1.3 _ . t 5.5.1.4.“ . . .. 1 . ‘ . . . rm“... 1:. .. 3 ‘ . 3a.? law 3.“. ... ”.0. 85 . amt". 1! c A. ; 8.3.? P.» as,‘ Sf} .1 :31. f: t? . , .‘ . 5!. .F; . V a i .7. «.1 : Luz. .fi 2 .2 7 . . . J. .‘wwuhin Wk»... .1 L J ‘ L 52" 1‘": vzt'xihfénf :59“: » an... .. «may! . z a s v." ’: .“Ptlmtnhww : : . . h‘tihufifi ‘ 3:...3. 3m x “figurinfizfl \a: {£6}... Jun“ :1 5.1:“... . I; u. 3.. 4h : “a!!! z W'AI? \nl livtnz v; ; iii}. A»; i 1. f 1.9.? . 1.: ‘9». a. I .3 03.7 1.. .‘ LP}, 1 K L.)- 0 r. ‘J J This is to certify that the thesis entitled AN HISTORICAL AND DENDROECOLOGICAL ANALYSIS OF A LONG-TERM REFORESTATION EXPERIMENT IN GRAYLING, MICHIGAN presented by Jason Scott Kilgore has been accepted towards fulfillment of the requirements for M.S. degree in Botany & Plant Pathology Major/professor Date/64 2? 400% 0-7639 MS U is an Affirmative Action/Equal Opportunity Institution LIBRARY Michigan State University PLACE IN RETURN BOX to remove this checkout from your record. To AVOID FINES return on or before date due. MAY BE RECALLED with earlier due date if requested. DATE DUE DATE DUE DATE DUE 6/01 cJCIRC/DateDuepes-pJ 5 AN HISTORICAL AND DENDROECOLOGICAL ANALYSIS OF A LONG-TERM REFORESTATION EXPERIMENT IN GRAYLING, MICHIGAN. By Jason Scott Kilgore A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Botany and Plant Pathology 2002 ABSTRACT AN HISTORICAL AND DENDROECOLOGICAL ANALYSIS OF A LONG-TERM REFORESTATION EXPERIMENT IN GRAYLING, MICHIGAN. By Jason Scott Kilgore The long-term survival, growth, and reproduction of planted trees yield insights on physiological responses to local conditions and potential for colonization. The Grayling Beal Plantation was established in 1888 to test 41 native and nonnative species of trees in the pine barrens of northern Michigan. After 112 years, survival of original stems was greatest for the nonnative Picea abies (L.) Karst. and the native Pinus resinosa Ait. and P. strobus L.; no hardwood stems survived. Regeneration was dominated by P. strobus, P. resinosa, Picea abies, and the nonnative Pinus .sylvestris L., while P. strobus and P. sylvestris were successfully regenerating in the old agricultural field adjacent to the plantation. Low site index values (28-45) reflect low productivity associated with the infertile soil. Although the high-frequency variation in radial growth for all four conifers was positively correlated to April temperatures, the native conifers were more complacent to variations in precipitation and temperature. P. .sylvestris was sensitive to variations in January, April, and September precipitation, while Picea abies was sensitive to variations in precipitation of the previous December. These results suggest that the nonnative conifers, P. abies in closed-canopy and Pinus sylvestris in open-canopy conditions, have the potential to colonize the pine barrens ecosystem. ACKNOWLEDGEMENTS I would like to thank my major advisor, Dr. Frank Telewski, for introducing me to the Beal Plantation and entitling the completion of Professor Beal's historic experiment to me. His relentless enthusiasm and support of my pedagogical pursuits were greatly appreciated. Dr. Peter Murphy has long been a mentor and friend who continues to look out for my best interest. Dr. Donald Dickmann presented pointed discussions on pine barrens ecology and, like the aforementioned committee members, advocates consideration of the important role humans have played in the forest landscape. Dr. Rich Kobe has forced me to think out of the conventional statistical box when considering trees in the landscape. I appreciate the resources all of these men have provided to this project and my professional development. This research was funded in part by a grant from Sigma Xi (The Scientific Research Society) and with logistical support from the Division of Campus Park and Planning (CPP). Grants to present this work at three conferences (Disturbance Dynamics in Boreal Forests, Kuhmo, Finland; Michigan Academy of Science, Arts, and Letters, Mt. Pleasant, MI; and Ecological Society of America Annual Meeting, Tucson, AZ) were obtained from CPP, Paul Taylor Fund (Dept. of Botany and Plant Pathology / Plant Biology), The Graduate School, lntemational Studies and Programs. and the Ecology. Evolution, and Behavioral Biology Program. Locating and tying together historical data presents many challenges. Fred Honhart (Michigan State University Archives & Historical Collections) assisted me in iii locating assorted files on Professor Beal's many activities associated with the early Michigan Agricultural College. Bill Botti directed me to a wealth of information on the State's interest in the Plantation in the 20th century, while Roger Rasmussen related the activities of the Beal Plantation Committee. Besides being a friendly face at the end ofa long field day, Kevin Gardiner (Camp WaWaSum) provided his detailed account of the 1977 Windstorm and its local effects. I could not have completed the collection of increment cores. tree metrics, and countless data points from oak seedlings without field assistance. My loving wife, Pam Kilgore, spent more time than any other person with me at the Beal Plantation; not only does she write legibly, but the time together was invaluable. Robyn Ast, Steve Nimcheski, Steve Lettero, Denise Kemp. Frank Telewski, and Alex Ztimig also helped me in the field, usually in suboptimal conditions. I received outside assistance in the interpretation of the modern ecological data. At the Laboratory of Tree-Ring Research (University of Arizona). Chris Baisan, Rex Adams, Tom Harlan, and Dr. Harold F ritts allocated time and their expertise to reviewing some of the dendrochronological data presented in this thesis. Marty Kroell (Natural Resource and Conservation Service) provided interpretation of soil-related data at the Beal Plantation. Finally, colleagues in the Departments of Plant Biology and Forestry critiqued several presentations of this project as it unfolded through the last few years. iv The Beal Plantation would no longer exist if Roger Rasmussen and Dan Sikarskie (both from Huron Pines Resource Conservation and Development Council), had not taken the initiative to protect this important piece of forest history. I also interacted with Ann Stephens and Susan Thiel (both from Michigan Department of Natural Resources). Denise Kemp (Kirtland Community College) took the lead on sponsoring the Beal Plantation to foster local academic interest. These folks, along with the rest of the Beal Plantation Committee and others before them, must be acknowledged for persevering in the face of industrial expansion. Finally, I would like to recognize the support that my family has provided. In particular, Pam forms my foundation in recognizing the value of life beyond the University. The rest of my close family. especially Bob and Kathy Reynolds, Lucy Dueck, Kim Reynolds, Justin and Jackie Kilgore, and Jason Tallant, provide the encouragement and diversions necessary to enjoy my work and life. I appreciate all of their unbiased support as I continue toward my goal of teaching at the collegiate level. ‘7 TABLE OF CONTENTS LIST OF TABLES .................................................................................. viii LIST OF FIGURES .................................................................................. ix CHAPTER 1 EARLY EXPERIMENTAL RESEARCH IN THE PINE BARRENS OF NORTHERN MICHIGAN: HISTORY OF THE GRAYLING AGRICULTURAL EXPERIMENT STATION Abstract ....................................................................................... 