. p 10. p 10. n 10. p , No. 3 No. x '“ = . ' ‘ 1 1 L Species 2 Populus Sflfldidentata - _ _ _ : Po§515§‘_"—‘-"‘ tremuloid°s 1 95 5 3-51 2 50 22 0.07 2 50 as 21.21 1 21 . 1.«) 2 to 9.53 117 29.25 13.11 V8.VO 10 078 19.77 21.975 59.000 50.01 71.505 \"7 L) k L H H k ) xi \ a C \ ') L— C\ I \ 1 \‘1 I\) \fL O bJ C3 C3 C\ . b.) ("x \A) K] \‘1 C\ H C r O \R l\) C) 0 |-’ 0“ CO H I O\ O\ 1111 . 28 2115' /O/ rubrum 2 50 37 20.2 2 50 102 50.02 3 75 21 5.78 _ _ _ _ 3 75 11.28 220 55.00 25.51 73.33 0.000 0.20 8.250 Quereus mll,_ ( ‘EESEE 1 25 30 21.2 2 50 31 10.02 2 50 23 0.33 3 7g ‘.70 87 21.75 10 00 29 00 1.333 1.08 5.115 tula . £221£2£233 l 25 l '70 l 25 23 7'1: l 25 29 7'98 - — - 1 2: 1.76 53 13.25 0.09 53.00 0.010 S Tinus bawksiana 1 25 1 .70 1 25 g 2,g3 l 25 2h 6.61. l Finns resinosa - - - _ 1 25 1 .31 _ _ _ _ . 1 25 ’ Quercus . alba l 25 7 1.90 2 50 10 3.12 2 10 0 2,20 1 Irunus . serotina lflolnrchier ca‘adensis 2 SO 11 7,30 2 ; Ylmus anericana I‘O \"L 1.) CI\ O 5:0 H F0 U1 h.76 36 9.00 h.l3 36.00 7.531 7.12 9.h75 T ~J ;\ H I ro \J’L I l—l H H O O H O\ E) H C O f\) O " J \ m H {‘3 Q "J .4 I\) \.'l O “ 3 . \ ,‘L b.) {‘0 G\ C\ e \."‘L O F0 0 \G ('1) H b.) H e \3 if.“ I'\) I-" O C") [‘J N O 52' PO \I‘L r O \ I O 6 3 \i ‘% H {‘0 \fl b.) . 3 W H l‘ 3 \TL H U1. 0.38 _ _ _ _ 2 50 9,53 52 .13 .97. .26 .161 .15 .300 p .111 .13' .180 .10? .01 .010 .01 .058 L.) ("\ \. O ( k I I I I I I I I N \1 O ‘0 C \ "L W N .4 (j‘x C \3 U1 \JJ 0 H O H Lo 0 \fl 0 ‘_J r J \fl l—J O .\1 O H I D UL K») O ‘ U k.) I I I I I I H {\x ‘I r. d? L" H O O 0 he b ,,.... , , .A Fulmmh 111 10.21 320 30.00 1 1 ‘ 0“ 11.02 ; i 11 $.07 525 100 100 100 100.117 132.053 I v ’4 "W”; ._.._ S Dons. No . ,3: A n r} 1’ f 9".) UK) 9 J 2 1.51 1 / Q) J v o I 1 2.27 Rh 5.07 n) “J “J". ‘31 P) \fl 111113 Dens. flIS "J 2h5 01.25 117 2?.25 220 55.0 100 Dr C-) 13.hh 100 Abun- dance Basal Area : P ~ gel. /icre ‘ I ‘ TI"\_ 10 (~70 1' '7 oh 076 l .520/ U o. '— o/ 2 6.690 1.912 .lCl .lhh 10§.hh7 .9 1 . "__.-. 11008 Sch E" 5.02 7.550 (‘x O to O ( t‘ O r .) I-' U1 0 3373.;fl‘73 1“. A_ c. _. ..r.:m"’2;" 7.1: 9.115 .01 .020 1.82 ‘2.125 I I .15 .200 " 3 .13 .130 .01 .058 32.033 1. -. ..3 bvvowow' M. q Hex F. ‘1 i /’ 40 Figure 4. Phytographs of Important Species in Aspen Type V.x - \- Populus tremuloides gear rubrum O O Betula Le epxgifera Finns banksiana Legend: ODr-aRelative density in percent OFq--Qnadrat frequency in percent oSc--Size Classes in percent OBr--Relative basal area in percent 41 Table 7 and 8. Data of the vegetation in the milacre quadrats of the aspen type as recorded in percentage of coverage or number of plants by height classes Station 5‘ Distribution of Coverage— Classes for the Typew location SE2, 5ec.9, Quadrat T25N, RESIN. __ 1 2 3 5 Height Class in Feet Floristic List 0-1 1-3 3—6 0-1 1-3 3-6 0-1 1-3 3-6 0-1 1:} 3-6 Ace; rubrum 4* - - 12* - - 1* - - 3* - - Apocynum a‘ndrosaemifolium - _ - 1 - - Carex.gpp. l — — O - ~ 1 ~ — 1 - - Qanthonia spicata l - - O - - — - _ - _ - Qgpltheria'procumben§_ - - - O - - l - - O - - Ehpulus grandidentata' 8* - - 17* - - 13% - - 5% - - Prunus pensylvanica_ 11* - - 11% - - - - _ 1* - - _P_teridium aguilinug 2 - - 2 - - 2 - - 2 - - _Qgercus alba 2%- - .. .- - - - - _ - _ - fie reps gubra 8* - -. ‘. .. .. 1e:- - - 1e; - .. Rubus 322. 0 - - 1 - - 1 - - o - _ Smilagina stellata' - - - O - - 1 ~ - l - - Wings; MEREEEBE - - - 2 - - - - - 2 - - ““ .1. .1112... 0:11.31". "m”- "m"- Station é .fi_ Distribution of Coverage Classes for the Type Location 53%, 580.7, h..._* Quadrat T26N, Raw. l 2 “.3. M--._.-_....._-,A,...._ Height Class in Feet Floristic List «__‘ 011 173 3-6 0-1 1-3 3-6 0-1 1-3 3-6 0-1 123 3-6 Amelanchier 3gpadensis - - - - - - 1‘7‘~ - - - - - Apocygum androsaemifolium - _ _ - - - 1 - - - - - Aster laevis 1 - - 1 - - o - - - - - Carer E22- 3 - - 3 - _ 2 - _ - - - girsium Hillii, - - - 0 - - O - - - - - Comptoni pgregrina 2 - - 2 - - 2 ' ' ' ' ' Danthonia Spicata l - - - - - 1 ' ' ‘ : : Erigeron annuus - - — l - - 0 ~ - - Eggltheria procumbeng' - ~ - - - - 1 ' ‘ ‘ ‘ : Hieracium venosum 0 - - - - - 0 ‘ “ : : _ Fanicum‘égribneriannm, - - - l - ‘ ‘ ‘ ' Prunus‘pgnsylvanica 1* - - - - ‘ ' ’ ‘ ’ ' Prunus serotina 6* - - 1* ~ - ‘ - - : : : lgteridiggiaguilinum Qgercus rubra 5% - - Rnbusigng, 2 - - Egpcinium angustifOlium l - - _._l 3.: Ahhbfi‘l a l bayou A>hfu I e number of plants #2 Table 9. Data of the vegetation in the.mi1acre quadrats of the aspen type as recorded in percentage of coverage or number of plants by height classes Station 7 Distribution of Coverage Classes for the Type location Sflfi, Sec.31, Quadrat T2411, RBW. =1 1 2 j 4 Height Glass in Feet Floristic gist __ 0-111-3 3-6 o-_1_ 1-3 3-6 0.1 1-3 3-6 0:1 1.3 1.6 £2.93 &w 1% .. .. 5-2:- . .. (3-2:- .. .. 2-)!- .. .. finelanchier_canadensis - — - 3* - - 2* - - 2* — - Aster cordifolius 1 - - O — - O - — 0 - - Danthonia spicata - - - O - - O - - 1 - - Gaultheria procumbens - - - 1 - - l - - - - - Lonicera canadensis - - - - — - O - - — - - Honotrgpa fixpopitfiES - - ~ 1 - - l - - l - - Erunus‘pensylvanica 1% - — 2% - - 1% - - 2a - - Pteridium_aguilinum l 3 - 2 2 - 1 3 l 1 2 - Quercus alba - — - 2* - - - — - - - - gmercus rubra 2* - - 1* - - 2* - - 2* — - Rubus £22. ‘— l - - - - - l - - l - - Smilacina stellata_ - - — 1 - - - - - - - - Solidago nemoralis - - - l - - O - - — - - Maccinium,myrt;115ide§» 1 - - l - - l - - - - - - - - _ - - 1 - - Viburnum acerifoliug '3‘ number of plants In station 8 of this type, which was situated in a transitional zone of well- and poorly-drained land, birch has become such a dominant species that is quite striking. Also in this particular sampling plot, the number of herbaceous Species has greatly increased due to the more favorable moi- sture conditions. species like Galium, gguisetum which never appeared in other stands showed up in the tally. A3 Table 10. Data of the vegetation in the milacre quadrats of the aSpen type as recorded in percentage of coverage or number of plants by height classes Station 8 Distribution of Coverage C1a::es for the fype Location NEfi, Sec.36, Quadratr ‘ T23N, H3W. , 1 2 3 h Height Class in Feet Floristic List 0-1 1:3 3-6 Obl 1:3 3-6 0-1 1~3 3-6 0-1 1-3 3-6 Acer rubrum 13W - 138T — 50 ~ - 9W W - Achilles Millefolium - - ~ - - - - Amelachier canadensis ' — 1* - - 1* - - Antennaria neglects - - - Astergsagittifolius Athyriim‘. Filix—femina Botrychigm‘dissectum Care: spp. Cgptis groenlandica Cornus E22- Uiervilla Lonicera Fragaria_virginiana Galium triflorum Hieracium venosum Iactuca ERR- ;onicera canadensis Iytopodium.9bscurum Unocleaeggnsibilis ngulus grandidentata Prenanthes alba Eyrola rotundifolia RUbUS S omilacina stellata solidagg canadensis §91idqgggcurtisii Trifolium dubiu_1_1 913112;) '1 1eripana Viburnum acerifoliun Vaccinium angustifolium Viola blanda I I I I 11a<3<§1 1 I .1:- I I A)F1l I I #11 1 \ol 1 1 1 F'FJI p41 kn 1 IOOOHI 11 11 10101 1 1 I IHOOHHI 1 1 1 1 1 1 I I I I C>C>C)C)I I I I I I F’CJCDFJI I I I C) I I I I l I I I I I I I I I I I C) I I I I F‘I I HIHIOOOHHI 11 11 00 1001 HH1g101HHH01 11 11 IOIOHOI I I my“ wen-on- 1......“ ., .., . .‘ . -fl._-._.__-_._*~ - . .% number of plants Northern pin oak type. Five distinct tree species occupying con- siderable area in this type of plant community. Three of these species belong to the Fagaceae; namely, Quercus alga, gmercus rubra and.Q. ellipsoidalis. The other two are Populus grandidentata and Acer rubrum. Cutting, fire, and grazing of the past are responsiblefor the generally depreciated stand composition, understocking, and large amount of defect in most of the oak stands. This type generally occupies certain red pine and white pine sites that have deteriorated severely as a result of at least two or three fires. Many investigators (62,95,31) have com- pared this type to the aspen type in that it is a temporary type into which the original occupants of the site gradually encroach. However, this reversion to the original type of tree growth seems to have to wait for a longwhile before it becomes a reality, as Judged from the quantitative data obtained from the area. First of all, there are not enough reproduction or parent seed trees to provide a satisfactory nucleus for a pine forest. Secondly, the fact that twentyaseven percent of the tree species are in size class two; thirtybfour percent in size class three; thirty-siILpercent in size class four; and only 2.6 percent in size class five, definitely suggests the young character of this so called temporary oak type forest. Six tree species make up the understory With Amelanchier canadengég appearing more frequently. Advance reproduction of oaks in unmanaged stands is often sparse. Such is the case in the quadrats studied in this region, since oak seed trees must be reasonably close together and uniformly distributed because the seed is disseminated only a short distance. Table 11. Qucrcus ellipsoidalis QEEFEI§__"'"— _;1hra Eucrcus alha Fonulus frandidentata “kW-g ruFrum Fetula papyrifera jfiflanchicr caqadcnsis Frunus gerotina yamanelis Viriiniaqa Erunus pensylvanica PM 1- ,. ‘ Via 1.516 .hs :ravis Corpus SHL Di: i .1118 Banksjana Surnr the x "[1 urea. , v {V’ 7 N Pb.) I\) VT if \m \‘a (A) -\1 \fl .4 UL H O n‘ 0 \FL P.) \ '3 in l") \[1 \ w: -iZC e h) finrthern d field lin a Dens. H0. fl r r) (3 .Cu 1 ’0 . .4] b 16.75 1 .h” C 37.32 6 :7} 995 Data 0: the fiour 2:1: 133»? lg’mjizc 3T4; Freq. Hens. To. x No. " 2 50 “O 7.78 3 73 31 l".OQ 257 3h.31 (‘1 . x 4L) in Fre No. L 4 b.) —n \ l/S icre iuadrnts Slass>:. ‘3 (A 1.4, I Dans r $T ' ' L HO 9 w 5 r 0 30.0 3 MO.D 6 30.0 _ _ _ _ _ _ _ - _ _ - _ _ _ _ _ _ - O "V U 0 \JJ Pb.) tr 14 k» 14 l4 *1 [‘3 \JI \,"L l\) VI 100 f \3 UT [\3 f\) \J‘L \f‘L 7‘) U1 10.3h 121 ”0.25 :.h, 2 3.50 0.3h 71 7.75 3.K§ l 0.2fl 13.79 8 20.99 3.h§ 6 x A) I L7 \FL 0\ 0.25 kg P [\D O o ‘51 P.) ”l? . lV/.‘s/\J on (‘0 L_\/ n , ,1 45 Basal Area 2le 11.95h 19.735 3.an .116 .297 .110 lie]. . /1‘LCI‘C 100 85.583 7 TOTALS 7 Freq. 7 Jens. ‘ Vibun- ;:‘S . 7:, F; F1 Ts D Dr dance 9.61 18 100 13.80 72 18 t u: ~ 7 :0 6.89 6h 16 8.55 32 20.52 LS Basal .‘.rea Ftl Lei. /ncr’ 1 I 20.h59 29.? '$ 110797,, ~;‘ ‘ \" #6 Figure 5. Phytographs of Important Species in Northern Pin Oak Type f&‘ 4 r \\\ vi—‘r .‘ / FQ \ I \ X 1. 73c. .Quercug_ellipsoidaligl lgggrcug rubra Querggg’alba ggmmgg§_grandidentata Acer rubrum Legend: 0Dr--Relative density in percent OFq-—Quadrat frequency in percent OSc--Si2e classes in percent oBr—-Relative basal area in percent A7 Table 12 and 13. Data of the vegetation in the milacre quadrats of northern pin oak type as recorded in percentages of coverage or number of plants by height classes gtation 9 Distribution of Coverage Classes for the Type— Location swi, Sec.25, Quadrat T25N, MW. l"w _ 2 3 1+ Height Class in Feet Floristic List -- 0-1 1-3 3-6 0-1 1-3 3-6 0-1 1-3 3-6 0-1 1-3 3-6 Acer rubrum 1* .. - 1* .. _ _. _ _ __ _ _ Amelanchier canadensis 1* - - - - - 11%:- - - .. .. .. Apocynum androsaemifolium l - - l - - - - - - - - Aster yievis l - - l - - l - - — _ .. Aster sagittifolius - - - - - -- 0 - - .. .. .. (Tarex spp. 2 - - 2 - - 1 .. .. - .. .. Danthonia spicata - - — 1 - — .. — .. .. .. - Hmnelis Virginians 8* 3* 2n 6-:- [1* 3 c 5%:- [1K 11".- .. _ .. Ionicere oblongifolia 1 - - - - - - -. - .. .. .. Pteridium aquilinum 3 - - 2 - - 3 — - .. .. .. Quercus rubra 1* - - 1* - - - - - .- - - gubus allegheniensis - - - - - - 1 - - - - - Solidago nemoralis - - - O - - - - - - - - Vaccfii’ um angustifolium 2 - - 2 - - 2 - - — - - * number of plants §tation 10 Distribution of Coverage Classes for the Type Location NW2‘;, Sec.l1+, Quadrat T2611 , RAW. 1 2 3 1+ __ Height Class in Feet ’ , Floristic List 0-1 1-3 3-6 0-1 1-3 3-6 0-1 1:3 3-0 0-1 1-3 3-6 Acer rubrum 1;):- - .. 3* .. .. 3* .. - 5* .. .. anZx app. 1 - - 1 .. .. 1 _ .. 1 .. .. Comptonia peregrine. .. - .. - .. .. 1" .. - ]_‘L .. .. nglus americana 1* 13:- 1e:- .. _. .. 1..- .. .. 3.. _ .. Hamamelis Virginians. - - .. 1* .. 1* 1* .. - .. .. - Hieracium venosum - - - - - - - - - O - Gaultheria procumbens l - - 1 "' " 1 " " :1, ' '- Prurrus Ensflvanica - - - - - — - : - 1 - Pteridium aquilinum l 2 - 2 2 - l a - l _ _ gaggle alba 3* 2 - 2* " " -.. - - 5* _ _ Quercus rubre .. .. .. 1-x- .. - 3.. .. .. - - - Smilacinewfistellata - - ‘ 0 ’ " - - : 0 - .. Sohdangemoraus - - - 6 ' " 3'- : - 1 _ _ Vaccinium ingstifoliugg - - - -* “‘w * number of plants 1+8 Iable 1h. uata of the vegetation in the milacre quadrats of northern pin oak type as recorded in percentages of coverage or number oi plants oy height classes Station ii Distribution of Coverage Classes for tne Iypp Location dEg, 500.16, .—’—' Quadrat - 126:3, Ids“. .._. 1 2 3 1; Height Class in feet Floristic LiSt 0-1 193 3-6 0-1 1:3 3-6 0-1 11§_3-6 O-l le3 5-6 Acer rubrum 11* _ - 21* 1a - 55a 2* - 18* 2* - Agropyron trachycaulum - - - ~ - - - - - 1 - - Amelanchier laevis 03* - - 1x - - 17s _ _ 22% - - Aster ciliolatus l - - l - - - - - - ~ - Ester laevis - - - 0 - - l - - o - - Carex spp. 1 - - 1 - _ 1 - - 2 - - ggnvolvulus_§pithanaeu§ — - - O — - - - - - — - thaegufipp. 1% 1* - — - - .. - - 1s .. .. fipigaea repens - - — 1 — - - - - - - - Fragaria Virginians 1 - - O - — - - - 0 ~ - Gaultheria Lgcumbens 2 - - 2 - - 2 - - 1 .. .. 'nnaea borealis 0 ~ - - - - 1 — - - - _ Lonicera oblongifolia l - - 2 - - 2 - - l - - Melangyrum_ lineare - ~ - U - - - ~ — - — - nonarda Iistuiosa - - — — - - - - - - 1 1 Populus'gragdidentata - - - - 1 1* - - - _ - - Prenanthes alba 2* - — - - - - - _ - _ - Prunus pensylvanica 1* - - - - - 1* - - - - - Pteridium__guilinum - 3 - - 3 - 0 3 - o 4 - 01a minor - - - 1 - — — - _ - _ - gyrola rotundifolia - - - l - - O - - - - - Quercus alba 2n - - - — - - — - _ - - Quercus ellipsoidalis - - - 1w - - 1w - - 1w - - Rubus allegheniensis 2 - - 2 l - l l - - - - Smilacina stellata‘fi' 1 ~ - O - - O - - l — - Egalictrun_divicum l - - - - - - - - - - - yggcinium angustifolium 1 - - l - - 2 - - 2 - - A r number 3? plants Again on the ground cover layer, bracken fern maintains its dominance over other herbaceous plants. Although big tree relics can hardly be found, yet numerous ground plants which remain as relics are still plentiful, such as Eacciniug angustifolium,Gaultherie‘procumbens, Aster £22. and Uarax'gpp.. 1+9 Table 15. Data of the vegetation in the milacre quadrats of the northern pin oak type as recorded in percentages of coverage or number of plants by height classes Station 12 Distribution of Coverage Classes for the type Location dEfi, See.lh, Quadrat T2611, 33w. 1 2 3 ..- b, Height Class in Feet Flori stic LiS‘b 9:11 .1173- 316*9-111 }:2 .2‘6 9:]. 1‘3 3-6 0-1 1'3 1:2 - - _ _ _ 1* _ _ - - 1* _ - s - _ Acer rubrum - - - Amelanchier canadensis — - - fintennaria neglecta l - - gpocynumLandrosaemifqligm 2* 1* - Arctostaphylos Egg-ursi Aster cordifolius Asterilagxig Carex‘gpp. Comptoniegperegrina Crata?52§.5£§!l£ Euonxmgg spp. gregaria_girginiana Gaylussacia baccata Gaultheria_pgpcumbens Eagamelis virginiana Hieracium venosug_ ngzopsis asperifolia Phlox‘pilosa Prunus serdtina Pteridium aquilinum .Quercus alba Quercus ellipsoidalis Rubus canadensis Solidago nemoralis Vaccinium angustifolium Viburnum;_acerifolium Xiola adunca I I I II II I I I I II II II II II II II II I I II-‘I-‘OOI-‘O I I II-‘OII-‘NOI II‘JNI I I N I N I I I II—‘OI M I I I I O I I I E' I: I I I AJCDI I I I I INO@OI I I I I INI I I I I IONI I I INWI I .1. II-‘I-‘I I I I '3! l I I I-' 3‘: h) b I I \uI can C?CDI I I i~* u '. I I I I \nI-l'I-d I I II-‘OIIBI—‘I-‘I I I l-' I I I I II-‘MI I I O I I I I I I I I w .fi" - D--- c “”7 ’vficfl— thdéfiber. of plant 5 50 Jack pine type. By the time the field has been abandoned for some years, the forest has encroached considerably on the area occupied by various herbaceous growth. Jack pine is the doninant member of the flora and poplar and oak are also present. In places, a thick growth of jack pine appears and there are but a few young oaks growing among them. This is the general picture of invasion of different plant conmunity which is well depicted by the data on station 11 on the pine type study. PhytOgraphs and summary of field data show definitely the dominance of the jack pine, as indicated by the basal area and other qualities are concerned. The open crown of the pine is reflected in a rather rich ground cover of herbs and the scarcity of seedlings of trees (Table 16,17,18,19). In quadrat 14 and 15 of all the Species present in the herbaceous layer, seedlings of the dominants in the arboreal layer were poorly represented, accentuating a decided intolerance of these conifers. It is, therefore, very evident that tolerant broadleaved Species could easily crowd out the pines if climate were favorable to their invasion. This is so gra- phically eXpressed in the pollen profiles of some Michigan bogs.