1 Objectives .................................................................................... 2 Introduction ................................................................................... 2 Grayling Agricultural Experiment Station ................................................ 8 Agricultural Research at GAES .......................................................... 10 Fruit Tree Research at GAES ............................................................ 12 Silvicultural Research at GAES .......................................................... 13 Agricultural and Silvicultural Research at Brinks’ Farm, Grayling .................. 21 Research at Other Substations in the Pine Barrens .................................... 22 Grayling Beal Plantation, Hartwick Pines State Park ................................. 27 CHAPTER 2 RESULTS FROM A LONG-TERM REFORESTATION EXPERIMENT IN THE PINE BARRENS OF NORTHERN MICHIGAN Abstract ..................................................................................... 53 Objectives ................................................................................... 54 Introduction ................................................................................. 54 Methods ...................................................................................... 58 Study site ........................................................................... 5 8 Survivorship and growth ......................................................... 59 Demography ....................................................................... 61 Statistical analyses ................................................................. 62 Results ........................................................................................ 63 Survivorship and growth .......................................................... 63 Demography ........................................................................ 64 Discussion ................................................................................ 68 Survivorship and growth .......................................................... 69 Demography ........................................................................ 74 Invasive potential of non-native species ........................................ 78 Future composition of the plantation and old field ........................... 80 vi CHAPTER 3 COMPARISON OF CLIMATE-GROWTH RELATIONSHIPS FOR PINA CEA E IN THE PINE BARRENS OF NORTHERN MICHIGAN Abstract ...................................................................................... 94 Objectives ................................................................................... 95 Introduction .................................................................................. 95 Methods ...................................................................................... 97 Study site ........................................................................... 97 Sample collection and data analysis ............................................ 99 Results ....................................................................................... 101 Discussion .................................................................................. 1 05 APPENDIX I ....................................................................................... 119 APPENDIX 11 ...................................................................................... 122 APPENDIX III ..................................................................................... 124 APPENDIX IV ..................................................................................... 126 APPENDIX V ...................................................................................... 129 LIST OF REFERENCES .......................................................................... 133 vii LIST OF TABLES Table 1-1. Number and mean diameter and height of surviving trees from Weldon Montgomery’s survey of the Plow Treatment in 1968 and from the whole Plantation survey in 2000. .............................................................. Table 1-2. List of participants in the Beal Plantation Committee, which was formed in Fall 1996 to protect and recognize the historical importance of the Grayling Beal Plantation ........................................................................ Table 2-1. Allometric biomass equations used to estimate total aboveground biomass (AB), including foliage and branches, of individual surviving Pinaceae ....... 20 ...... 38 at Grayling Agricultural Experiment Station (Ter-Mikaelian and Korzukhin 1997) ......... 60 Table 2-2. Equations used to estimate site index for surviving Pinaceae based on mean height at Grayling Agricultural Experiment Station (Carmean et al. 1989) ...... Table 2-3. Mean size and growth statistics [i1 SD] for originally planted and surviving Pinaceae at the former Grayling Agricultural Experiment Station, Crawford County, MI ........................................................................... Table 2-4. Rank in dominance by Importance Value of woody species and their Success Index in the plantation at the Grayling Agricultural Experiment Station, Crawford County, MI .................................................................. Table 2-5. Rank in dominance by Importance Value of woody species and their Success Index in the old agricultural field south of the plantation at the Grayling Agricultural Experiment Station, Crawford County, MI ................................... Table 3-1. Dendrochronological characteristics for the raw ring-width data and ARSTAN (ARS) mean chronology for each of the Pinaceae at Grayling Beal Plantation (GBP) and Hartwick Pines State Park (HRP) ................................... Table 3-2. Pearson correlation coefficients (r) and significance (Bonferroni probability) between the ARSTAN master chronologies developed for each species at Grayling Beal Plantation (GBP) and Hartwick Pines State Park (HRP). . . . . viii ...... 