(73). Though the numbers of species appearing in the ground cover is large, frequency is low for most of them. This is, however, what one would eXpect according to Raunkiaer's law of frequency (71). One of the most significant fact observed in station 13 is the apparent ability of the conifers to reproduce and develop well under the parent group, evidently acquiring more light than.many of the herbs and a a few of the broadleaved tree species received. This agrees with the finding of Shirley (80) who reports the same competition for pine seedl- ings by aspen and bracken fern. ...... in; Quercns ruhra Populus tremuloidgs M Quercus ellipsoidalis Finns rcsinosa Irunus serotina ——-_..~__.__ Amelanchie Canadonsi I I Quercrs alba l ryzpym - T m J. DJ. 1.1.“) $4: \flw‘i'eirtha W m ummarizrd he Jack line ’D _ Freq. Dene. Y V no. , .o. , ,QO 2 W0 20o J r 2 5“ 1d ’1 r/ m a; V 50 PL >fi o _‘/ / '5: fear l/S Xcre wuwdrnts of s ‘ classes. 1+ .u9 H ro \W |'\) I‘O ‘Q‘L F C) \1 \3'1 [‘3 ibun— dance 131.5 7.33 Lo 3‘ 66.132 Basal Area. 2 i Jel. £99.20 ‘\ lOO 82.662 51 52 Figure 6. Phytographs of Important Species in Jack Pine Type S Pinus banisiana Qggrcus rubra Eopulughtrgmulqides D Q Pinus resinosa Prunus serotina Legend: 0Dr--Relative density in percent OFq—-Quadrat frequency in percent OSc~-Size classses in percent oBr--Belative basal area in percent 53 Table 17. Data of the vegetation in the milacre quadrats of the jack pine type as recorded in percentage of coverage or number of plants by height classes Station 13 ' Distribution of Coverag§“Ei§§§€§”f6E“£Be Type—“- Location SEfi, Sec.18, . Quadrat - T26N, Raw. 1 2 3 A Height Class in Feet Floristic List 0-1 1-3 3-6 0-1 113 3-6 0-1 1-3 3-6 0-1 1-3 3-6 Andropogon Geraggi. l - - Andropogon scgparius 2 - - gpccynum androsaenifolium - - - Artemisia caudata fi——'O - - Aster laevis 2* - - Aster sagittifolius - - - Campgnulalggtundifolia Egrex1pggnylvanica Cirgium Hillii ngptonia peregrine Qanthoniagpicata gubus hippidusifl_ Vaccinium angustifolium 2101a adunca - - - > I I \n )I; I I I I I 3 I I I I ’3 I I I I I II-‘I-‘WIOI I I J:— a I I -__ H -.-—- H' --—n—.~- -. M .0-0 .— con-m- "‘ "" v -' * numberfofuplanbs The data show that Pinus banksiana is by far the most dominant species With a frequency of 100, a density percentage of 92, and making up 99 per- cent of the total basal area. The tree species with second rank is Qgercus EEEEQ: frequency 75, but with a rather low density percentage of 2.9, and the total basal area per acre is negligible. On the basis of size-class representation for all tree Species recorded on this forest type, those of size class 2,3 and L are evenly distributed. Furthermore, advanced young growth of red pine has appeared in some plots, adds the necessary elements to suggest that the coniferous species have gradually returned to their sites after lumbering and subsequent fires. 54 Table 18. Data of the vegetation in the milacre quadrats of the jack pine type as recorded in percentage of coverage or number of plants by height classes Station It; Location SE}; , Sec . l7 , T26N, RAW. Floristic List Amelanchier humi; j 5 Andropogon scoparius Antennaria neglecta gpocymim androsaeudfolium a,“ DOOO‘ -.. ARtOStaEhylos Uva-ursi Aster laevis" '""‘ ”'“" _C_J';rsium Hillii 99mptonia Beregrina Convolvulus spithamaeus fitaegus Dodgei Danthonia spicata Epi 3. aea Yepens Fragaria virginiana Gaultheria Lrocumbens fleliav'dlms occidentalis Hieracium lenosum Liganthaiwn canadensc‘_ @ampymTlineare flgnopanthus mucromta Prunus Ensylvanica Prunus Ella Pteridium aouilinum Rubus gnaw Rubus canadensis Smilacina stellata Solidago 3W chcinium angustifolium Vaccinium vaczillag n—‘k 1 Quad rat 2 3 Distribution of Coverage Classes for the 13336 6* 1 11* 2* 3 i:- 17‘:- 1+ 3 '31- 20* Height Class in Feet 0.1 1-3 3-6 0-1 1-3 3-6 0-1 1-3 3-6 -~-- .—. -—‘- .. v 1 .. .. .. 15: - - .. 43* .. - _ 7-:- - .. .. 2-;:— .. - .. 1-x— - _ - B '7‘: - - - 1 - - number Br plants h 0-1 1-3 3-6 1 - .. 10-3:- .. - 21‘!- - - O .- - 15* - - 30* - — l-X- .. .. 11: - - 1* _ .. o - .. 3-:- .. - 2 - .. 2-3: _ .. The principal Species forming the ground cover are Pteridium Equilinmn , Em Beregrina, Rubus spp., Epigaea remns and _Ggultheria procumbens. The higher shrub layer is well develOped and is composed 0f 92332592 REESE; flelanchier spp. and two Species of the genus Vacciniun}. % Table 19. Data of vegetation in the milacre quadrats of the jack pine type as recorded in percentages of coverage or number of plants by height classes Station 15 Distribution of Coverage classes for the Type Location NW , Sec . 12, Quadrat T2 N, RSW. 1 2 3 h Height Class in Feet Floristic List 0-1 1-3 3-6 0—1 1-1 3-6 0-1 1-3 3-6 0-1 151-6 m trachlcaulxm Andropggon sceparius Antennaria neglects Apogynum androsaemifolium Arctostaphylos Uva-ursi Aster laevis _, Cigium H1111; Danthonia spicata Qiervilla lonicere Epigaea repens Erigeron annuus Frageria Virginians lgnicera'oblongifolia W ”Mme Pteridium.aquilinum. Prunus pensylvanica Salix humilig Solidago nemoralis ggilacina stellata II solidago Juncea Vaccinium vacillans Viola adunca OII-lI-‘I II-‘I" I I OIOOI II-‘H I I II-‘OIH II-‘OOO I I I I NII-‘IO H OIOOI I I “z I I IINI I l 9 I I I I HHOIObIHHoo o HOHIHHOH I I INIOOI I IIII INIIOI I I I l INOOOI I I * number of plants % Table 20. Data of the vegetation in the milacre quadrats of the jaCk pine type as recorded in percentages of coverage or number of plants by height classes Station lE' Distribution of Coverage classes for the Type location NEfi,Sec.36, QuAdrat T25N, Raw. 1 2 3 4 Height Class in Feet Floristic List 0—1 193 3—6 0-1 123 3—6 0—1 1-3 3-6 0-1 1-3 3'6. Amelanchigr,laevis - - - — - - - 1* - - - - Andropogon scqparius Danthonia spicata h - - 3 ' - Pea compressa Antennaria neglects 0 - - - - - Apocynum androsaemifoliump - - wemisia caudata 0 - - Arctostaphylos Uva-ursi Asclepias syriaca Aster laevis Cirsium Hillij Comptonia paragrina Hieracium aurantiacum Hieracium venosum. Pteridium aquilinum Pterospora andromedea Rubus canadensis Salix humilis Solidggo nemoralis Vaccinium angustifolium Viola adunca' w I I I:- I I E? I I I I I I II-‘OII-‘I I I II-‘OOOI-‘I-‘I WII III IOI NII III I I IOHHHHHHONHII I I owogn I I I I ONHI I I OHHI I I I I ooowHHIoooan I l * number of plants 57 Northern-hardwood-hemlock type. Conifer-hardwood group, generally includes the northern hardwood type, characteristically composed of sugar maple, yellow birch, basswood, beech, and hemlock, occurs princi- pally on loamy upland soils. The oldgrovrth stands, contain trees rang- ing in age from 160 to 300 years or more, in diameter from 18 to 30 inches and second-growth sawtimber forests, in height from 80 to 100 feet. Commonly residual stands from heavy partial logging, often include a disproportionate share of inferior Species and cull trees. The pole stands and sapling> areas, likewise, contain scattered saw-timber trees left at the time of logging, but are made up mainly of young trees of more or less even age. The four stands included in the study of conifer-hardwood type were located where no recent-burning evidence could be found, and were dis- tributed over a wider area than the pine forests; they also involved more stages of development than other type of forests, making possible observations frozr: early youth to decadent maturity. Representative of the typical mature stand was Station 20, a decadent stand which is situated on hi 18 to Rosconmon. Incidentally, the stand is also a representative of a forest after some disturbance by cutting; and the one at Station 17 pictures secondary succession from complete denudation of a former hardwood stand. The youthful character of the stand of maple-beech forest in the station nurber 19, is expressed by the small diameter of the stems and also by the large number of stems per unit area. These features are shown in Table 20. Though the complex of species playing a p'Irt in the crown cover 58 is not so simple as in the pine forests, it is still very simple when compared with the southern nixed hardWood community, such as found in southernLhichigan and Indiana, where up to 40 Species play a part. In the hardwood type, three layers can be recognized: (1) tree layer, usually consisting of superior and inferior strata; (2) shrub layer, consisting of shrubs and tree saplings; (3) herb layer, composed of herbs, pteridophytes, seedlings of trees and shrubs. On the Whole, there is sone clear cut separation of layers but sometimes it is rather overlapping. In general, the canopy is quite irregular, since it represents an uneveI-aged nixed stand of trees. A few of the larger trees, occasional— ly reaching a great height, at nd above the superior layer as "doxinentsu he tree species that are ;ost co;ron, and present in rest of the stands are Fagusggrandifolia, Acer EEEELYEEE: and £99?.ENP£EE- Other Species of tall trees at times boco e codoninant in the crown cover are Populus grandidentata, Tsuwa canadensis, but their distribution is scattered and erratic. Under the Open crown cover of the mature stands, there are some shrubs and small tree strata devoloyed. The tree species present at all regularly is Aoer rubrum and it oztnufibers all other species in repro— duction. This could be an indication that the original composition of the stand had been disturbed and recession of the climax species thus resulted. In Station 19 where young trees over 9 feet in height, but under one inch in diameter, size class two, and also in l—3.5 inch class, Size class three, were tabulated, Acer saccharum had a greater represen- 59 tation than all other Species combined. Tilia a4 ricana and Prunus serotina shared this abundance in stews of l-3.5 inch and 3.6-9.5 inch U.B.H. in class 3 and A. It is striking how abundance of stems per unit decreases With maturity, such is shown in station 20. Even though hemlock and beech were lOW'in number of stems per acre, their basal area showed doubtlessly that they were the dominant species before the des- tructive logging operation hit this stand. One thing worth notice in this particular instance, is that there were quite a number of large decayed and dead trees in the sampling area. Although they were not tallied, this may show that some of matzre or overmatured stands in the area should be operated under a better management. The flora of the deep hardwood forest is interesting. Some under~ growth of inferior tree species were found here, and the gloomy recess of this type of forest nourish plants that thrive in the shade. Here the Lycopodiums find a congenial home and flouriSh luxuriantly, While Trientalis covers the ground. The great Ulintonia with its broadoval leaves close to the earth, is also frequent and striking. We shall also meet Rubus, Snilacina, Maianthemun and a few ferns particularly'Dgzope teris spinulosa. Other Species occur, of course, but not so abundantly. Table 21. Species grandidentata Farms vrandifolia 3—“.— Tsuwa canadcnsis ___._..____ Betula ._____ 1711.113e3a Abies labalsanea 1cer saccharum , , \M— 1.11 . Vlrzlnlana M 3111a americafia Frunus ______. serotlna fl_,___._._ ~m€TCUS ' ___-_. Elli soidalis Phnms 11__, . fensylvanlca M Qusrcus —“‘—~r velut1na Ca ‘ w_ Carollnlana M Plnus ‘— Strobus 1 WK— linus ____T Tes1nosa , N :mfllanchlcr K Canadensis 1_._______7 TOTAL 7 w r a , {_ . Summarized Field Data of the your l/) fgre 411d11zs of the “orthern Hardwood-Hemlock Pype by 5126 Cluome 2 T .. 7 v No.\ ” no. r , , 25 MS 15.30 3 25 h 1.hh 2 25 30 10.36 1 50 9 3.76 2 :5 1 .36 2 25 1 .36 1 SO 1&5 59.73 2 - - — 1 - — - 1 25 15 5‘0h37‘ . 1 ? 276§2h.38 'Freq. \,'L C) w [‘3 C? \fL P) O) P” re \0 I‘ 5) \fl R) \51 S 4-771%§q.vaxp ,, , ‘ I .,No; N. No. b ‘ 1 SO 3 3.12 l 25 l 4.32“ ‘ 5.92 1.97.,1 .5 2 2.76“ _ _ _ 21.3h ’; _ _ 8.69‘ _‘ _ _ .39— ~_ _ _ f V 1 r .- 7 _ .- I 1’58 ‘i‘?‘2“5‘“ 3‘ 1.18 "'1": .1 *' '3' ”13—31, I‘D r0 U1 I‘D \Q I\) UT m ~ V1 \n [‘3 UT 1‘. ".1.-. T LTALS Dens. .u7un— d anc e D Dr h5.75 16.18 61 o r’ I") O 3.36 12.“ 12.75 b.51 51 7.75 2.73 15. 60 T LTALS S 7 Freq. 1 Dans . .Xbun- Basal Area Deng.” 3T; F1 F3 Ts 1) Dr dance 21:2 .161. /.10re N" g ,0 ! L . 1 . 12 26.66. 3 75 9.69 183 h5.75 16.18 61 20.869 21.26 26.335 3 5.5513 7; {3:39 38 . 9.50 3.36 12.66 11.033 11.21; 13.789 1 2.22 1 25 3.22 51 12.75 b.51 51 1 1.938 1.967 2.123 0 20.0 2 50 6.h6 31 _ 7.75 2.7 15.50 11.n70_ 11.68 11.3h0 3 6.66 3 75 $.69 73 13.2 6.16 21.33 0.h07 6.61 3.110 7 15.55, 2 53 6.16 31 7.75 2.73 15.70 3.757 g n u .: 7L,",h 2 SO - - 1 25 2 ‘f'lll 3 T‘: - - ‘1 5 - - 1 r: - - 2 50 $ . 3.22 .—.1__ __-4r., - P - 7'; 1 25 3.22 775 100.0 .\ . , 61 Figure 7. Phytographs of Important Species in Northern Hardwood- Hemlock Type Dr Acer rubrum Egggggg,rubre Pogulus grandidentate .Egggg_grandifolia Acer saccharum Legend: ODr—-Relat1ve density in percent OFq—-Quadrat frequency in percent OSc--Size classes in percent oBr--Relat1ve basal area in percent 62 Table 22 and 23. Data of the vegetation in the milacre quadrats of the northern hardwood-hemlock type as recorded in percentage: of coverage or number of plants by height clesees ’o-cror‘ II. -.——.--.--o-- -0 on- O-Q‘-~ - Stationr l7 Distribut19n_of coverage Classes for the type Location Hug, 366.10, (Quadrat __ 22517, Raw. 1 "“2: 3 __ 'flHeight Classes in Feet ‘* Floristic List 0-_1_1-3 3-6 0-1 1-3 3-6 0-1 1-3 3-6 o-1 1-3 3-6 Acer rubrum 28* - - - 26* _ _ 25* _ - 10% - - Amelanchi§§:canadensis 7* - - - - - - - - - _ _ Uarex Spp. "_ - - - 1 - - 0 - ~ - - - Danthonia gpicata 2 - - 2 - - 2 - - 2 — - Gaulthér§a_procumbens 0 - - 0 - - 1 - - _ _ - Koeleria_3ristata ‘*fi 0 - - - - - - - - - - - Lonicéra'canadengis 0 - - - - - O _ - - _ _ Egpuln§—'randidentata 1* - - - - - - - - . 1* — - Prun§§_serotina h* - - 4* - - 11* - - 3* - - Quercus alba 1* - - 2* - - * - - ~ - - ggggggg FEBFE' 1* - _ 4* _ - 2* - - 12* _ - flaccinium augustifoling. - - - - - - l - - 1 - — ~‘fi" “4"-..U‘ * number of plants w giétion 18 Dietribution of Coverage Classes for the Type Location SE7, 5ec.14, Quadrat T26N, RAW. 1 2 3 4 Height Class in Feet §1oristic List _*_NO-1 193 3-6 0-1 123 3-6 0-1 123 3-6 0-1 1e3 3-6 Acer sazharum - - - 4* - - 2* - - 73% _ - Aster lucidulagn - - - - - - - - - o - - Gare; blanda 1 - - 1 - — - - - - - - Carex gpp. 1 - - — - - 1 - - ‘%L - - Uogxlus cornuta 1* - - - - - - - - C - - Famgegm .. - - 2 - .. 1 - - 1. :- : géichella repeng 1 - - g" - - g - : 6* - _ Ex? virginiana * - - W - - - W - - ____Prunu=.mnsy1van1ca 5* - - 15 7* - - 1+ - - 1 _ _ Smilacina stellata O - - - - - - - : _ 1’ - Eilig,amer1cena - - - - - - - - L * number of plants 63 Table 2A. Data of the vegetation in the milacre quadrats of the northern hardwood-hemlock type as recorded in percentages of coverage or number of plants by height classes Station 19 Distribution of Coverage Classes for the Type Location SWL, SEC.9, Quadrat T20N,R5W. 1 2 3 A Height Class in Feet ElOristic List 0-1 1-3 3-6 0-1 1-3 3-6 0-1 1-3 3-6 0-1 1-3 3-0 Acer rubrwm 27* - - 21* - - 13* - _ 10* _ _ Acer saccharum - — .- - .. .. 54.;- _ _ 10;:- _, _ Amelanchier canadensis 2* - - 1* - - - - - - - - Amelanchier app. 9% - - 2* 1* _ 20* _ _ 7* _ _ Antennaria neglecta - - - - - - o - - _ - - Apocynum androsaemifolium. - - - - - - O - - - _ - Aster 1uciE§t&$'~7 1 - - 1 - — o _ - 1 _ - Carex 222, 1 - - 1 - - 1 - - 1 - - Chimaphila umbellata 0 - - - - - 0 - _ - _ - Clintonia borealis - — - - _ — o - _ - _ _ Corylus americana 3* — - 1% - - 1* 2 3% h - - Fagus grandifolia 1* - - - - - _ - - - - - Fragaria virginiana - - — O - — - — - - - — Hieracium venosum. - - - O - - - - - - - — lactuca 222° 0 _ - - - - - - _ - - - 1gpopodium.obscurum. -. - - - - - O - - - - - hitchella repens — - - _ - - o - - 0 - - Mitella diphflli o - - - - - - - _ - - - Ostrya Virginiana 1% — - 7* ~ - - — - - - - Finns strobus - - - - - - 2 - - - - - PediculariSQ_§§padensis 1 - - 1 - - - - - 1 - - Prenanthes alba 2% - - _ - - _ _ - -, - - Prunus penalyvanica — - .. .. .. .. 