60 ...... 64 ..... 67 ...... 68 ....103 104 LIST OF FIGURES Figure 1-1. Location of the former Grayling Agricultural Experiment Station in Crawford County, Michigan .................................................................... 39 Figure 1-2. When Professor Beal arrived at the location for the Grayling Agricultural Experiment Station in early 1888, little more than stump fields remained .............................................................................................. 40 Figure 1-3. The land prior to clearing for the Experiment Station in Crawford County, Michigan, was covered with scattered jack pine and scrubby oak resprouts from frequent fires ....................................................................... 41 Figure 1-4. The southern 40 acres of the donated land were fenced, and the southern 20 acres were cleared, grubbed, and prepared for agricultural experiments ........................................................................................... 42 Figure 1-5. Layout for the 80-acre Grayling Agricultural Experiment Station as reconstructed from various sources ............................................................ 43 Figure 1-6. Experimental design for the south 20 acres of the Grayling Agricultural Experiment Station ................................................................... 44 Figure 1-7. Experimental design for the north 20 acres of the Grayling Agricultural Experiment Station for the years 1888-1890 ..................................... 45 Figure 1-8. View into the west end of the Plow Treatment of the Plantation from the old agricultural field, August 1902 ............................................................ 46 Figure 1-9. The Grayling Beal Plantation, as seen in 1998 ................................... 47 Figure 1-10. Experimental design for the agricultural plots at the Oscoda (losco County) substation, which contained 10 acres deeded by Mr. James Barlow ............... 48 Figure 1-1 1. Red pines remaining from the Oscoda Substation. now located at the trailhead to the Eagle Run Trail System, Huron-Manistee National Forest ............ 49 Figure 1-12. Experimental design for the agricultural plots at the Walton (Grand Traverse County) substation on 5 acres rented from Abram F. Philips ............ 50 Figure 1-13. Red pines remaining from the Harrison Substation. now located on private property just north of Budd Lake ..................................................... 51 Figure 1-14. Entrance kiosk to the lA-mile interpretive trail at the Grayling Beal Plantation, now an annex of the Hartwick Pines State Park ............................. 52 Figure 2-1. Location and detail of the former Grayling Agricultural Experiment Station in Crawford County, Michigan ........................................................... 82 Figure 2-2. Climate diagram (sensu Walter and Leith 1967) based on data from the Grayling, Michigan, Station (COOP ID 203391), National Climatic Data Center ................................................................................................. 83 Figure 2-3. Number of stems (>1.37 m tall. per hectare) for each shrub and tree species surveyed in the plantation (solid line) and old field (dashed line) at the Grayling Agricultural Experiment Station, Crawford County, MI .............. 84 Figure 2-4. Relative density of saplings (0-20 cm dbh) and large trees (>20 cm dbh) by species in the original Grayling Agricultural Experiment Station plantation and old agricultural field south of the plantation, Crawford County, MI ..................... 92 Figure 2-5. Distribution of Importance Values (IV) among woody species in the plantation and old agricultural field at the former Grayling Agricultural Experiment Station, Crawford County. MI ....................................................... 93 Figure 3-1. Great Lakes regional map showing the location of the Grayling Beal Plantation and Hartwick Pines State Park in Crawford County. Michigan ............................................................................................. 1 10 Figure 3-2. Climate diagram (sensu Walter and Leith 1967) based on data from the Grayling, Michigan, Station (COOP ID 203391), National Climatic Data Center .......................................................................................... 1 1 1 Figure 3-3. ARSTAN ring-width chronologies constructed for conifers at the Grayling Beal Plantation (GBP) and Hartwick Pines State Park (HRP) ................... 1 12 Figure 3-4. Correlation and response functions ( 22 PCs retained, R2=0.4021) for the ARSTAN chronology (6 trees. 25 series) for Picea abies against the mean monthly temperature (T) and monthly precipitation (P) for the previous (p) and current growing seasons at the Grayling Beal Plantation ................ l 14 Figure 3-5. Correlation and response functions (22 PCs retained, R2=0.3380) for the ARSTAN chronology for Pinus resinosa (23 trees. 82 series) against the mean monthly temperature (T) and monthly precipitation (P) for the previous (p) and current growing seasons at the Grayling Beal Plantation ................ l 15 Figure 3-6. Correlation and response functions (22 PC retained, R2=0.2835) for the ARSTAN chronology for Pinus strobus (10 trees. 21 series) against the mean monthly temperature (T) and monthly precipitation (P) for the previous (p) and current growing seasons at the Grayling Beal Plantation ................ 1 16 Figure 3-7. Correlation and response functions (22 PCs retained, R2=0.4680) for the ARSTAN chronology for Pinus sylvestris (5 trees, 20 series) against the mean monthly temperature (T) and monthly precipitation (P) for the previous (p) and current growing seasons at the Grayling Beal Plantation ................ l 17 Figure 3-8. Correlation and response functions (21 PC 5 retained, R2=0.5238) for the ARSTAN chronology for Pinus resinosa (28 trees, 52 series) against the mean monthly temperature (T) and monthly precipitation (P) for the previous (p) and current growing seasons at Hartwick Pines State Park ................... l 18 xi CHAPTER 1 EARLY EXPERIMENTAL RESEARCH IN THE PINE BARRENS OF NORTHERN MICHIGAN: HISTORY OF THE GRAYLING AGRICULTURAL EXPERIMENT STATION ABSTRACT The need for experimental research in agriculture and forestry was recognized coincidentally with the 19th century logging of Michigan’s northern forests. Supported by the Federal Morrill Act of 1862 and Hatch Act of 1887, Michigan Agricultural College established five Experiment Stations across the sandy pine barrens of northern lower Michigan to determine how to improve the soil for agriculture and to reforest the land. Forage crops were planted at the Walton (Grand Traverse County) and Baldwin (Lake County) substations, which have not been located, and forage crops and trees were planted at the Harrison (Clare County) and Oscoda (losco County) substations where red pines remain. Forage crops, vegetables, fruit trees, and a variety of forest trees were planted at the main Grayling (Crawford County) substation. The agricultural research lasted less than 5 years, and the mixed-species plantation in Grayling has been irregularly surveyed for over 110 years. Recognized for its importance as the birthplace for reforestation in Michigan, the Beal Plantation in Grayling is now listed on the State Register of Historic Sites and supports an interpretive facility as part of the Hartwick Pines and North Higgins Lake State Parks, Michigan. OBJECTIVES The objective of this chapter is to compile all information relevant to the establishment and operation of the first Agricultural Experiment Stations (AES) in northern lower Michigan. This chapter will serve as a base from which historical research on these AES can be conducted in the future. Sources have included files from the Forest Management Division (Michigan Department of Natural Resources) and the Michigan State University (MSU) Archives & Historical Collections, newspaper articles from The Avalanche in Grayling, Annual Reports of the Michigan Board of Agriculture, and published literature. Additional materials not investigated in this study may exist at the Michigan Historical Museum in Lansing. INTRODUCTION By the end of the 19‘h century, loggers had harvested most of the native pine forests in the Lower Peninsula of Michigan and had begun cutting hardwoods (Whitney 1987). After traveling through the State, A.A. Crozier, a professor from Michigan Agricultural College (MAC), said that he “cannot now recall having seen in any one place as much as a single standing acre of white pine in good condition” (Dempsey 2001). Imperfect logs and timber slash left on the land and recurrent droughts created conditions suitable for chronic wildfires set by land-clearing operations, locomotive sparks, careless settlers, and berry pickers. When Chicago (IL) and Peshtigo (WI) were burning in October 1871, a series of devastating wildfires stretched across the northern Lower Peninsula from Lake Michigan to Lake Huron (Sodders 1997). As Professor Liberty Hyde Bailey, also from MAC, wrote in a Detroit Free Press article and later cited by Beal (1888b), “What little herbage the plains afford is burned off, instead of passing into the ground to enrich it. In this manner a continual impoverishment of the soil is progressing.” Consequently, much of the northern Lower Peninsula was rendered infertile, acidic, dry, and covered with windblown sand. Because the logging companies did not pay taxes on their staked and now deforested land, the property reverted to the State of Michigan, who in turn encouraged agricultural settlement by selling the property at low prices (Dempsey 2001). However, most settlers were unsuccessful at farming the sandy soils in the 15-county region known as the Pine Barrens (Figure l-l). The farmsteads were abandoned, and the property again reverted to State ownership (Willits 1888). Reforestation efforts had already begun in the East, but governmental support for reducing the rate of logging and reforestation efforts was minimal in Michigan. Representative Robert C. Kedzie, later an MAC professor, led an early effort to preserve the remaining forests of Michigan and to begin replanting trees at the same rate as removal by logging (Dickmann and Leefers in press). Kedzie appealed to landowners that forests have practical value by controlling extreme fluctuations in rainfall, protecting crops from harsh winds, and preventing sand and snow from blowing over agricultural land (Dempsey 2001 ). In 1867, Michigan passed legislation to sanction shade tree planting along roads and to penalize removal or destruction of these trees. A decade later, another early conservationist. Representative Charles Garfield, introduced legislation to credit roadside property owners for a portion of their highway tax if they planted trees. Still, in the early 18803, legislative efforts to limit logging of the northern forests were minimal because a majority of legislators had financial ties to the logging or mining industries, which considered the forest resource limitless (Dempsey 2001). With concern over the cutting of Michigan's forests, Professor William James Beal of MAC began growing trees in 1873 for the purpose of reforestation (Telewski 1998). Specifically, Beal wanted to determine the growth requirements for a variety of tree species so that second-growth forests could provide income in the future (Beal 1878). For 17 years, Beal planted trees and shrubs from 215 taxa in The Arboretum on the campus of MAC with the purpose of educating visitors on the value of growing trees as a crop (Telewski 1998). As a result of his efforts, the public began to pressure the State government to take legislative action against the wanton forest destruction (Garfield 1905) In 1887, Hon. N.A. Beecher introduced a bill in the Michigan legislature to create an Independent Forestry Commission to investigate “the extent to which the forests of Michigan are being destroyed by fires, used by wasteful cutting for consumption or for the purpose of clearing lands for tillage or pasturage” (Reynolds 1888a,b). The Commission was also charged with reporting the effects of logging on the water bodies and climate of the State and to consider “the protection of denuded regions, stump, and swamp lands.” Within two years, the Commission. directed by Beal and Garfield, was to report back to the Governor with the results of their inquiries and proposed legislation in an effort “to preserve and restore the forest wealth of the State.” The Commission began its inquiries by asking every township supervisor a series of questions relating to the conditions of the forests and wastelands in their domain. Any fires occurring and the extent of their damage were also to be reported back to the Commission. Of the 1 185 sets of questions mailed, 722 were returned, and most were incompletely written. Over half. or 488, of the supervisors saw no need for forestry legislation (Reynolds 1888b). In January 1888, the Commission organized a Forestry Convention in Grand Rapids to “awaken a general interest in forestry matters and to aid in gathering and disseminating information on this subject” (Reynolds 1888d). Prominent