11 .. .. 1.: - .. Pteridium equilingg: - 2 - - 2 - - 1 - - - - PYTola secfifida o _ - o - - - - - - - - fiercus alba z-x- .. .. .. _ .. _ - - g.”- .. - Quercus ellipsoidalis - - - 1% - - - - ~ 3; 1* : Qpercus rubra - - - 1* - - - - - _ - - Salix discolor - - - O - - - - - O - .- Smilacina stellata O - - - - - 0 ' ' 0 _ _ Solidago nemora1i§_ - - - - - - 0 - ‘ _ - _ Trientalis borealig - — — - - - 1 ' : _ _ _ Viburnum acerifolium. l - - - - ' 6 ‘ _ O _ _ Viola blanda 1 - - 0 - - ‘ * number of fiiafiié M Table 25. Data of the vegetation in the milacre quadrats of the northern hardwood-hemlock type as recorded in percentages of coverage or number of plants by height classes Station' 20 Distribution of Coverage Classes for the Type Location NW£,Sec.2, Quadret TZSN, RBW. 1 2 3 I. Height Glass in Feet Floristic List 0-1 123 3-6 0—1 1-3 3-6 0-1 1-3 3-6 0-1 1—3 3-6 Acer rubrum 3 - - 2 - - 2 - - - - ggrostis alba 0 - - — - - - - - - - _ Agrostis scabra - - - 1 - - 1 - - - - - fippcynum_androsaemifolium. - - - - - - — - - 1 - - Aralia nudicaulis - — - O — - - - - - - - Usrex intumescens 0 - - 2 - - - - - l - - Carpinus caroliniana_ 1* - - - - - - — - - - - Centaurea maculosa O - - — - - - — — _ _ _ Clintonia borealis - - - - - — 1 _ _ 1 _ _ Uoptis trifolia - - - - - - - - - 1 _ - Cumswmwfih Cornus stolonifera Dryqpteris spinulosa Fagus grandifolia Galium triflorum mpodimn ob scurum Maianthemwm canadenst“. hitchella repens Polygonum cilinode Polygonum Convolvulu§g Prunuglpensylvanica Eteridium aquilinium guercus rubra Rubus hispidus Rubuslparvifloggg flubus Strigosus Smilacina stellata Trientalis borealis Tsuga_canadensis Eiturnum cassinoides Viola blanda , - - - Viola scabra ' - - ~ I I I I If I I I I I I I: I I I I I I 3 I I Icn I I I E? I I I AJI ~3AJI I I IOIOHOWI I I I I I I-‘ I I I I I I If: I-‘ I I I I I I I I I-‘ I I I I I N I I N I I IHOHHII I I I I 9HHHIOgHmIHOI I II I II 0 II I II I II IHOHIII I II I I I I I I I I I hi I I I P'ADI I I I I I I I h * number of plants 65 V. THE STUDY 1.? .‘IUAPHIC FACTCRS 1. Physiographic Features and Description of boils Under Different Vegetation Types The glacial action Which this region has been subjected has erased most of the sharp features of the topography. A large proportion of the region is relatively level to gently rolling, with slopes less than 25 percent. In some sections the land surface is practically level for miles. These areas, situated above the general level of the surrounding country, have excessive drainage and become dry in.nidsunmer. Dotted through the region are swamp areas. A great variety of soils, all podzolic, are represented by many soil types. Sand and sandy loams predominate. Locally peats and mucks or heavy clays occupy extensive areas. Generally speaking, the soils are deep as the result of extensive deposition during the glacial period. stones and boulders are abundant in some soils, but almost entirely lacking on the sand plains. Roselawn sand. Roselawn sand comprises the loose, yellowish-gray sand, which is dry to a depth ranging from.3 to more than A feet, on the pine and oak hills. The deeper underlying material consists mainly of send but contains scattered gravel and boulders and in places pockets of clay. The soil is fairly uniform in that it consists dominantly of a.ndxture of medium and fine sand, With only a small proportion of silt 66 and clay. The land is for the most part free from stones but in places boulders and gravel pocket occur (Table 26). The gravelly phase of Roselawn sand is distinguished by the pre- sence of an appreciable quantity of rounded gravel and small stones on the surface and throughout the soil. The gravel consists of waterwonn fragments of granitic rocks, chert, and some sandstone, with very little or no limestone. This type of soil is one of the more extensive soils in the region. It occurs as low swells, smooth rounded ridges, and hills with.broad dry swales or valleys intervening. The land is dry or well drained, OWing to the perviousness of the soil and free downward percolation of water. The original forest consisted dominantly of red pine, with perhaps a few scattered white pine, oaks and jack pine. The pins had been entirely out by lumbermen, and the old cut-over land has grown up to a poor or fair growth of Small oaks, red maple, aspen and white birch. A few scattered old red pines and most of the pine stumps left by the lumbermen remain. Roots of the tree extend visibly down to about 20 inches on average, and the fine roots continue to penetrate the soil to a depth of 30 to AA inches. Bracken fern, sweet fern, and lOW’blueberry are common, and in places there is a dense thicket of briers. Some hardwoods,.maple, beech, and hemlock grew on the sandy loam areas which have been included with mapped areas of Roselawn sand (58). The gravelly phase of Roselawn.soil under the Northern pin oak type consists of the following layers; (1) A surface layer, A1 and A0 (The latter being a thin layer), varying from 1 to 2 inches in thickness, 67 of mold or a loose xzu'xture of gray sand, charred organic matter, and plant roots; (2) The A2 horizon, consists of grayish leached incoherently sarfl from 2 to 1}- inches thick; (3) The horizon of 131 with dull-yellowish slight loamy sand, tinted by organic colloidal matter; about 16 inches in thickness; with scattering gravels here and there; (1.) B2 horizon is a gradation from the loamy sand, with clayr pockets fragments of granite rocks, stones, and gravels in places down to a depth of about 50 inches where tree roots usually end their extension, and (5) the weathered parent glacial drift material, the U horizon, consisting mainly of sand with some scattered gravels and boulders and here and there a pocket of clay (Figure 8). The content of organic matter is uniformly low and not durable. Where the soil is grass covered, as in spots on old cut-over land, the dark-gray color from organic matter extends to a greater depth than under trees. The moisture-holding capacity and the average quantity of water hold is low, but it is probable that a. high proportion of that present is available. The total quantity of essential plant nutrients, such as calcium, magnesium phosphorous, and potassium is lower than in the heavier soils and those containing a greater proportion of lime- stone, but there is no evidence of an abnormal deficiency of essential plant-food elements. The low fertility is compensated to some extent by the penetrability of the soil and greater freedom of root development. The reaction is strongly acid to a depth ranging from 3 to 5 feet. The gravelly areas, spots in which some clay is present between depths of 3 and 5 feet, and the dry valleys or swales between hills may be Slightly more productive than typical. 68 Grayling sand. This soil type comprises the deep, yellowish-gray sand soil of the drier pine plains. The forest soil of this type found under an aspen stand consists of the following layers: (1) A very thin layer of A0. (2) The Al layer of mold and humous soil ranging from one-half to 1% inches in thickness; (3) Underlaying the A1 is the A2 horizon in which gray sand changes to light gray sand from 2% to 3 inches in thick- ness; (h) The horizon of Bl has a dull-yellow loamy sand which becomes lighter in color with an average thickness of 16 inches and grades into (5) the B2 substratum of light yellow, wet coarse sand, sand, and fine gravel at a depth ranging from.20 inches downward. The distinguishing characteristics of this soil are its loose consistence, incoherent, or single-grained structure, sandy texture throughout, and its perviousness and nonretentiveness of moisture. The average moisture content is very low to a depth ranging from.3 to h or more feet, and the fertility is correspondingly'low; The reaction is medi m or strongly acid to a depth ranging from 3 to h or more feet. Penetration of some tree roots extend to a depth of about 30 inches (Figure 8)(Tab1e 26). In the Open grass-covered areas such as the sample plot on grass- land type, station A, the soil layer colored by organic matter is appre- ciably thicker and the tint is darker than in the same layer under jack pine or aspen. This soil is fairly uniform throughout the region, but the texture of the sand varies somewhat, from.nedium to coarse, and the amount of gravel varies from place to place. These variations may have some slight significance, in relation to cultivated farm crops and in relation 69 to average moisture content and amount of plant nutrients affecting plant growth. There is a suggestion of a little higher natural ferti- lity in areas of the gravelly phase, indicated by a somewhat more thrifty tree growth, and owing, perhaps, to a high preportion of minerals other than quartz in the parent materials or to a somewhat higher mois- ture-holding capacity. Areas of this soil are level, plainlike or very slightly uneven, owing to shallow dry depressions and hummocks of wind-blown sand. The land is excessively drained and dry, due to the perviousness of the soil and the underlying geologic formation. The water table or perma- nently wet sand probably lies at a depth of more than 15 feet. The original tree growth probably consisted mainly of jack pine and red pine; there were probably a few white pine and oaks. The pre- sent growth consists of mainly of jack pine, either in thickest or scattered in association with small groups of oaks, and aspen, and also some good aSpen stands. In the more open areas, the characteristic and more common shrubs and herbs are blueberries, low willoW, sweetfern, bracken fern, species of bluegrass, oatgrass and bunch graSS. Kalkaska loamy sand. Kalkaska loamy sand includes the lighter and deeper sand soil of the dry sandy plains and valleys, Which supports a hardwood forest. Its chief visible differences from the sands of the pine plains is in the darkAbrown or amber-colored layer Which underlies the pale lavender gray A2 leached layer at a depth of a few inches and in the characteristic slight cementation. Beneath this layer, begin- ning at a depth ranging from.15 to 20 inches, is the B2 horizon which 70 consists of pale-yellow or gray loose penetrable comparatively dry sand mixed with stones and Travels which extends to a depth ranging from 8 to more than 10 feet. The soil is not highly fertile, and is moderately or strongly acid to a depth of 30 some inches, but apparently has a higher average moisture content and hence is a little more productive than the sand of the pine plains. The land is nearly level but is pitted here and there with shallow dry depressions and lake basins and is fea- tured by low terrace escarpments (Table 26) (Figure 8). The forest cover consisted principally of hard maple, beech, yellow birch, elm, and ironwood, together with a few scattered white pine. A few tracts of virgin forest still remain in the Kalkaska county, but the greater part has been cut over and has grown up to a second growth of maple, elm, and other Species of the original forest, or, where more severely burned over, the growth is more largely aspen or the land is covered with grass. A considerable acreage has been cleared for farming, The soil is easily plowed, is nearly free from cobbles and large stones, and large bodies are comparatively uniform. Its chief deficiency is probably low moisture content. The sand also has some tendency to blow in clearly cultivated fields and, as on most of the sandy soils the fields are likely to be infested with quack grass. ggpmaw sandy loam. From.the surface downward it consists of (l) a thin surface layer of about i inch of forest mold and a one to two inches layer of dark gray loamy sand. (2) A leached, grayiSh or‘whitiSh, loose sand layer, varying from.h to 6 inches in thickness. (3) Dark, yellow or coffeeébrown sandy loam, in places gravelly or cobby and 71 slightly cemented sand, about 20 inches in thickness. (A) Levched grayish wet sand; and (5) comparatively impervious pale reddish-brown clay. The dark color at the surface, the dark-yellow or brown color of the third layer, and the presence of clty at a depth varying from 21 to A feet below the surface are distinguishing characteristics. The basal part of the sandy soil is more or less permanently wet, and the average moisture content is higher than in other sandy soils such as Grayling sand, Roselawn sand, a d Rubicon sand, and the fertility is slightly higher. In most areas, the surface is acid, although in a few places it may be nearly neutral or slightly alkaline; the underlying sandy layers are strongly acid; and owing to the presence of calcium carbonate the clay substratum.is alkaline (Table 26). This soil occurs in Small patches on nearly level plains in asso- ciation with the heavier soils. In places the land is very uneven on account of the small mounds and pits caused by the uprooting of trees. Under natural condition some of the land is wet aid imperfectly drained. The higher average noisture content is shown by the darker color and greater accumulation of organic matter, as compared with some other soils, the aSpect of the present vegetative cover, and the composition and nature of the original cover. The original tree growth consisted mainly of White pine, With a variable mixture of hemlock and hardwoods, such as elm, ash, basswood, and beech and more or less Spruce, fir and arbor vitae. The present cover of the uncultivated land consists of a dense brushy growth of poplar, aSpen, maples, alder, willow, and briers, and scattered clumps or individuals of hardwoods and other original species (FigureB). 72 RUbicon sang. Rubicon sand consists of deep-yellow comparatively dry sand on pine plains and sandy valleys. The typical profiles of this soil type consists of (1) A0, a thin layer of organic matter; (2) A1, a hori- zon.composes of forest humus mixed with dark gray sand with a thickness of from 1 to 3 inches; (3) the grayish leached sand which is designated as A2 layer; (A) Bl horizon which is a darkébrown layer of loamy sand occurs at a depth ranging from 6 to 24 inches below the surface, and beneath this is (5) the B2 layer of pale-yellowish, incoherent pervious sand (Table 26). Areas of this soil are level, and the soil has a thicker gray and brown subsurface layers unlike those in the Grayling sands. This soil is free fron.stone, moderately acid and the fertility is low,'but sup- ports a little greater amount of vegetation because of slightly higher average moisture content. At present, there is a poor to fair second growth of oaks, aspens, white pine and white birch, together'with a few widely distributed individual relics species such as white and red pine (FigureB). 73 Figure 8. Schematic Presentation of Typical Profiles of Various Soil Types é“ ‘ 6‘ . .\ V e vvl°b Roselewn Grsyling Grayling Kalksske Ogamel Rabicon Northern Grassland Jack Pine Northern Aspen Northern Pin Oak Aspen Hardwood Paper Hardwood Aspen Hemlock Birch Homlock I.: Jv'“ qr.- u.’ r‘ O --I n ‘1‘ 0'181 7h Table 26. The Depth, pH value, Content of Jrvr3 Ac Ziattcr ard Vechanical Analysis in Various Horizons. The Boil Type and Cover Type in Six hmplin; Stations. ' ' 1 Y ‘ Inca- Sail ; Gover Type and Kori-{Depth' pH Org. Kaghanical analysis (fl) tion Type i Important Spp. zon .(in.) 'Kat.)2mm. <2mr. ‘ fl sand ‘ sand 4SQM,4SM,4HK, 1 _ _1_._ 7 5 , r 1.. _....._, l I i Grassland Type 11 0-2 5.1 1.21 3.12 88.27 3.97 1. 26 3.38 ion 3 Sand;0 . AndrOpogon spp. 32 2-6 5.7 1.02 12.70 83.60 1.59 O. 2h 1.33 1 (Gravelly= Danthonia spi- Bl 6-38 5.3 0.30 1.21 96.95 C. 39 0.01 1. 38 phase) cata B2 38- 5.0 0.13 6.25 92.30 0. 28 WOO 1.17 ERmt-‘Grayling Aspen Type A 0-5 1.1 1.63 0.16 67.13 8.15 0.91 3.11 1011 ‘ sand ‘ POPUlllS grandi- Bl 5-23 503 0.85 20:22 86.20 (1.09 0006 5033 S (Gravelly ’aeniata 92 20-36 5.0 J.27 1.32 90.38 b.77 0.30 2.9 phase) Quercus alba Efiat- Roselawn Northern pin oak Al 0-2 3.6 6.66 16.18 70.89 5.35 1.02 2.56 ion _ sand gzpe A2 Z—h h.0 l. (2 6.h3 8h.89 3.02 1.00 2.h6 12 '(Gravelly Quercus rubra Bl h-l9 5.0 0.02 10.51 31.09 3.00 1.30 3.55 phase) Quercus 9111- 32 19-50 5.3 0.26 b.05 55.60 2.03 1.5h 2.39 psoidalis Iopulus :ra.di dentata _tat. (“ema* ASpen Type A 0-7 5.0 0.72 1.58 37.70 7.33 0.93 3.05 ion LOW Populus tremfl- Bl 7’18 507 1.18 0036 91050 Sol-L9 0020 2.245 3a.d loides B1 18-27 5.0 0.71 2.61 90.00 5.37 0.68 1.33 Pepulus grandi- 132 27-30 5.9 0.3 0.06 97.011 1.30 0.00 1.60 dentata ~Tetula papyrifera Acer rubrum -@at— Kalkaska Northern-Hardwood-l 0&5 5.0 1.33 3.73 53.30 10.10 0.79 1.98 1:“ Loamy , Hamlock Type 31 6-20 5.0 1.06 7.39 75.53 10.03 1.28 5.17 1“ Sand I 1cer saccharum. 32 20- 5.7 0.16 10. 01 83.10 3. 69 0.90 ..2 Fagus grandi- folia Tilia anerioana :Rat- iRubicon Northern-Hard- A1 0-3 .00! 0.62 82060. ion 13333 5.3 3 3 9.22 0.55 1.01 10 I I {W‘OOd‘ bGDflOCk W A2 I 3— 6 5.50.5]. lohh 88005 6095 1037 2019 I } linus strobus :§_} 6-2h 5.3 0.37: h.§8 85.62 5.90 1.95’1.9§ .1cer saccharum 32 2h,36 5.1 0.hh 11.70 85.80 0.02 0.00 1.98 75 2. Topography Although tepography affects vegetation indirectly by modifying other factors of the environment, it has nevertheless a significant ins fluencc upon all plant communitieS. Uniform.ve3etation may be eXpected in area that is level provided other factors being equal. Usually irre- gularities in topography such as depressions or different slopes or exposures will produce entirely unsimilar light, temperature and mois- ture conditions, and the resultant of this interaction is the various types of plant growth. Furthermore, slopes affect surface runoff and consequently will determine the degree of erosion and soil moisture condition. Geologically Speaking, if the topography is immature, drainage is relatively poor and depressions contain ponds or lakes which Vin turn will support different species of plants. In a region where moisture is rarely a critical factor, slope and eXposure produce scarcely noticeable differences in vegetation. Whereas, local topogra- phic effects will become apparent and result in vegetational zonation. Two transition tendencies of plant growth are obvious in the study of this area: one, which occurs where there is increasing soil moisture, is toward the northern hardwoods; the other, which occurs where there is decreasing soil moisture, is toward the pine community. The hardwoods, too, consist of two variants in which one or the other of the two domi- nants, plays a more important role. 0n upland sites, with decreasing soil moisture, Acer sacchargm: predominates; such is the case in station 19; Whereas in the more moist level land habitat, Hemlock {22233.2222f densis) and beech (Fagps gzpndifolia) predominate as in station 20 of the Conifer-northern Hardwood type. 76 3. Soil Reaction There are various designations for soil acidity. Generally it is eXpressed in terms of pH values. Thus, in a glossary of Special terms in the United States Department cf Agriculture yearbook of 1938 an acid soil is defined as: "A soil giving an acid reaction (Precisely, below pH 7.0; prac- tically, below pH 6.6) throughout most or all of the portion occupied by roots. More technically, a soil having a prepon- derance of hydrogen ions over hydroxyl ions in the solution." Likewise, an alkaline soil is defined as: "any soil that is alkaline in reaction (Precisely, above pH 7.0; practically, above pH 7.3)". In the development of acid soils the soluble bases are removed by conditions of high rainfall and the resulting leaching processes, while alkaline conditions are accounted for by precisely the lack of leaching during the weathering of parent material. Contributing factors in the development of acid soils are the organic acids produced by plants, the low base content of residue materials added to the soil, and the character of their decomposition. The pH of a forest soil is not constant but shows variations dur- ing the course of a year. Seasonal variations in the pH of soils are probably related to climatic conditions and to the nature of the vege- tation. The degree of drying and the kind of soil also have a marked differences in soil reaction. It seems that drying of soils is general- ly in the direction of incre sed acidity. Furthermore, altitude, aspect and drainage influence this re ction of soil. In general, pH values decrease with increase elevation, the soils of north Slopes tend to be 77 more acid than those of south and west slopes. Under conditions of poor drainage highly acid conditions are apt to develop. There is very little evidence that low pH pg£.§g is responsible for poor growth of forest trees, for in northern Europe, soils having pH values of h.0 or less are known to be highly productive, even for hardwoods, such as beech. It seems, however, that seedlings are more sensitive to soil reaction than old trees, and they develop best in soils having reaction values between pH L.5 and 6.0. It has been known that sometiwes it is necessary to acidify nursery soils in order to produce satisfactory coniferous stock. The forest soils of rest of the upland stands are strongly acid in the upper portions of the profile, especially the A layer, With a slight but gradual increase in pH with depth. The pH of the horizons from the B1 upwand was definitely lower in the poplar and oak stands than in the open grassland soil. These soil reaction tests confonn With the findings of Hilde that the pH values range from.5 to 6.5 (98). The pH values of the several forest soils are given in Table 26. Table 27. Analysis of Variance of pH Value in Different Soil Horizons and under different cover types. Source Degree of Sum Squares Mean Squares F ___ Freedom Total 17 3.6700 Between Hori- zon means 2 0.6132 0.3066 2.39 Between Type means 5 1.7652 0.3530 2.78 Error 10 1.2916 0.1292 78 There were no significant differences in different soil horizons and under various vegetation covers as far as pH value is concerned. h. Soil Temperature Soil temperature is a highly important factor, since it influences to greater or lesser degree, the physical,chemical, and biological pro- cesses in the soil. The principal source of heat is solar radiation, and its intensity in.turn will vary according to seasonal changes, at- mospheric conditions, latitude, eXposure and slope and also the living and non-living cover of the soil. Thusly, radiant energy reaching a given spot on the soil is not constant. Beside all these external factors that can influence the soil tan- perature, certain characteristics of the soil itself should not be overlooked. For properties of soil such as color, content of water, Specific heat and conductivity of heat etc. also may exert an impor- tant influence. From the ecological View point, one .ay find that soil temperature influences the germination of seeds and the survival and development of seedlingS. Many instances show that even if moisture and aeration con- ditions are favorable, seeds will not germinate or the period of germi- nation will be prolonged, if the temperature is too low. This phenome- non may explain some dense forest stands are frequently Without enough rePrOduCtion. On the other hand, in situations eXposed to intense solar radiation, surface soil temperatures may rise high enouéih to be fatal to young seedlings. 79 Data Obtained during this study show that the highest soil tem- perature at six inch under a 30-year-old jack pine stand were from ho to 7°F lower than in the open grassland during August and Septenber; and in October the differences were 60F. There seem to be no significant differences in soil temperatures among different types of forest stands, except the mature hardwood stand of station 20 showed a slightly lower temperatureS. The soil temperature data previously described are given in Table 28 and Figure 9. Table 28. Soil temperature (OF) at six inches below surface in six different cover types on different dates. TYPO Grass- Jack Northern Aspen ASpen Nerthern Hard- land ‘pine pin oak wood Hemlock Station No. A 15 9 6' _§ 18 July 26 77 7O 67 68 68 68 Aug. 2 62 6b, 62 61. 6t. 63 Aug- 9 70 63 64 6h 6h 63 Aug. 16 66 6 6h 65 65 65 Aug. 23 69 58 59 59 60 59 Aug. 30 68 61. 61. 6A 65 63 Sept. 6 61. 60 59 59 60 5? Sept. 13 72 65 66 68 65 60 Sept. 20 60 5h 55 5h 56 55 Sept. 27 57 50 53 51 52 51 Oct. 5 51 A6 A6 AA A6 #6 E53. 13 52 A8 AB 46 A6 A6 Table 29. Analysis of variance of soil temperature. Source Degree of Sum of Mean of ___ Freedom Squares Squares F Total 71 #186 . LA Between Type 5 259.25 51.85 23.89““ Keane ‘UL Between Date 11 3807.62 3h6.13 l5h.89“" Heans §f?°r 55 119.77 2'17 ** Significantly different at 1% level. Figure 9. The Relation of Soil Temperature to Date in Various 80 Cover Types 50 ................ Evasslmv‘d T As an Nmthem Pin Oak ——————— Jack Pine ~ — _____ Nowhere MdWOod-Hemtock '15 not '1 ." l,’ '\ ’\ ‘ " ‘\ e “‘ I: “‘ 0: \ ‘1 ' \\‘ /’ ‘\ \ "I /\ ‘\ d .\ ‘ I \\‘// \ I‘ / u ; 65 b- \ ‘\ - \ 'I '/\.\ ‘\ 4' \e I! ’7» /'\ \ , // ‘ 2 ‘ :7 — — I I / \ \ \‘ \‘ g. \‘—‘ / . '/' \ \ g l \ / \ \\ / , \ . F2 ' 4 \ / \ ‘. .— 60“ ‘J \‘ \ \\ '6 \ U) \-\ \‘\\ ' /\\\ 55 L ./\ \ . \\ \ ‘ “ \\‘ \ \ a , \\\ V,” 5° ' \ \. ‘\ \ \‘ ” 45* \/ a i 3! T? {5‘ f 3 TL? 2‘0 it, 3. '13 Ian], A5. at 81 Analysis of variance of soil temperature in six.sample plots indi— cated that both covering types and dates caused significant difference. The grassland soil had definite higher temperature than other covering types, Which were not significantly different aLong the selves. 5. Soil hoisture As a result of tranSpiration, large amounts of water in the fonn of water vapor pass into the atmosphere from the leaves of trees and other plants. This water loss must be balanced by uptake of water from the soil. In.xany regions, the occurrence of forest types is largeLy controlled by the supply of water. An excessive amount of water may be quite as unfavorable for tree growth as a deficiency. Usually site quality improves with increasing a cunts of available soil moisture. In an evaluation of the moisture relation of Soils, consideration should always be given to the nature of the deeper soil horizons and underlying strata. Layers of finer-textured material lying several feet below the soil surface may be highly important to support a good growth of certain type of vegetation. Stoeckeler and Sump (86) found that direct seeding of conifers Wis feasible on sand-pla'n areas in the Lake States, there a Hater-table was situated from 2-5 feet below the surface, that did not fluctuate greatly. In such situations they in- variably found more available moisture in the surface 6 inch of soil than was found in areas where the water table was deeper than Six feet. Precipitation is the principal source of water in soils, and its infiltration into the “round is influenced by the nature of soil. This 82 effect is not cnly affected by the total volunc of non-capillary :ores E of the Soil, but also by their Size and arrangerents. Beside the scil itself, other things such as the incorporated orjanic matter in mineral soils usually increases their permeability to water as a result of in- creased porosity. Losses of soil water by evaporation may be substantial if the rela- tive humidity of the atMOSphCPC decreases. The loss of water fron the soil, on the other hand, generally increases with Wind novcfient. hethods for MBQSHTBwCNt of soil moisture nay conveniently be con- sidered in tan JTOUPS: (1) those involving re oval of a szgple from the soil bocy, and (2) those in which the moisture of soil is measured .£§.§;§g. In this study of natural vegetation type, the need of a tech- nigue whereby moisture changes in undisturbed soil can be followed "“ e 1 -~‘ -~\«|-‘ q M ‘.- ' -o r\ r~~. ‘ J\ '. ~-, v~ r~ '__ r‘fl'.‘ ',' 3" -. \w . tnIOigflcab the 5rouin5 season wflfi airinu uCCCCJulfC dears is Most deSir- able. Thus, electrical resistance method of souyoucos was used and *4: readings were obtained by means 0 a.ned fied Hheetstone bridge. The electric resistance of soil at the depth of 6" and 18” in six sampling plots, i.e. station n1. h,5,5,9,15,18, were recorded weekly, then corrected to 70°F and 60°F in order to convert the data to percen- tage of noisture content by using Bouyoucos colona sand curve and Della- Bianca's Grayling snnd curve respectively. The results are sunmarized in the following table 30 and 31. Since Della~nianca's study (3) was done in the vicinity of this location and the nature of clinutic, edaphic and phytosociological factors were closely related with that of the present research, thus fifi Q a mnwk Northern 1m an 6n M" an @ meme E m" Q mu By Coloma 83nd Curve Q Grassland typos MNMm mmww m 0 mn Table 30. Percentage of soil moisture content in different cover Hemlock an analysis of variance of percentage of soil moisture content can.be carried out according to Table 31. Station %flh hm 00555555_ 0 o . :776665555 Shun/KJAUc)nwnuK/:)C2nw WWWWW56555AA 8. 7 3 5. h 2 5 6 6 .266h2632322 . .005000555 .“_57232l222 055055550500 767362222225 2 .005550300 “n.37221252l 550550550005 52536221122L 505550555 — - u 0000000 ._.78h322222 550550050550 857662612222 055500551 ___3W222lh22 555000055500 727632322222 6296306B075 W“*_WWWWW6.55 mmfiR l“ mnMR @ “Fumn M mu Rmn mm M Q_ B“ Jack pine M m" By Della-Bianca's Curve Grassland @ Wma m" m Hmbfi MNMm Mflmw m Table 31. Percentage of soil moisture content in different cover h 2 l . 0 o t o o W%&8&8mefitt uuuueeeecm JAAAAASJSSOC Station W? 3 76553266 00005982159 unnnu8689m7 08516165 “._87465L6W 9371286830 .609Wh55253 079h5h97 ._5W3h3233 07005268592 H6US7h56333 20890502 _:h8333353 0701908002l uh85W232233 003959h8 ___885h3243 00073200895 n9fl003W2233 20092537 _. "W8h33253 09056226768 n3n9935h3h2 5 %29mamsnmm x.&&&&mmmmt muuuuueeeqwmw JKAAAfiSSb 81+ Figure 10. Soil Moisture of Various Cover Type by Using Coloma Sand Curve Determined by Bouyoucos — - ——— —-—-- massmnd __ _ ASPen —_ Noxfhern Pm OcLK q . ______ _ Jack Pine _ . _ . _ NOVThQYn Hardwood—Hemlock a -‘\ m :0 \ l/ 4’\I\ ‘35— ‘\ ‘ l \ / ‘\\ or \ \ ' 'I .5 “\‘ [ll/I \ \ V // “y / —- — ’— §°+~“ III \ ,1 :\\\. /, \ / ‘6 I V) \ I ‘ / I x / \ // L‘ \ \ ///,, v:~_, ‘ / \ l ' \ ‘ -X;\//4 \ \ x/ ' \\ \t \ l :6 2. q IS 23 50 6 13 20 2’7 5 13 Jul) Aug. 5!?“ 0d“. Dole 85 Table 32. Analysis of Variance of soil moisture in different cover types, depths and dates. Source Degree of Sum Squares Mean.Squares F “w“ Freedom Total 107 586}12 Between Type 5 175 .95 35.19 22.h6** Means Between.Date 8 261.08 32.6h 20.83** Means Between.Depth l 3.25 3.25 2.07 Means Error 93 1A5o84_ 11;? ** significantly different at 1% level. The result indicates that the moisture contents of soil were dif- ferent under various venetation types and on different dates, but not at different depths. Further "t" test shows that the moisture content of soil in oak forest was highest and that of jack pine forest lowest, and followed the order as: northern pin oak type>aspen type>grassland and noethern hardwood-hemlock type) jack pine-aspen type >jack pine type. The fact that the jack pine type has the lowest moisture content among the various types of vegetation may be due to its possession of a rather open canopy and Sparsely spaced trees, consequently the evapora- tion rate of soil uoisture would be higher during a windy and hot period. 