landholders, academics, politicians, and lumbermen were invited to participate in this “first public step” toward addressing Michigan’s dwindling forests. Hon. Beecher opened the Convention by telling of the facts relating to Michigan's forests that inspired him to introduce the bill calling for an Independent Forestry Commission. Adding to his own experiences, Beecher consulted with others concerned about the forests. including "Dr. Beal, Prof. Bailey, President Willits, Senator Holbrook. Senator Monroe. Governor Luce, Chas. W. Garfield, Representative Watson, Representative Cross. Hon. Geo. M. Dewey. Dr. Palmer, of Grayling, an ex-member of the Legislature, and many others" (Reynolds 1888d). The rest of the Convention was given to professors. land owners, and politicians such as Bernard E. F ernow. Chief of the United Stated Division of Forestry. presenting Ur papers on topics that included fruit tree growing, statistics on the remaining forests, use of timber products, wildfires, tree and wheat culture in "waste places," woodlot management, proposed legislation, water use, maple syrup production, forestry reserves, and civic pride in tree-planting (i.e., Arbor Day). The Convention was closed with "A Convention of Michigan Trees." a humorous and educational narrative on forest use played out by members of the Convention acting as different species of trees. In June 1888, Beal organized a 2-week expedition to survey the condition of northern Michigan’s forests firsthand from Harrisville on Lake Huron’s shore to Frankfort on Lake Michigan’s shore (Beal 1888, V035 and Crow 1976). Accompanying Beal on this expedition were two recent graduates of MAC (L.A. Dewey and DA. Pelton) and two prominent botanists, Professor Liberty Hyde Bailey (MAC) and Charles Wheeler. Two reporters from Detroit newspapers. Mr. Fisher from the Free Press and Mr. H. Parish from the Tribune, were invited along to present their readers with an impression of the devastated northern woods. Their teamster and guide was W.W. Metcalf from Grayling (Beal 1888b). In the Pine Barrens. the explorers found vast stretches of logging slash and open sand fields dominated by scrubby oaks, which resprout after fire (Beal 1888, V053 and Crow 1976). After considering the current status of Michigan‘s forests and actions taken in other states and Canada, the Forestry Commission made several immediate recommendations to the State (Reynolds 1888c). First, data on forest conditions should be collected from sources considered most reliable by the Commission rather than from township supervisors. Second, to minimize the number of and damage from wildfires. the use of fire when clearing land should be prohibited from April 1 to November 1, unless the township supervisor provided written permission. Third. the Commission should be authorized to purchase “cheap lands to be set aside and maintained as a preserve.” Finally, the Commission recommended additional funds to continue their work for two more years. The need for experimentation was recognized several decades before the organization of the Independent Forestry Commission. Supported by the Federal Morrill Act of 1862, the Agricultural Colleges had been promoting “agricultural and mechanic experimentation” through the State Farmers’ Institutes and published circulars (Beal 1915). By the mid-18805, representatives of Agricultural Colleges were politicking for Federal funds to support stations devoted exclusively to experimentation. In 1885, the Michigan legislature authorized experiment station work, and, in 1887, the Federal government provided funds for these stations through the Hatch Act. Given the $15000 allotment, MAC organized its first experiment station network in 1888 under the directorship of then MAC President Edwin Willits (Beal 1915). In February 1888, among other appointees, Professor Beal was appointed Botanist, and Professor Kedzie was appointed Chemist of the newly formed experiment station network (Willits 1888). According to Willits’ (1888) opening report, he believed that “a special and thorough knowledge of the laws of nature as allied to agriculture will reduce the reducible uncertainties to a minimum and raise the productivity to a maximum.” The main experiment station was located on the campus of MAC, while substations were established throughout the lower peninsula of Michigan. The South Haven substation was launched to identify hardy varieties of fruit trees, while a series of substations were instituted across the Pine Barrens to investigate their “restoration to agricultural possibilities” (Willits 1888). These substations were located at Walton (Grand Traverse County), Baldwin (Lake County), Oscoda (losco County), Harrison (Clare County), and Grayling (Crawford County) (Willits 1888). In the hope of making the Pine Barrens agriculturally productive, Professor Kedzie wanted to test the effects of inexpensive fertilizers on the production of forage and food crops. Since animal manures were not available due to lack of forage for raising livestock and the use of superphosphates was cost prohibitive, Kedzie used readily accessible fertilizers: lake marl, salts, and plaster (Kedzie 1888d). Professor Beal had two clear objectives for these substations (Beal 1888b): “To find one or more grasses or other forage plants that shall be better adapted to the soil and climate than any heretofore in general use in such places; and to test many kinds of forest trees to learn which are best fitted to plant for timber on the sandy plains.” GRAYLING AGRICULTURAL EXPERIMENT STATION In early 1888‘, the Michigan Central Railroad Company2 donated “eighty acres of wild land” to MAC for the Grayling Agricultural Experiment Station (GAES, W'/2 of ' The last year Michigan Central Railroad is named as payer of taxes on this parcel was 1887, thus the State Board of Agriculture assumed ownership by 1888 (Crawford County Equalization Department 1887). NE‘A of Section 17, Crawford County, T.26N., R.3W., Figure 1-1) (Willits 1888). This property, located approximately 1‘/2 miles southeast of the existing village of Grayling, had been cut (Figure 1-2) and exposed to recurrent fire, especially on the north and south ends (Kedzie 1888b), which left scattered jack pines and oak root sprouts (Figure 1-3). The GAES was considered the base substation because the area’s climate and conditions would “best represent the average. - neither the best nor the poorest of the sterile land” (Willits 1888). Two problems were identified with the Grayling location (Willits 1888). First, because of Grayling’s relatively higher elevation (347 m above sea level) on the High Plains and other climatic factors. “frosts come late in