6. Organic Matter The organic matter of the soil represents an equilibrium or balance between the agencies supplying fresh organic debris and those leading to its decomposition. The balance is of course dynamic; the adjustment is probably closest in climax plant communities. In forests, the principal 86 sources are the leaves, stems, branches, roots, bark, fruits and seeds of trees. The contribution made by shrubby and herbaceous vegetation is smaller, but not without significance. In grasslands, by way of contrast, a large amount of organic ratter is deposited annually in the soil body by the roots which die. Humus is highly important from the standpoint of forest production. It functions in soil fertility, serving as a storehouse of nutrients necessary for plant growth. The physical, chemical and biological pro- perties of soil are irproved by humus with the result that conditions for the :rowth of higher plants and microorganisms are made more favor- able. Urganic matter tends to make fine-textured soils more porous and binds together the particles in coarse-textured soils. Internal drainage and aeration of soils are facilited by organic matter, and the field capacity is increased. The dry'combustion method was used to determine the content of organic matter in the soil of six sample plots. The procedure of the nethod is as follows: To 2 grams of finely ground air—dry soil (0.5 or 1 gram for $036 of the soil whose organic matter is very high) were added 0.25 gram manganese dioxide and 5 grams 60 mesh carbon free alundum. They were nixed well and placed in an alundum combustion boat and the boat was inserted irectly into the hot silica tube of fur- nace, previously heated to operating temperature of about 9500‘. The flow of oxygen was adjusted to about 100—200 c.c. per minute. Carbon dioxide evolved in combustion passed through the purifying section of the train, and wzs absorbed in a previously weighed tube filled with ascarite. Grams of carbon dioxide evolved was calculated and than converted to percent of organic matter by using the converting factor 0.471. Analyses show an average organic hatter of the A horizon of 0.725 to h.l9%, with the Al layer practically always higher than A2. In well developed Podzol profiles the A2 is frequently the poorest as far as the content of organic hatter is concerned as the result of leaching. Table 26 shows the distribution of organic matter of different horizons as feund in six different locations under various vegetation covers. Analysis of variance of organic matter in different horizons under various covers shows that (l) the covering vegetation did not cause any difference to the content of organic matter in soil in the six sampling plots. (2) the percentages of organic hatter in various horizons were significantly different. Further ”t“-tcst verifies that the organic matter in horizon A was significantly higher than that of 51 and B2, but there was no difference between that of 51 and 32. Table 33. Analysis of variance of organic matter (5) in different cover types & horizons. Source Degree of wfiSum squares Mean squares F ___ Freedom Total 17 16.58h0 setrreen type ‘ means 5 2.1903 0.h8h5 0.69 Between horizon means 2 7 .1235 3 . 71.18 5 .362? Error _ 10 6.9702 0.6é7o s significantly different at 5% level- 88 VI. THE STUDY (F CLIELTIC FACTLR 1. Temperature Temperature is like water in its action upon plants in that it has more or less to do with nearly every function, but as a working condition and not as a material. All the chemical processes of metabolism and also mahy physical processes such as diffusion, precipitation, and coagulation as in cell-wall formation etc., are dependent upon temperature and accelerated by its increase up to an optimum. With a decrease of temperature to a certain mi.iruh, growth of the plant may be retarded, and at a further decrease, death of the plant may even ensue. Plants that miQrate northward, such as the case here in Higgins Lake region, in addition to a corresponding adjustment to lower tempera- tures and a lower sum, run an increasing risk of encountering a fatal minirun. Prolonged periods during tie sniper with deficient precipita- tion and high terperature cause Lore damage to trees which are establish- ing themselves in their new environment, capecially those of the size of seedling class. However, it has been noted that even under very severe conditions on this sand plain region, trees with a diameter of more than onc~half inch at the root collar are seldom killed outright by heat, "1- though larger trees up to A inches in diameter at the root collar often are injured (78). In the course of studying, the highest temperature occurred in June 16 with a reading of 95°F although the average maximum temperature ’1 89 of that month was only 79°F. The average temperature of the month reached the peak in July with a figure of 70.20F. Table 3A gives a good indication of the distribution of daily temperature by months. 2. Precipitation A light rain usually does not affect soil moisture, for most of it will be intercepted by vegetation and will evaporate quickly. It may raise the humidity temporarily and reduce tranSpiration for a short time. If rain falls heavily for short periods, much of it will be lost by run- off, the anount depending upon the steepness of slope, nature of soil and amount and kind of cover. But if rainfall is uniformly distributed throughout the growing season, moisture conditions may'be far more favo- rable. Therefore, the total annual precipitation of an area is only a rough indication of roisture conditions for plant growth, but the sea- sonal distribution of rainfall is of ruch more importance than the total amount. Different types of vejetation usually are controlled partialky by the kind of precipitation the area enjoys, for if precipitation is regularly seasonal, there certainly is followed by a limited type of plant growth. For instance, grasslands characterize those areas where rainfall is rather light and concetrated in the spring and early summer, Winter rains with dry su hers, characteristic of several coastal regions, support shrubby vegetation. During the tine of studying, the month of July was very wet when compared with months like September and October which had a total rain- fall of 1.26 and 0.75 inches respectively, while July enjoyed a total 90 rainfall of 3.2\ inch which exceeded the normal rainfall by 3.37 inch. In one day, July 8, the area received 1.70 inch of rain. The following table 3h summarized the precipitation data. Table 3b. Climatological data recorded at Higgins Lake state forest weather bureau for the months of June, July, August, September, and October, 1952. Temperature (0F) an h wIverage Tverage Average Highest Lowest Haximum. Kininum Reading Departure from Reading Date Reading Date normal June 79.0 5 .9 69.5 3.2 95.0 16 9.0 1 July 82.3 58.0 70.2 3.h 92.0 7 38.0 31 Aug. 77.? 51.7 6b.? 0.1 89. 28 36.0 23 Sept. 71.? h0.0 58.9 1.2 90.0 11 33.0 2 Oct. 53.9 30.6 h2.3 44.6 79.0 1 17.0 18 Precipitation (in.) Henth. Total Departure from Greatest Snow, Sleet, hail normal Reading Date Total Max. Depth Date on ground June 2.35 -0.78 c.6h 17 0 0 - July 6.2h 3.37 1.70 8 o o - .‘LTI’T. 3029 0.35 1.22 14 O O .- Sept. 1.2 -1.99 0.h1 1 o o - Get. 0.75 ~1083 0.26 19 T T 19 n... 91 3. erative Humidity The moisture of the air which is in the form of vapor is termed humidity. it is one of the most important factors since it directly affects the rate of transpiration of plants. The amount of water that a plant losses frequently determines whether it can or cannot grow in a "iven habitat. The reneral humidity of a habitat depends upon climate and location with respect to bodies of water. Humidity is affCCth by temperature, wind, altitude, exposure, cover, and water content of soil. Hifh temperatures increase the capacity of the air for moisture and consequently lower the relative humidity. Hind has a powerful effect upon humidity in that dry winds lower the amount of air moisture by removinf the moist air ahout plants and mixinf it With dry air. —'1 b ‘hese two elements account for the decreased relative humidity in the air of the Kic'ins Lake area, especially during the months of summer when the temperatures and wind velocity were hign durin; the day, I '\ \Table 35). h. :Naporation lithin certain limits the combined effect of atmospheric humidity, atmospheric pressure, temperature, solar radiation, and wind is indicat- ed by evaporation. It is impossible to determine the transpirational water loss of trees directly from evaporation, althoujh in some trees the transpiration and evaporation trends are comparable. Different trees, owin~ to differences in stomatal movement, cell-sap denSitv, colloidal content of cells, and incipient drying, respond differently #9 .I'.~\lt" 92 to the enviromental factors controllinf evaporation. The CVaporation rate has a marked inf-uence not only on trunspirational water loss from trees but also on reduction of water content of the soil, especially in dry regions. Evaporation markedly determines the effeciency of rain- fall, especially where the annual precipitation is less than 30 inches. Evaporation is measured fly the United ”tates Heather Bureau by means of large Open tanks of uniform size and depth, hut this method is not quite satisfnctory for most ecological pur oses. The writer used the well-known Livingston atmameter which consists of a porous clay sphere or bulb connected to a reservoir by means of a tube. deter eVAporates from the clay urface and is constantly replaced from within. The sphere and tuhe then are filled with distilled water so that no air bubhles are present, the water "ill be drawn from the reservoir through the tuhe. '“ a”ditional small-bored tuhe is placed throujh the stopper of the reservoir will permit equalizing of pressure by nerlijible loss of water by evaporation. The reservoir is marked near the tOp and filled to the mark h; lifting a sto per. Subsequent fillings 0f diS' tilled water made at regular intervals indicates water lost to the air by evaporation over the period of -ime involved. The weeklv evaporation rate was measured COHtiHUOUSlX at four stations of different plant communities in this area durinz the period f ltmometers were set up with bulbs from July 16 to October S, 1922o w— . ° o’np‘a placed 19 inches above the ground. Jvaporation data 1” diiielent types 3_ . -. l. ' wr . Of veietation are summarized in Table 35 and 5‘1 an urapnicalld 1n F'gures 11 to It. 93 The data show that evaporation increases rather uniformly with in- creased temperature and decreases with increased amount of precipitation. For instance, durin" the week of Sept. 6 to Sept. 13, there was hardly any rainfall, and the evaporation rate before correction was 95 c.c. per week in the aspen stand while the rate increased to 233 c.c. in the open grassland. The highest mean daily atmoneter loss in the grassland community was 25.63 c.c. after correction, but only 13.92 c.c. in th jack ine forest comruuity. ilthouih evaporation rates were less in forest than in grassland, the evaporation rate in the pine forest is higher than the mixed hardwood and almost one and one half times as ireat as those in the densely canOpied hC-year old poplar stand. The writer finds the trend of evaporation rate in this region, namely, tithest rate in the Open erassland, intermediate in pine forest and lowest in a more or less densely stocked hardwood forest. During the growing season in this region, the evaporation rate is :enernllr “if“. This is caused by rather hi~h temperatures, low relative humidity, and generally constant wind during the daylight period. How- ever, when conpwrin" these data with the ones which were collected oy other investigators, the? seem to be low in their values. BOT instance, . . ‘ T 1" h {.1 ‘ yo Jhitfieid reported an fivsrase WGOKlI atonOJGter 1°55 0* 3‘“ ('C' f‘om white porcelain atmometer during the summer of 1948 on the plains near . ~ -- - L c 5 an . e are Colorado Sprlngs (96;, hhereas Jilliams and..blch re orled 1v r o . . J 4 w ' 9 s * 3 4 "“I‘EISS- daily evaporation rate of 35.2 C.C. en a sine LOTCSU: 33-3 ‘°C' *n u 1 ‘7‘ r1 / land and 29.3 c.c. in Chaparral near tee Black -orest, Colora o \100). ' v p11. . a“? V The fact that the rate of evaporation from the chamaepnitlc 1 Jer 9h is decreased in the developmrut of meSOp¥ytism has long 72c: demonstrat- 1 ed :y naay workers if tFis field. Tu ler (37} says the decreased rate of evaporation caused hy the heavier vegetation is the direct cause of succession bctueen different associations. Thereforc, 'th the develop- ment of the invading species th: evaporation conditions of the ground layer are changed which is usually also accompanied by a change in the ground flora such as the ones we have witnessed in this study. Table 35. J ckly total of evaporation (c.c.) in four different vegetation types, relative humidit35air temperature and total precipitation in Higgins Lake area (25). Evaporation Ave. ave. Totaffi TEfifiTfifi?F‘Tfifi§§f2 JaCK’_fi§§En Aclet. Temp. krccip. Date Hardwood land pine Ehmid. fixation O 19 3 15 5 g E in. July 16-July 2:3 65.11 136.3 97.0 17.7 51.1. 01.3 JU17 27-Auc. 2 131.0 10u.0 119.0 05.3 51.1 31.5 0.19 Aug. 3- Aué. 9 23.8 10.0 L0.7 *1?.0 05.3 51.1 2.2h inf. 10-1u . 16 u9.2 109.0 70.1 33.5 5&.7 53.1 0.12 flu". 17-1uc. 23 71.5 lOL.C 5b.? 52.3 Sl.h h3.& 0°52 iu~. 21-1u~. 30 cr.2 1 2.0 11h.0 0!.0 62.1 51.9 0.00 by. 31-".Cpt. 6 5.1.11.9 ,~;-:',‘7.1 50.2 35.1. 05.7 33.1 .98 mm ‘ a y; 4 {.f’ I r31. (1 l 0.00 scpt. 7-»ept.l3 K...5 179.0 131.0 0,.0 20.7 ’/‘ A SSpt.lh-ert.?0 50.8 100.0 7h.5 19.0 0L.1 13.1 0.31 Sept.21—Sept.?7 30.8 73.0 13.0 30.3 05.0 30.; 0.30 nept.28-0ct. 5 91.0 172.0 125.0 70.? 57.0 39.0 0.¢0 133 % Estimated missing data (Qnedecor 5- 95 Table 36. Analysis of variance of evaporation in different cover types and on different dates. Source Degree of Sum of Mean F freedom squares Squares Total h3 09079.57 Between Type means 3 30202.02 10057.3h 76.27%w Eetwcen Date means 10 35503.62 3550.38 26.79‘~ Error 30 3973.72 132.h6 The ever jrasslcnd was no difference 0 Op. 0 J SlTNlalCZfitly different. "\ hijtest and gal conifer-hardwood type lowest, and r different at 13 level. :ration rate among various covering types and dates w re As for covering types, the evaporation of there was Between that of jack pine type and poplar type. M" w. I Ill, .. 7 I PHI h H». A .. 7 7 . If. 1 x 7L I RV ‘ 3F... uh.» shy 91.. .. 1|..l 1 In ULA||\|I1\ ill 1|. .Ab\ . 11.1.} 5+7 .NtP-(s ‘. .I. 17,..1h7 11 lull $2.4 nl . w l a 2.9: o Rufth Ia} flkmbsu‘w .0 ? 1|. .1. a. -- L». .11.- W Mn 7.: > . v, 6 ..v (Pon,‘1liu).AVPO ‘8 7‘ 713‘- N illu1la 96 Figure 11- Graphs Showing Interrelationship Among Various Recorded Data on Soil Moisture. Precipitation, Air Temperature, Relative Humidity and Evaporation on the Grassland Type Cover WT ‘ i . ‘60.: I II ‘9 I '1 1 |4o~ ’ i t I / 7 . i i 7 ‘°/ ’ 1 g 7 2‘ / “ . ’ £ 7—7—77 1 i 3'00/ {\E E/ 7 “J E i )/ ‘ I /—/7—‘ I 100 / .' r‘ , 7 . . ‘ ‘ I 3 LE ea y +0 5. g 3 4a: ’ O :3 3“ /’ ' m 2' 7 . '5; E / '60 2' 3°: 2 . g: ’3 / ~33. 3- I/ f. “i 0 a 3 ' / (I k d: . /‘ [/7 ’ ‘, S‘s / f l r/ M._ 4 1/ / . 4.] V __ b, Colomo. 50nd Cuvve (if b ‘ lq \ ’/\ . \‘ -\“ __ __b, Colomméond Cuwe at l8" 7' ‘ ‘ \ I, \/ \\ _ . _ b) 6-mjliv3 Sand Curve (If 6 ., as ‘ ‘ V b) ijlinj Sound Cum/e at l8 :1 61- / \ _ _ _ _ _ _ y / \ \ 43-, . '6 6_ Z T's VI 4", 2r 1?: A.lP-.V fi.fi~o9i\§.ogufii§“ ) Evaem‘fim (c.c PtQCEP-‘i'afiou (hm) Soil Moisture L‘Va) _ \oo 80 97 Figure 12. Graphs Showing Interrelationship Among Various Recorded Data on-Soil Moisture, Precipitation, Air Temperature, Relative Humidity, and Evaporation on the Aspen Type Cover '70 “60 ‘\///’-‘\ ‘50 .\\ . f . _ ‘ “7440 I ; 2 430 ‘7 1 ‘1 ' t { ; L~ «i l l_ 1 ._1.i--.._l_._._._;- 2. . /' /%\\//. /\ \ 11 5'0 by Coloma Sand Curve. at 6” /' \ ._ __ by ColomaSamd Curve at I8" / \\ _. _. by Growing Samd Curve mt 6" ' ‘ _____ b 6111 M Sand Curve oi- Io" .- / \ \.. 3 3 3 6- 4.— 3.. 5 13 5 25 2 «1 IE 2.5 30 6t I 3° 1" 0d J'uly Aua. (3.) unimadwuai (961 ‘mmH “3411133 I‘IL‘I.I| y I \‘ .. sunk. w... 4....» m ”A 1 fI°~+flethAwlu>w A.a).— ‘nt.$u.T-A¢.—.d.vlfi\ A.|U. UV 98 Figure 13». Graphs Showing Interrala tionship Among Various Recorded Data on Soil Moisture, Precipitation, Air Temperature, Relative Hmnidity, and Evaporation on the Jack Pine MW Cover Type / . ' Izo— j / ,l '00 7 I 160 3: ‘1 /\ ’ 5 8° / \ 7 5° 3‘ 5F I / 3 S / 7 \ ‘40? u’: " / x / E. / . I ,7 / , ‘ 50 n ‘0‘- ]. 1+ / . l // I y/ g / / / ‘ / "/ 7/ l .. / / V / /J_ _ ‘_7____‘ 20 7o 3- ' r '2 g a / so 0 g 1/ / E: E ’ '2‘ + , o 2: .