the spring and early in the fall.” Frosts occur later than May 30 and earlier than September 17 in half of the years in the period 1951-1980 (Weirlein 1998). Second, recurrent fires following logging reduced the humus content of the soils: “there is but little vegetable matter in the soil.” In the Spring of 1888, Professors Beal and Kedzie supervised the clearing, fencing, and preparation of ground at GAES (Figure 1-4). The barbed wire and board fencing was intended to exclude cattle from the experiments, while a 5-foot strip of plowed ground along both sides of the fence was to prevent fire from entering the property (Kedzie 1888a,b). Kedzie (1888a) opened bid for materials and labor in the local newspaper to remove roots and stems (“grub”) from the south 20 acres by May 1, plow the south 20 acres by May 12, deliver 50 cubic yards of marl by May 23, and construct a board and barbed wire fence around the south 40 acres by May 23. Contracts 2 The tax rolls indicate that Jackson. Lansing and Saginaw Railroad Company (J.L.&S.R.R. Co.) paid the taxes on the property known as the “State Farm” prior to 1887; an assumption is made that this railroad company was later known as the Michigan Central Railroad Company. were let by the end of April to Alpheus Slaght (Grayling Township) for grubbing, James Gallamore (Ball Township) for plowing. and AI. Rose (Grayling) for fencing and marl (Avalanche 1888a). An additional contract was let in May to clear and plow another 20 acres on the north side of the GAES (Avalanche 1888b, Kedzie 1888b). All but the middle 40 forested acres were cleared and either plowed or grubbed. Except for a “tool barn” constructed on the northwest comer of the property in 1888. little infrastructure was developed at this and the other substations. AGRICULTURAL RESEARCH AT GAES The north and south 20 acres of the GAES were dedicated to agricultural experiments (Figure 1-5). The goal of this research was to determine how to increase the soil fertility with affordable fertilizers and green manures. Consequently, Professors Beal and Kedzie tested the survival and growth of different grasses, clovers, and legumes under the influence of marl, plaster (gypsum), salt, and no fertilizer. The marl was dug from the bottom of the nearby School Section Lake, while plaster and salt were shipped to Grayling by rail. Both fields were plowed 7 inches deep and harrowed to remove small woody plants, which were raked into windrows for burning (Figure 1-4). The fields were then rolled to "compact the soil." and the harrow-rake-roll sequence was repeated (Kedzie 1889b, Kedzie 1889a). Without knowing which plants would survive, the researchers sowed seed from about two dozen species in carefully measured plats in the south 20 acres at the end of 10 May 1888 and in the north 20 acres on July 5 of that year. The south field was divided into one-acre plats for each of 20 species (Figure 1-6), while the north field held plats of varying sizes (Figure 1-7). Few of the sown plants were able to tolerate the frosts and droughty conditions at the GAES. In addition, insects destroyed many of the crops; for example, red clover was devastated year after year by cut worms (Kedzie 1890). Marl was an effective fertilizer, as was plaster "in most cases," but salt was of little value (Kedzie 1890). Spurry, winter vetch. clovers, and field pea showed the greatest potential for forage and green crops (Kedzie 1890). Tall fescue, perennial rye, and tall meadow oat grass were the best performing individual grasses. but combinations of timothy and redtop with other crops like clover produced a better "sward" (Kedzie 1890). The only vegetables tested at GAES were spinach (Kedzie 1889b, 1890) and sugar beets (Kedzie 1890) in the northwest plat of the north field (Figure 1-7). Overall, the results suggested to Beal (1889b) that no crop can be grown "profitably. . .without the aid of some fertilizer." A shift in the management of the experiments at GAES may have led to a discontinuation of the agricultural research. In August 1890, Professor Kedzie (1891) was relieved of his duties at GAES by the Board of Agriculture, while Professor Beal was completely relieved of Experiment Station work the next year (Beal 1910, 1915). In 1892, Dr. 0. Palmer, a local agent and GAES agriculturist, was directing the agricultural research in Grayling (Harwood 1892). The last Experiment Station report mentioning this work summarized the continuation of activities begun by their predecessors. Tall meadow oat grass, orchard grass, fescue, and spurry showed the most promise as forage ll and green manures (Harwood 1892). No subsequent reports of these experiments at GAES were located. FRUIT TREE RESEARCH AT GAES The north 20 acres of the GAES were unforested but were included in the fenced 80 acres (Figure 1-5). For the first two years, “recuperative crops” were planted to increase the organic matter in the sandy soils (Avalanche 1888d, Taft 1890). After plowing under a crop of spurry in the Fall of 1889, the land was harrowed in the Spring and then planted to test hardy varieties of fruit trees. Taft (1890) reported “four trees” each of “45 varieties of apples, 5 of pears, 12 of plums, and 5 of cherries were planted.” The only fertilizer used was “one half pint of ground bone and potash to a tree” (Taft 1890). A 4-5 foot diameter circle around each tree was raked every 10 days. All of these trees survived their first winter but put on little growth in their second summer, which was particularly dry (Taft 1891). Another hundred trees, “mostly Russian varieties of apples,” were planted the following year (Taft 1891). After two years, the orchard had not lost any trees due to the climate. as reported by Dr. 0. Palmer, who was directing this research for MAC’s Horticulturist. L.R. Taft (1892). The orchard was not mentioned again in the annual Experiment Station reports but was noted by others. Professor C rozier wrote in 1895 that the “orchard has made but little growth” (Anonymous 1913). Most of the trees had died, except some of the 12 uncultivated trees. By 1900, the orchard of mostly Russian apple trees had been abandoned (Anonymous 19 1 3 ). SILVICULTURAL RESEARCH AT GAES Within the middle 40 acres that were not cleared, Professor Beal began his Silvicultural experiments. To test the effects of common site preparation on tree growth (Figure 1-5), he prepared three plots of land and planted trees from 40 species: 3 native to the jack-pine plains, 21 native to North America but not the plains, and 16 exotic to North America (Appendix I). All of the stock for the GAES Silvicultural experiment was obtained from W.W. Johnson of Snowflake, Antrim County. Michigan (Beal 1888b). The first “newly broken” l-acre plot just north of the grass plots was 4 rods3 wide from south to north and 40 rods long from west to east, and was “well broken up” (Beal 1888, Anonymous 1913). Fourteen rows were plowed approximately 4 feet apart, thus the author calls this the Plow Treatment. On May 21, 18884, a total of 2145 seedlings and cuttings from 39 species5 of native and exotic trees were planted on 4-foot centers in this plot (Anonymous 1913). 