§' I 5 J cg ____t_._ L___. t ,Efllzjj’fiog o 1 __ ’ -mf ._ . IO- \ b3 Coloma Samd Cuxve (if 6" Soil Mois‘fuve (73) __ —-bj Colomo. Sound Curve. at l8” __.__bj angling Sand Cuwe at 6"" \\ _____ b3 ijlinj Sand Curve at I3 31.1.1.1: mu $0ml’16klgfiw .‘)M II L .31.... fl . v |P. 1 pi] v ..-. « \uui..!d.u$a”.:.v-I\ \... 99 Figure 14. Graphs Showing Interrelationship Among Various Recorded Data on Soil Moisture, Precipitation, Air Temperature, Relative Humidity, and Evaporation on the Northern Hardwood-Hemlock Type Cover Recifii’afi'm (£71.) m / [ / To ..; x ”<2. . 3 9% .r/ /’\7/\ r\\ .50 5‘ S ' - § .9 / C/ \ a2 ‘3 ‘3?“ 3/ » are 40; " / / / 4 > ‘ . m 4419/ / (,4 r,” 30;: " I] A 4 .I_...._ ..._..__ L i 10 '10 , 51° / i. /\/ 60 3 I s: 3. » n. so“? 1 l % r—Vl—u 3 .. - l . , . .. - ~_-. . 4.... i. 0 So” Moisture. (%) _.__. by Coloma Samd Curve at 6" ._ __ b5 Ca‘omaSand Curve wt 18" ’ \‘1 __ . _ b3 Guglins Somd Curve at 6" A |\\‘ — - —— bf) G’Yajfina Sand Cuwe‘at 18" / \ .\\ / A _ ,'\\ \‘ l,/' ‘x .4 1. \./,7 /\\ . ‘\ / \ \‘xV‘ “.«__ ~ I, / \\ ”/I' \~\’ ___— .‘h g \— K .— ‘13 R \ Duffi -_ . . \\. ( \ \ 100 VII. THE STUDY CF BIOTIC FACTURS We may say that the initial cause is biotic when the series of suc- cession starts because an area has been made bare by living agencies. Thus when an animal or plant pest produces a bare area by killing all the plants of the community as sometimes occur in forests growing in pure stands, the initial cause of the series is biotic. Sometimes the initial cause is destruction of vegetation by man. Lumbering does not usually initiate a serious succession except when followed by fire or a long period of over-grazing. Complete denudation.by animal is of in- frequent occurrence. seavers may initiate a series by daming the outlet of a lake, causing denudation by overflow and drowning of vegetation. A very destructive barkbettle k93ndroctonus_frontalis), common in the Southern Appalachian region, has killed shortleaf and pitch pines in groups and often over relatively large areas, hastening the normal succession from.pine to hardwoods. The human interference of a certain Stage in a forest succession is of great importance in forestry, the typical example of this is represented in the coastal plain, where sub- climax.pine forests are maintained indefinitely by the constantly recur- ring fires to which the pines are resistant and thus keep'down the growth of hardz'loods. In the Higgins Lake region, the associated organisms having impor- t are described taut influence upon each other and to their environmen as follows: 101 1. Kan Man, to be sure, will modify some kind of catastrophe which visited the virgin forest. Fire, for example, and not so surely epidemics of insects and disease, will be less extensive under the present condition, and the chances are that windthrow and glaze storms, the frequency and devastation of which in virgin forests are only beginning to be reduced, Will not ravage th shorter and more compact short-rotation forests of the future to as great a degree. On the other hand, with axe, saw, and fire in the slash, man can produce a forest catastrophe which surpasses in its thorough destruction of the past. Euch more subtle, but no less profound, changes in the forests are bound to result from man's upset- ting the natural balance between forest-dwelling squirrels, rabbits, mice, and deer, and their predators. The natural forest landscape was subjected to a marked change by the complete cutting down of large areas of forest. Not only does such cutting considerably change the lighting conditions, but wind and pre- CiPitation are given.freer rein, and the soil structure itself is changed markedly. While the power for nitrification increases, air capaCity and permeability of the soil are decreased. "5 The degree to which the original forest was logged is also a very inPortant influence on second-growth Species composition. Light cutting ' ' ' - ' u~ . eech sucir 0f the Virgin forest is followed by greatei abundance of b ’. gc . r- r“(113113, and hemlock; Whereas after both heavy and clear cutting the pe . . ' n ; c t associated centage of black cherry increases. Clear cutting, frecu n 1y 'eet With fire, is followed by increases in the abundance of red maple, SW \I I -11 102 birch, black cherry, and, to a less extent, of yellow'birch. Here is an excerpt from.Herbert's study on a logged tract of hard- wood forest near Johannesburg, Kichigan (h7); "This was what remained of the original timber two years after logging in a typical section of the hardwood region of the northern part of the Low Hichigan. The ground was densely covered with reproduction weeds and underbrush that had come in after the logging or just prior to it. In that particular place, there was an adequate number of forest trees of seedling size to assure a future crop. The larger trees left by the woodcutters were either defective or too small to be merchantable. host of the larger defective trees (three per acre) will die or be wind thrown before the second crop is merchantable. Those that do remain will be worthless, and will be occupying space upon which valuable timber trees could be grown. ‘hany of the trees in the smaller diameter classes have been injured in logging or are suf- fering from the changes in the physical and biotic factors brought about by logging.“ Eigg;.&s pointed out by'BrauneBlanquet (1A), the most remorseless asso- ciate of man in the destruction of native vegetation is fire. Fire is particularly destructive upon very thin, sterile soils and especialLy in the transitional region between forest and prairie, where both types of vegetation are struggling for control. Fires are always followed, whether the original vegetation is destroyed partially or entirely, by a secondary succession, which tends anew toward climax. Studies made on the old hichigan literature show that fires ran through the heavy pine slashing almost everywhere within a feW'years after logging. hany of them were set intentionally to clear the land. Some of the fires, like that of October 8, 1871, which started by des- troying the city of Manistee and swept east entirely across the state, are matters of historic record. Kittredge reported that (54) out of 77 plots chosen for the most part to represent areas as little burned l (:v Pr 103 as could be found, 66 showed evidence of one or more fires with an average of three fires since the origin of the predominating stand. Fiftyhthree of the plots had been heavily burned two or more times, With an average period of nine years between fires. Further evidence of the prevalence of fires up»to 1929 is found in.the fact that over 60 percent of the area of oak stands in the five counties in the Lower Peninsula covered by the Land Economic Survey were in the 0-3 inch diameter class. These sizes correspond to ages of less than 15 years. The fires which burned on the average every nine years were hot enough to soar or kill rany of the oaks of moderate size in spite of the fact that owing to their thick bark they are the most fire resistant Species with the possible exception of the larger pines which have their crowns above the reach of a ground fire. These fires killed all of the smaller pines and other Species, leaving only an open stand of fire- scarred oaks. The new crops of oak sprouts after each fire have formed the prevailing present stands. Repeated burning since logging has been the fate of large areas in the Higgins Lake section with the result that open degenerated stands of aSpen-pin cherry have developed. Much of this land is occupied by an understory of shrubby and herbaceous growth of which bracken fern, blackberry, dwarf bush honeysuckle, goldenrod, sedges and grasses are prominent. The occurrence and particularly the severity of forest fires.fol- lowing original logging also influence the Species composition of second growth. A destructive slash fire soon after cutting greatly 55 Cu 10h reduces the percentage of sugar maple, beech and hemlock in the subse- quent stand and is followed by an increase in the abundance of light- seeded hardwood Species such as the birches and red maple. If fire comes shortly after the aSpens have germinated, it may be sufficiently severe to kill them outright and necessitate reseeding. In such cases the land may be with little or no vegetation for two or more years. In the better types of soil, if fires recur at short enough intervals, the aspen association will maintain itself indefinately. If the fires are at long intervals the stump Sprouts of beech or maples compete for dominance and may attain it. If fires occur often, the aSpenS are favored, unless the fires are so frequent that even the aspens are killed, as is the case when fires occur annually. Under such condition a meadow of grasses usually develops and maintain itself as long as such conditions continue. In the case of sandy land, the seriousness of the situation lies in the fact that a light fire is rather certain to kill all small, pine seedling. Fortunately, under the present rigid and scientific system of fire Control, also with the cooperation from the public, there had not been a single big forest fire in this region for more than 20 some years. 2. Animal Grazing by-donestic livestock is of such minor importance that it does not need special attention. Rabbits are a menace to jack pine seedlings, eSpecially where there are abundance of shrubs. The leaving of a few scattered trees or snags 105 as roosts for hawks and owls may provide adequate control in the less critical areas. Elsewhere this may have to be supplemented by controlled hunting, trapping, or as a last resort, poisoning. Postponement of planting until the rabbit population is lOW'iS also advisable where ar- tificial reforestation is necessary in heavily populated areas. Reports from various sources show that it is very likely that more nipping damaye is done by deer than hares or rabbits, »specially in open areas Where the latter are never common and during years of low population. This is eSpecially true in the case of plantations(77). Although deer probably cause much nore do age to plantations than they are credited with, they cause relatively little mortality other than in local areas. Many surviving trees, however, lost part of their growth and deformation resulted. Sowetines, nipping in such cases sets the trees back enough to nullify the effects of the relatively expensive release opor¢ticn. is for as the d_0r population is concerned, emphasis should be placed on the prevention of overpnpulation to check the damage. But the planting and the maintenance of occasional groups of plants that make acceptable browse for deer are often adviSFble (78). 3. InSOCtS §pittlebugs: Hush more serious damage is caused by the naratoga spittlebug (Aphrophora sar3t0"ensis Fitch) to young jack and red pines. OnJv the adults of the saratoga spittlebug feed on the pines, usually from July to October. Theg puncture the bark and suck out the plant f-‘D 106 juices. Damcse is more severe in open-grown stands with considerable W.) ground cover. The eggs of the spittlehu; are laid in dead and living 'A bud scales of the host pines and other trees, such is osks ind nuples. The nymphs descend to the ground and develop on the stews of sweet fern, or other low plants, in masses of spittle, Closed canopy stand or those with an ad ixture of her woods sufficient to shade out th ground cover, are not at acked seriously. This provides the key to prevention of Seratoga spittlebu; damage. more natural control is provided by late spring frost since to persture of 180 F. or below are known to kill the nymphs. Recently, aerial spraying with DDT before the adults lay their eggs has proved effective. Pine tip—moths, The larvae of pine tip moths (ghygcionigflgrustrang Schiff.) work on the soft tissue of the new shoots, tunneling through then and later emerging from the buds or shoots, and the trees growth consequently reduced. Evidence of tip moth work appears in the summer as a small pitch mass or a dead needle at the base of the bud. Later the tips turn brown and will break off easily because they are hollow. Sometimes red pine is attacked quite severely, even in the case of mature trees. It is known that in a red pine plantation in.this region, nearly 60 percent of all the living trees were affected by tip moths in 1937. Spraying with a Penetrol-nicotine corbination or with DDT have given good control against these tip moths. Sawflies. The jack pine snwfly (geediprionpbaksianae doh) is a defoliator confining its efforts to jack pine. ‘hore common in lower 9 Michigan is the red-heeded pine sawfly (Neodiprion lecontei Fitch). ‘2 107 The former's larvae begin eating the leaves of the previous season's growth late in the spring, but the latter consumes both old and new foli‘ge. Damage often is most severe in trees growing under or adjacent to hardwoods. Complete defoliation, of course, reams death for the tree, as does repeated partial defoliation. A lead-arsenate spray is effective if the infestation is small, but is too expensive if wide spread attacks occur. Airplane spraying with DDT in oil solution has provided fairly effective nnd cheap control of these sawflies. Spruce bud worn. The Spruce budworr, Qrchi£§_fumife§ana Clem.) for quite a number of years, has been a serious pest on open-grown jack pine because such trees produce a lirg quantity of staminate flowers, the food most favorable to high survival of budworm larvae. Partial or complete defoliation may cause death or stagheadedness of, and occas- sionally reduced seed production.by, the infested tree. The attack is most likely to be fatal when the budworm works in conjunction with the jack pine sawfly-the budworm consuming the neW'fdJJege while the latter attack the old leaves. This post may be controlled most effec- tively'by'maintaining closed stands and by removing any wolf trees that bear large quantities of staminate flowers. As a rule, red, White pine or young jack pine are attacked severely only when they are growing under or very near infested older jack pine. A. Diseases Rot-producing fungi are the worst enemy of the hardwoods. The Shoe-string fungus frequently invades trees that are injured in logging, 108 playing a secondary role in their decadence. Luch of he windbreakage in mature trees is traceable to trunk rots. Beech, yellow birch, bass- wood are more susceptible to decay than their associates, and their woai rots more rapidly, once they are infected. The following are the most troublesome diseases that are common in this area: .Egd ring rottred heart,_or pecky rot). This rot is caused by the ring scale fungus, Fomes,pini. This fungus is world wide in its distri- bution in the North Temperate Zone; in the Higgins Lake region, losses resulting from this pecky rot far exceed those from any other types of decay. This fungus infects many jack pines, particularly damaged trees, the decay progresses rather rapidly. The incipient stage appears as a pink to reddish discoloration of the heartwood, which is usually retain its firmness and toughness. The advanced stage appears as few to many elongated, sore what spindle-shaped pockets parallel to the grain and separated by apparently sound wood. In the pockets the wood is reduced to a white soft fibrous mass of cellulose. Prevention of injury to the trees and adjustment of the rotation are practical control.measures. flypgxylonficanker of poplar. Hypoxylon1pruinatum causes this disease. ASpen and largetooth aSpen are most commonly attacked, infected trees are finally girdled and killed, or the trunk may break off at the canker before girdling is completed. Trees less than 30 years old and trees on poor sites suffer most. Infection occurs through wounds, particularly insect punctures, but also through ax.cuts, breakages caused by wind and ice, and through dead branches. In order to control this disease from killing the poplar and aSpen stands, infected trees should be removed 109 in thinning. Trunk conkers often occur high enough so that they are difficult to see when the trees are in leaf. Consequently, thinning should be done, or at le st