3 Rods are antiquated surveyor units of length with 1 rod equaling 16.5 feet, or 5.0292 meters: 3 chain is composed of 4 rods and is equal to 66 feet. or 20.1 168 meters. 4 The actual day of May 21, 1888, is assumed to be the planting date for the Plow Treatment based on the referenced planting date for the Harrow Treatment (Anonymous 1913). The actual date is likely within a couple of days prior to May 22, 1888. 5 European/Scotch elm (Ulmus montana/glubru) was not planted in this Treatment. The second “unbroken” 2-acre plot lay just north of the Plow Treatment and was 5 rods wide at the east end, 11 rods wide at the west end, and 40 rods long (Beal 1888. Anonymous 1913). The orientation of the north and south boundaries for this treatment is unknown but is assumed to be oriented as shown in Figure 1-5. Since the plot was “passed over once with a spring-toothed harrow, which seemed to tear up the soil considerably, though most of the wild shrubs and other perennials were still left in the ground ready to grow” (Beal 1888b), the author calls this the Harrow Treatment. Before planting, the plot contained some jack pine, scrub oak, “blueberries, one bear berry, trailing arbutus, Wintergreen, eagle fern. sweet fern, some dwarf service berry, choke cherry and a few grasses and other perennials” (Beal 1888b). Although Beal did not replicate his plantings, “an assortment of the trees obtained of Mr. Johnson” (Beal 1888b) “are placed here as in plowed land” (Anonymous 1913). Based on the 4-foot center planting rate observed in the Plow Treatment plot, approximately 4290 seedlings and cuttings could have been planted in the Harrow Treatment; however, this number is too large when considering that a total of 5260 seedlings and cuttings were planted in the entire plantation in 1888. Unfortunately, the trees in the Harrow Treatment were “mostly plowed up” the year after planting (Beal 1889a), thus unknown numbers of unknown species remained in the Harrow Treatment plot. The third 0.4-acre plot lay north of the Harrow Treatment, was “six rows 40 rods long,” and was not cultivated (Anonymous 1913), thus the author calls this the Control. The record (Beal 1888b) provides an ambiguous account of species planted here: “another lot of the same kinds was planted on a piece where there had been no cultivation.” If Beal planted at the 4-foot center planting rate as in the Plow Treatment, 900 seedlings and cuttings could have been planted in the Control plot. However, as for the Harrow Treatment, which species and how many of each were planted in the Control plot could not be ascertained. The records for the plantation in the few years after planting are found largely from notes transcribed on January 21, 1913, from Beal’s handwritten notes by an unknown person (Anonymous 1913). Beal’s September 21, 1888, report states that the Summer had been very dry, cultivation occurred two or three times, and hoeing occurred once. Trees doing well included locust, Russian mulberry, green ash, black cherry, box elder, catalpa, coffee-tree, beech, Norway spruce, and honey locust; trees in fair condition included weeping willow, red cedar, red pine, and white poplar; and trees doing poorly included white pine, Scotch pine“, basswood, hackwood, black ash, and European larch. A " good many" of the trees died during the first Summer and Winter (Beal 1889a). The unusually warm and dry Summer (Kedzie 1889b) was faulted, especially for the mortality of nearly all of the cuttings (Beal 1889a). The following Spring, Beal (1889a) plowed and planted a one-acre plot with "small trees," including white pine, red pine, Norway spruce, box-elder, and locust. The location of this small stand is unknown and was not again reported. He also planted seeds of pitch pine in "two or three places" and chestnuts; two 19th century pitch pine trees have survived in the plantation. Another inventory (Anonymous 1913) on July 4. 6 Scotch pine is both an historical and Silvicultural name for Scots pine (Pinus.s~_i=lvc.s'1ri.s'). The former name will be used when referring to historical texts. 15 1889, reported that the cultivation with horse and hoe the previous Fall had removed “some trees,” that the trees in the Harrow Treatment were “mostly plowed up,” and that additional pines and Norway spruce were planted in the “south rows” to replace trees that had not survived. By this second Summer. trees reported to be doing well included white pine, red pine, red cedar, box elder, and Scotch pine; trees in fair condition included green ash, red cedar, beech, elms, sugar maple, and black cherry; and trees doing poorly included locust, Russian mulberry, willows, European larch. and catalpa. Subsequent inventories appear to refer only to those trees planted in the Plow Treatment. In his report to the Director of the Experiment Station, Beal (1890) only noted that “some of the best so far are Norway pine7, White pine, Jack pine, Red Cedar, and box elder.” On November 13-14, 1895, A. A. Crozier reported the number, and sometimes condition, of trees within each of the 14 rows of the Plow Treatment (Anonymous 1913). While he considered Scotch pine to be doing the best, many other species were also surviving in good numbers: white pine, red pine, green ash, American elm, white spruce, and Norway spruce. Silver poplar was noted to be “thrifty” and sprouting up to “12 ft away from the parent tree.” In a report by an unknown author from November 3, 1900 (Anonymous 1913), red pine was said to hold the most promise followed by white pine, red cedar, and Scotch pine, on the “planted acre.” Norway and white spruce and white cedar were also growing well. Professor E.E. Bogue (1903) “studied” the plantation and concluded that the “native White and Norway Pine give the 7 Norway pine is an historical name for red pine. l6 best promise of success” and “are now 14 to 16 feet high and thrifty” (Figure 1-8). On a 1918 visit with then State Forester Marcus Schaaf, Professor Filibert Roth (1919, University of Michigan) noted that Scotch and red pine were the proven winners. Scotch pine was "the only one which produced a promising young growth or natural reproduction from seed, some of which are now 8 ft, in height," while "Norway [red] pine