the trees marked for removal, after the leaves have fallen. The first thinning should be moderate enough so that the area can be thinned again in about 5 years and a satisfactory stocked stand maintained. The second thinning is necessary to eliminate trees with incipient conkers missed by the first thinning find those with new infections. Since the fungus may live as a saprophyte on cankered trees cut and left, such material should be destroyed. Recent investigation (4A) have established a striking important fact on hypoxylon canker of aspen. In more than 95 percent of all cases observed in the Lower Peninsula of Michigan, where the cause of this type of infection could be positively determined, insects were reSpon- sible for the initial injury. By using proper silvicultural practices the abundance of these poplar borers may be materially reduced so as to minimize the infection of hypoxwlon. ’(1 110 VIII. DIJC”""ILU 0F FCiLST ”UC UJSSJI n I} THQ STU DI D Mill The different stages of a certain plant succession, generally speaking, can not be observed directly, but numerous other considera- tions offer convincing evidence of the nature of the process. A number of criteria have been used rather generally by ecologists in recognizing climax associations, and one or two others ha a seen proposed. Among these, the presence in the understory of the same species as in the overstory is obviously valid. A wido-age-distribu— tion, or the presence among the dominant species of young as well as old trees, is another way of describing the characteristics of the dominant species. Species which seed regularly and abundantly without being narrowly limited as to seed bed, which endure shade, and which reach optimum heidhts, are most likely to characterize the climax, althox h longevity may offset the absence of one or more of these characteristics. Another criterion is the number of species in the dominant stand; only those best adapted to a given site will be likely to survive indefinitely for the climax. A criterion of dubious value is the maximum volume of wood produced at a given aje or on a given site. A study of the literature reveals tha.t the virg in forest of the northern Michigan is now reduced to a very limited area, chiefly in Public ownership and intended for permanent preservation. It is the second- or third-growth forests which from now on concern silviculturists, and to which they will apply the sivical information lll gained by study of the undisturbed forests. The studied area is classified by Dice (31) within the illeghanian biotic province. The chief ecolozic characteristic of this province is the extensive development of the pines (Pinus strobus, E. resinosa and E, banksiana), which form very important successional stages, sometimes long maintained and which perhaps in some situations may never be followed by hardwood forest. Other investicators, such as Gleason (bl), point out that the pine forests are a part of a great series of associations, which have been closely related to one another since pre-glacial time, which are connected by certain definite successional trends, and which occupy the same general range, extending in a broad belt across the continent from Alaska to fiewfoundland. whereas, the hardwood forest is similarly, a part of another gr at vegetation complex, occupying most of he land between the Missouri Valley and the Atlantic Ocean and between the Great Lakes and the southern end of the Appalachian Mountain System. There is great competition between the various associations of these two complexes wherever they come in contact, and that in general the associations of the southern complex tend to displace those of the northern. For if a species is not able to hold its own in competition, and if the associations of which it is not a part, a retreating migra- tion is indicated. This is illustrated in the Northern Michigan by the renular replacement of coniferous communities by deciduous forests, indicatinc the retreat of the conifers and the advance of the hardwood trees. However, numerous relic colonies were left behind occupying *i 112 for the most part in extreme habitats, either xerOphytic rock hills and sand dunes, clothed with pines such as {inns banksiana; or a hog, characterized by species such as Larix lkaricina and Thuja occidentalis. In his more specific work done in the northern Michigan sand plain, MCAtee (67) classified the important upland forest types into four groups. The first one are areas where the dominant tree growth are oaks and maples. In this type, pin and choke cherry, witch—hazel and willow are most abundant; all of these plants are scattered also in the other types. The second type is the one which are dominated by two species of aspen, Populus tremuloides_and E. grandidentata and is better suppli- ed with sand cherry, service berry and choke cherry than the other groups. The most characteristic type of barren is that dominated by the jack pine; it appears to have more bearberry, New Jersey tea and bush honey suckle. The Northern hardwoods make up the remaining category with a little more complex species composition. The associations of the second and third-growth forests, which have followed cutting by lumberman and, in most cases, forest fires, are without doubt staging in secondary succession. Evidences all over the region indicate the profound changes which the white man has brought about. In the main, these changes are affected by: (l) the highly selective character of early logging, which removed only those species which were currently valuable; (2) the severity and extent of later logging, which removed send-bearing trees over entire region; (3) more severe and more frequent fires; and (h) great increases in the population of certain animals such as deer. 9’3 .‘1. .. . -1 _ S 113 The various stages of plant succession of this region can be post- ulated in the following manner: The introduction of the blue grass (Egg ggmprgssa) was doubtless contemporaneous with the logging, and the association now marks the site of feeding grounds and logging area. The ultimate fate of the blue grass association depends entirely upon its relation to continuous human activity. According to Thomson (88), plant succession on such.abandoned fields of hardwood lands can be summarized in these approximate steps: i.e., there is an abundance of ragweed (Ambrosia artemisiifolia) and sandbur (Cechrus pauciflorus) during the first three years following abandonment; then, they practically disappear, persisting in the fields only as a result of grazing or some other disturbances. This fact suggests that most of the abandoned lands in this region have lain idle for quite a number of years, hence they are in the advanced stage of succession. After some more years of abandonment, Canada fleabane (Erigeron canadengis) becomes more domirant, other species such as genothera biennis, Carex pennsylvanica, Asclepias tuberosa have risen in frequency. Species such as Hieracium, Euphorbia, Helianthus occidentalis, Aristida spp. can be found with the bluestem (Andropoggn scoparius). it this time, a younf forest develops rapidly at the margin of this blueirass association, and gradually encroaches upon it. In the meantime, scattered individuals of original species succeed in esta lishinf themselves within the main body of the grass association, and by their shade contribute to the destruction of the bluegrass and the eventual reappearance of the forest. Among these pioneers, 11h Rubus idaeus var. and leer rubrum.are conspicuous in the succession by the beech-maple forest, while Rhus glabra and Trunus pennslyvanica in the succession by aspens, leading ultimately to a pine forest. By the time the fiel“s have been abandoned for fifteen years, the distribution of each species has been fairly even throughout the area, but after that period, the various species be~in to appear in patches, rather than evenly distributed. This tendency becomes more accentuated in the old fields. Probably this tendency is due in part to the severe competition in the field and the micro-conditions within the field limit the distribution of each species. The growth forms of these erassland veqetation agree with the findings of Evens who studied the vegetation of an old field community in southeastern Nichigan (33A). In the nine region, when the field has been abandoned for thirty some years, the forest has encroached considerably on prairie flora. Jack pine is the dominating member of the flora, and Populus grandi- dentata is also present, shrubby plants including sweetfern, young Tfills' oak, and Rubus spp., Corylus americana occupy considerable terri- tory. Carex pensylvanica is common in the slight depression in the field. forming an almost continuous truf. In places, a thick growth of jack pine appears and there are but a few young oaks present among them. Since the oaks are more tolerant of shading than the pine, they would survive until some accidents happen to the pines would Open up a space for growth of the oaks when they would bejin growing rapidly and shade out the younger pines which might start. In this way an interspersion of oaks and jack pine would result, 115 The northern hardwood.type, as pointed out by Braun (13) is not made up of a number of distinct association, but it appears to be a formation with only one association. It is usually dominated by two species which have most of the characteristics of climax species, namely beech and maple. The latter not only the most ubiquitous species of this region, but appears most invariably in the underwood of all the stands. This is the inevitable consequence of its abundant seed pro- duction, ability to germinate and survive on varied seedbeds, and its tolerance of overstory competition. In this area, the beech maple communities of the hardwood forests may best he described as a local variant of the climax, or a stage just below the true climax-- hemlock and beech -- and seem likely to be invaded by hemlock at sometime in the future, and there can be no doubt that together they constitute a Climax association. Although hemlock is less able to become established on all kinds of seedbeds, but it over-comes this handicap by extreme longevity. Usually, stunted individuals of beech and hemlock species in the understory flTOW'Wee when the larger trees are removed. Beech-maple stands, are in some cases being invaded by hemlock as indicated by the presence of seedlings, and in other cases contain relics of a very old age class of that species. Because seedlings of hemlock is destroyed more readily by fire than the sprouting hardwoods, its absence from sampled areas today may be a result of past fire. Under the present condition, the beech-maple or maple-hemlock-beech forests are truly stable association that have great resistance to logging, fire and other destructive agencies. It is only where repeated 116 fires burn over the same area, or where poor air drainage results in frost pockets that the climax disappears completely and an entirely new association composed of pioneer species gains possession of the soil. There are few principal combinations or forest types in which the speCies of oaks occur on the sandy soils of northern Michigan. However, the Mill's oak-white-red oak type with small proportions of red pine, jack pine, red maple and black cherry, aspens are mostly found on the lepes and dry, morainal ridges in this studied area. Sherrard's (80A) work in the Roscommon County in 1902, distinguished oak flats, oak ridge and jack pine plain as the main classification for the study of forest distribution. In these early publication in 1929 (52), this oak type forest was described as characterized by~scrubby-looking clumps of oak sprouts three to fifteen feet high, scattered in the low brush. Today, one may still find the old charred pine stumps in this area, which in- dicates that the "scrub oak" lands were orifiinally timbered with pure red pine or a mixture of white and red pine. The present old growth forest in the Interlocken State Park offers an xample of the orijinal forest which has been preserved adjacent to a typical area of "scrub oak". This stand consists of a few large trees of white and red pine with an understory of varying sizes of oaks and other species. This evidence indicates that the original forest in general probably contained a large proportion of red pine and jack pine and less white pine, red oak, red maple and beech. Study (67) made on a pine forest 60 to 80 year old in the heart 117 of the oak hills region in Cgenaw County can also be used to supplement the theory that the present prevalence of oaks, particularly of small sizes, in the stands which formerly were pine. In some of the pine forest, more than 7,000 red oak seedlings to the acre were found in the sampling plots, and that illustrated the ability of the comparatively few oaks which were present in the old pine stand to seed in abundatly under the pine. A large part of the area of the aspen community in the region was occupied before human interference by a sub-climax forest which was dominated by white or red pine. Populus tremuloidgg_probably occurred as a dominant in limited areas and as an inconspicuous subordinate species in the climax forests wherever accidents or the death of large trees left opening. Logging and fires destroyed or eliminated most or all of the conifers, maple and other hardwoods. In the growing season followinrr the fire, the bare habitat was densely occupied by a group of pioneer species composed chiefly of those which came from light and wind-borne seed and from the under ground parts of the preceeding generation. The commonest of this group are, Aster spp., Betula, Corylus, Diervilla, Frajaria Virginiana, Populus tremuloides and Solidawo spp.. A large number of other species, relics which escaped the fires or those less efficient in their means of propagation are more or less frequent associates. Among them all the aspen suckers lead in heifiht growth from the first season. If the aspens are not sufficiently abundant to dominate the community after one fire, they are almost certain to do so after the seCond, third, or fourth fire. This aspen 118 type is by far the most important secondary plant community in northern Michigan, revegetatinr n:arly every type of location following the removal of virgin growth, thus standing between the devastated land and deveIOpement of the most desirable trees, whether for lumber or for recreational purposes. In qeneral a wide range of habitats may be occupied by the aspen type, from the very driest in the region to very wet although not persistently submerged soil, and from almost pure sand to a somewhat clayly soil. The species of ngulus grandidentata is most abundant in the drier sandy areas on which the aspen group occurs. The root system is widespread rather than deeply penetrating, which is somewhat favorable to windfall. Growth is rapid at first but within a few years decreases remarkably. Flowering and the maturing of seeds occurs early in the growing season. Trees from seeds or stump sprouts are soon thinned out on account of their intolerance, resulting in an open woodland. The other species, E, tremuloides prefers wetter sites, althoufh it does grow in just as sandy land. After ten to fifteen years or more under the closed canOpy of an aspen stand develOpin“ on an area followinfi one or two fires, usually gives place to a hi:her fenetic type in a comparatively short time. Numerous invading species are present in the aspen community. The most frequent are pines on the sandy upland and usually it takes longer (at least thirty to fourty years) for the pine to displace the aspens. Whereas on the better soils, this invading species would be the beech and maple which will take over within twenty to twenty five years. Such invaders must be able to grow in the shade of the aspens. As this is 119 usually less than in the virgin forest, the frowth of invaders is more rapid in the aspen stand unless the soil is too poor or too dry, or some other retarding feature prevents. Im:ortant invading species on better upland soils are suéar maple, Leech, yellow birch, and on poorer upland soils the species would be red pine, white pine and red oak. Further fires favor continuance of the aspen association indefi- nitely. Fires as often as once in twelve years favor the aspen at the expense of the pines. After the more mesic and shade-enduring species gradually invade, and become successlully established, and maintain themselves or increase in abundance and stature, the next stane of succession is thusly formed. The succession may take any one of several different directions depend- ing primarily upon the environmental conditions. Most commonly it pro- ceeds directly to the northeastern deciduous forest climax or to the northeastern conifer climax. is on more xeric habitats or on upland sites such as th: case in this ration, with decreasing soil moisture it may pass first to the white pine sta e or even to a red pine stage; whereas in the hardwood association, Acer saccharum would predominate. On the hydric side, the succession of hardwoods is often developed to the ashpelmpmaple stage or to cedar or sometimes to a mixture of these two. 120 7" r‘-‘ 1' xnm' T ("""1 1"“!er1 {\tT a...(. ..).LLV.L.C.J.L Uab'liJ L)‘; I'_I-.J.J.L V-‘ Knowledge of what constituted the orifiinal forests of a fiven locality, and of the trends which may be expected in the de'elOpment of the second growth, can he put to very practical use by forest land man- agers or silviculturists, to avoid costly mistakes in practice. They can also adapt timber stand improvement and harvest cutting methods as either to speed up natural forest succession or to hold it back, as seems required by the particular products or services desired from the land. It is a well-known fact that repeated heavy cuttings of all merchant- able timher deplrte the growing stock and cause retrogression in species composition. Clear cutting of cord wood from young second- firowth stands on short-rotation favors a great increase in the short- lived "weeds" or least desiratle species, such as pin cherry, aspen, hornbeam. Furthermore shrubby and herbaceous vegetation and poorly formed stump sprouts also cover the ground to the detriment of desirable seedlings of the better tree species. In addition to these, forest fires, like heavy cuttings deplete the forest growing stock and cause a severe retro ression in both species composition and site quality. Fire is much more common and more difficult to contral on heavily cut forest land, both because of the increased fuel and the more severe and unfavorable land condition. Significant fact which has been established in this refiion now is that fnrests are not fillinr local requirements. Lake States forests 121 currently supnly less than half of the saw timber products consumed in the region, and only about two-thirds of the total cubic volume of wood used. Shortages are most critical in softwood saw-timber products where the local forest lands supply less than one-fourth of the quantity used. Sijnificant shortages appear also in coniferous pulpwood, and high-grade veneer timber. Timber requirements fifty years hence have been estimated one-third greater than at present. The previous heavy cutting has created a poorly balanced growinf stock. The present extent of pine and hardwood forest under all kinds of protection does not indicate fully the area available for producing valuable timber in the future. Ho ever, under more aggressive management and probably with more avail- able funds, many areas now occupied by grass, brush and poor aspen stands can be restored to better types of tree species. Of course this reforest- ation pregram has to have the wholehearted cooperation of other afiencies in the field of conservation in order to achieve an allround result for the multiple use of natural resources. If protected from forest fires long enough, much of this land would restock with trees naturally, although chiefly with less valuable kinds. Such a process of natural restocking, however, would be so slow that it would be against public interest in many cases to allow these lands to lie unproductive for so many years. Hence a large part of ttis land must be restored to valuable forest growth by plantin:. Althouqh increased production of wood is the chief reason for reforest- ing the land, there are other important reasons. In some areas the planted trees would protect the watershed 1Ud prevent loss of the best 122 soil throuxh erosion. Alon: streams, lakes, roads, or other areas reforestation would e Lance recreational values; and trees and shrubs should be planted in some places to provide shelter and food for wild life. The need for reforestation in the refion had long h.en recognized hovever, even under the fireitlv expanded program tetween 1933 and l9h2, when Civilian Conserve ion Corps and other state helps contributed to the rork, the rate of reforestation was not 3reit enoug“ to replant the required ares. Looking ahead to large scale plaantingg operations, the planting situation SVOVld be siren careful thouNht and study by comparision to de's mine the least expensive class of stock which could be used to secure a well distributed stand. T‘e expem dent carried out in the "sand plain” test “round of the Huron National Forest showed th-t the transplants of 1-1 and 9-1 has a higher survival than the seedlings However, the 2-0 stock on the basis of survival distribution, and hei3ht 3rowth and the low final cost appears to he the most economical class of stock to plant. he l-O stock, is considerably below the 2-0 stock in survival and hifher in final cost, especially under severe weather Direct seedin: is not a Sal Mi factory 31‘9b1.1t8 for plentir: except under csnrc~ell~ Tavern le conditiors, nlthou‘ n jack line, red oak have 3 en successfullf seeded on sites free from afgressive, competing vege- tatior and seed-eating rodents. Due to the facts that an excessive Quan tit 3* of seed was used, the necessity of ground preparatioz, attack 123 1 ‘ V ‘ i on seeds by rodentS, birdS, fun3i and irsectr, “l ; tenperetrre at the (J‘ ‘ soil surface, and tne cost of the operation, t‘is m tloi was found at least as prent as planting would have teen. The present merchantatle stand of aspen cannot ‘e marketed advan- tafeouslv 1wec<~use the consinptitn of aSpen is comparatively small. ilthoujh the nerket for aspen -— chiefly as pulpwood, cratinj -- has already expanded substantially becuase of increasing paper mills that are equipped to utilize aspen and the impetus of the resewrch work done on hybridizing aspen, there is still little likelihood that more than a Y small proportion of availahle aspen will ever so utilized. In view of the fact that there is little chance of frouing aspen to saw-log size, except on the boot site such as station 5 of the aspen type in this study, decay usually sets in ty the tine it reaches raw-10g size and thereafter the rot pro resses so ra1idly that the trees soon Decone unmerchantahle. It is obvious from the manner in which untreated aspen stands develop that even where advance :rowth is present, this species will not develop into merchantahle size in sufficient quantit; to yield a profitable re- turn. This is true also in the case of northern pin oak and jack pine t3pe, ”Vic“ are beirf considered by some investi3ators as the So-Called temp-_ orary tvpes, They, like the aspeus, occupy certain red pine and northern wTite sine sites that have deteriorated severely as a result of at least two or three fires. In general, mam:r of the stands support some red or white pine re- production, but less than 10 percent of the area occupied by these temporary types provide enough reproduction to form a satisfactory 124 nucleus for a pine forest. Due to the facts that most of the tree species are of poor quality, frow slowly an” will yield little merchant- able material at th» tine firowth culminates, profitable use of the land depends on conversion to another forest type. From the view point of wild~ life management, conversion of an inferior forest type to u better suitable one can even change the type of animal and bird pOpulation quite favorably. Ecologically speakinfi, these temporary types may look flourish enoush to be classifies as a distinct type for the time being, never- theless, the force of succession “as alreaey set in as is shown by the fact that the oriéinal occupants of the site are actually encroaching back. During the studying of the Roscommon and Crawford Counties, Livinjston (63) found that scatterinfi seedlings of white pine were then evident on practically all areas originally covered by that species, which have not been recently subjected to the action of fires. Seedlings of the red pine, however, were more numerous on these areas than were those of the white pine itse f. They were plentiful throu hout the re- Sion on liéht soils excepting the very lightest. There were some EVidence that the red pine was gradually advancing its seedlings into the areas hold by the jack pine. The three species of pine native to the region, red, jack and white Pine, grow naturally on the sandy oak lands and are the logical species to be first considered in connection with the natural conversion of oak to pine. In the ori‘inal forest, red and white pine were the predominat- infl Species and the oaks were subordinate. The soil and climate are, therefore, suited to the restoration of the pines provided other factors 125 which have resulted from the cutting and turning of the areas are not prohibitive. fiethods are protection from fire, proper methods of cuttinc and plantinj. In this region, the evidence indicates that jack pine should be favored only on the sites less suitable to hardwoods, such as abandoned fields, or pastures, or aspen-pin cherry burns. Artificial planting of pioneer coniferous Species will often be necessary on such sites. Conversion of aspen to conifers should also be encourared. Aspen does not thrive on dry sandy soils; thus, reconversion to pine is highly desirable. Unfortunately, he rite of natural conversion is slow. With present relatively good fire protection, the rate of reseeding to pine should accelerate, but not enough to insure complete conversion, without considerable planting. much of the aspen has an understory of maple, basswood, elm, and some other Species, eventually most of it will revert to other hardwoods unless cutting, burning, pasturing, or otha? activities interfere. Limited areas may require planting. The conversion of good hardwood sites to white pine is not recomp mended, thou h silvicnlturally possible if cost is no object. Heavy hrrvest cuttinvs, in addition of leaving ample seed-bearing trees, and subsequent renovel of hardwood Sprouts and seedlings, are necessary measures for the perpetuation of pine forests on heavy soil. Nhereas, on comparatively heavy soils such as the one studied by Fisher (36) in central New Tngland, forests of pure white pine develop on abandoned fields as an intermediate stage in succession. They show, however, strong tendencies to revert to hardwood, rendering reproduction Of pine difficult. In this instance, the conversion of these forests 126 to stands of more valuatle hardwoods, or mixture of hardwoods and pine, can he accomplished by renovini the pine in one or several cuttings, followed a out four years later by cleanings or weedinrs to favor in- dividuals of the desirahle species. In the natural establisfimunt of seedlinjs of the hardwood species, decayins of forest litter seen to he of extreme importance. Fire not only Consumed these natural "seedling lads” but also deprived the shallow "l“ horizon of much of its humus content. Sugar maple, as shovn Ey dates (39) has a potentiality for a wide range in habitat suitable to invasion. Its prolific reproduction sometimes establish competition too :reat for species such as yellow birch and hemlock. ‘Partial cuttinr, not followed by fire, permits hoih species of Tsuga and Betula to reproduce in sufficient abundance to insure representation in the developnen and produce stands comparable to that in the primeval forest of the region. A. far as the silvicultural trestncnt on second-growth Northern hardwood-hemlock stands is concerned, partial cu tings are favorable to sufar nnple, heech and hemlock. A silviculture based on light and frequent harvestins of either cordwood or sawlozs will maintain a SpeCies composition similar to the virgin forest climax. Such a forest prohably is test able to meet the multiple demands for cellulose, control of water supply, wildlife, and recreation. Under t‘is form of manage- ment, trees dyinfi before maturity will be salvaged by light and frequent cuttings, growing stock will he maintained and high rates of growth in merchantable volume obtained. 127 X . SUi—lI-tliiY The successional trend of the major forest tvpes on the well- drained soil in the Higgins lake area is considered in relation to structural characteristics of the various concrete communities and also with certain physical factors of the environment. guantitative data collected from 15 one-fifth acre quadrats, representing four different forest cover types and 16 milacre quadrats of grassland type on six different soil types were analyzed and postulated to formulate the successional sequence of the existing vegetational communities. The major forest types were classified as (l) aspen type, (2) northern pin oak type, (3) jack pine type and (h) northern-hardwood- hemlock type. The other type of plant community included in this study beside these forest types was grassland which occupies open burned wild land. The associations of all these second and third-growth forests, which have followed cuttinfi by lumberman and, in most cases, forest fire, are staging in secondary succession. The blue grass association has invaded the abandoned field of hardwood lands with pioneer species such as lunhrosia artemisiifolia and Cechrus pauciflorus. But within 15 years, scattered individuals of orifinal species will succeed in estab— lishing themselves within the main body of the grass association. In the pine rejion, when the field has been abandoned for about thirty some years, jack pine, the dominating member of the flora, will encroach~ ed considerably on those prairie flora. The northern pin oak and aspen 128 type have been considere” temporary forest cover types, however, this reversion to the oricinal type of tree growth on hese land seems to have to wait for a lonr while before it tecones a reality by judging and studying the quantitative data obtained from the area. Some climatic, edaphic and biotic factors were also studi-92. m +0 ~40 (100) Uilliams, and 1. j. Holch, IQhC. 30019;; cf the Vlacn Forest of Colorado. 3001. 27:133-lh9. (101) Uilson, H. L., and Z. V. Berard, 1952. The Use of Aerial Iroto— :rnphs and Ecological Erinciples in Cover Type Hagping. Jour. Kildlife honog. 10(3):320-326. I“ H 0 PO \/ mitford, H. 2., 1901. The Genetic Bevelogn‘nt of the To rests Of ‘.ort.ern Kichigan. Tot. Gan. 31:139-139. (10;) Ioung, V. A., 1933. lb. wood Invasion in a Comparativelr Old Jhite kin Afforested Area. Ecol. 1hzol-CQ. Wfi, . _\ . .'.- i1“’..'.s.b:£k‘ ~"" Plato 1. An open accord-grown pure Jack pine stand. A fairly open 311;. in a requisite to the establishment of pine uedlinga. Plato 2. Tho stands of oak type occurring on ridges or. often of poor quality. late the thick growth or blueberries on the ground. r ' , pa‘. 1.‘ _ Plate 3. A typical stand of the upon typo. Some white oak, red oak and rod maple are in mixture. Phte h. A representative stand of northern hardwood-hemlock forest. A well-stocked understory of saplings and poles of verione epeciee has developed under protection rm firQe Plate 5. A typical soil profile of Kalkeeke loamy eeui under a hardwood stead consists principally of. hard maple, beech and elm. Plate 6. A dense stand of young Jack pine has successfully encroeched and developed on this grassland area on a. sandy soil. Demco-293 I lllWIN!illHllllWlWHllllllNIHIHHMHHIHHHI 31293101945685