made the best and most shapely trees, some of them over 25 ft. tall." White pine, Norway spruce, and larch grew fairly well but were less impressive than the two aforementioned pines. Black locust rootsuekers and seedlings were present but appeared to die back regularly, while ash, elm, and poplar made a "precarious and useless existence." No other planted hardwoods or conifers were mentioned. Roth declared the experiment to be a success in showing that the land could be restocked to a forest, even with minimal care. An unauthored report of "all planted trees on the 4x40 rod area," presumably the Plow Treatment, confirms Roth's report in that Scotch and red pine showed the largest diameter and height growth followed by white pine and Norway spruce. Given the temporal coincidence, this report may be Roth's field report. Shortly thereafter, Professor A. K. Chittenden (1921) wrote in a State Board of Agriculture Special Bulletin that he was to begin an inventory of MAC’s forest plantations. In that Summer, he inventoried and calculated the mean and maximum diameter and height for all surviving trees in the Plow Treatment. Chittenden (1922) wrote the following summary for the trees originally planted here: “Of all the hardwood species which have survived none have shown any possibilities. They have largely failed even to develop to tree form and are 17 mostly merely a scrubby growth, being either unable to withstand the severity of the climate or the infertility ofthe soil.” “Of the 9 species of conifers planted, specimens of all still exist but only a few have apparently demonstrated their ability to succeed under these conditions, the red pine, white pine and Norway spruce. Two others show possibilities, the Scotch and pitch pines. The others have either made such slow growth or grow under such disadvantages that their use for forest planting on a large scale in similar localities should not be attempted. The conifers arranged in order of their success are red pine, Norway spruce, pitch pine, white pine, Scotch pine, European larch, red cedar, white spruce, and white cedar.” After the passage of four more decades, interest in the plantation was renewed. A “reappraisal by the Forest Division of the Conservation Department in 1961 confirmed the validity of the 1918 report” (Daw 1968); the “1918 report” probably refers to Roth’s (1919) report, which was contained within the 1918 Report of the State Forester. The 1961 report, which has not been located. noted the slow growth of species that had survived since the original planting. On 24-25 September 1968, Weldon J. Montgomery (Area Forester, Forest Division) surveyed the l-acre Plow Treatment, apparently unaware that trees were also planted adjacent to this stand. By this forester’s estimation, at least 95% of the merchantable volume was composed of red pine, white pine, and Norway spruce, in that order (Table 1-1). Only two Scotch pine stems had survived. but this species was seeding l8 in adjacent to the Plantation. Four other conifers, pitch pine, European larch, eastern red cedar, and northern white cedar, survived in low numbers, as well. Several hardwood species, such as elm, ash, red and Norway maple, cherry, and black locust, survived in shrubby form but were gradually dying out. Although State employees visited the Plantation again in the next decade (Kreger 1978), the trees were not tallied or surveyed. No other inventory was made until 1997 when Dr. Frank Telewski, in his capacity as Curator for the W.J. Beal Botanical Garden and Campus Arboretum at MSU, visited the site (Figure 1-9). With assistance from Denise Kemp (Kirtland Community College), he located, flagged, and measured the diameter of all trees that were most likely to have been planted in the Plow Treatment. From 1998-2000, I relocated and measured all surviving trees in all three treatments (Table 1-1) and surveyed the plant community within the plantation and old agricultural field just south of the plantation. The 4-foot center rows of trees and cut stumps from windthrow removal in 1974 (TMR 1994) and after the 1977 windstorrn (K. Gardiner. personal communication) in the Plow Treatment and Control are plainly visible. Of the 41 tree species planted as seedlings, cuttings, or seeds in 1888-1889, no original hardwood stems survived, but original stems from seven of nine conifer species survived to 2000 (Table 1-1. Appendix 1). Of these seven species. only red pine (34.5% survival) and white pine (8.4% survival) are considered to be native to the jack pine barrens, Norway spruce (26% survival) is exotic to North America. pitch pine (2 stems from an unknown number of seeds) is native to eastern North America, and white spruce l9 (4% survival), eastern red cedar (3% survival), and northern white cedar (1% survival) are native to this region but not to this forest type (Voss 1972, 1985, 1996). The two conifers with stems that did not survive were European larch and Scots pine; however, Scots pine regeneration is very common under open canopy in the plantation and old agricultural field. Several stump sprouts of red maple, white ash, eastern red cedar, and northern white cedar can be found within the plantation. Wild black cherry seedlings and bushes in the plantation and old field could be derived from outside seed sources or original root sprouts, while white poplar and black locust, which could only have been derived from the plantation, are common root sprouts in the old field. Table 1-1. Number and mean diameter and height of surviving trees from Weldon Montgomery’s survey of the Plow Treatment in 1968 and from the whole Plantation survey in 2000. Diameter was measured at 1.37 m, while height was estimated in 1968 and measured with a Vertex Forestor hypsometer in 2000. A “-“ indicates no data available. 1968 — Plow Treatment only 2000 — Whole Plantation survey No. Diameter Height No. Diameter Height Species stems (cm) (m) stems (cm) (m) Ash 9 6.9 6.1 0 - — Cedar, eastern red 10 10.4 6.1 3 17.7 12.2 Cedar, northern white 15 7.4 4.6 1 16.3 Cherry, black 3 10.4 - 0 - - Elm 8 6.9 6.1 0 - - Larch, European 1 25.4 12.2 0 - - Maple, Norway 1 - 1.2 0 - - Maple, red 1 - - 0 - - Pine, pitch 2 29.5 15 2 2 21 0 14.6 Pine, red 62 3.8 17.7 69 40 21.8 Pine, Scots 2 32.3 16.8 0 - - Pine, white 99 27.9 17.7 37 42.0 23.8 Spruce, Norway 51 75.7 15 8 24 35.9 19.5 Spruce, white 2-3 - - 2 26.6 18.3 AGRICULTURAL AND SILVICULTURAL RESEARCH AT BRINKS’ FARM In addition to the GAES, MAC rented 8 acres of cultivated land (Brinks’ Farm, S‘/2 of SW'A of Section 8, Crawford C ounty. T.26N., R.3W.) from William Brinks (Crawford County Equalization Department 1884, 1886, 1888) within a mile of the GAES for additional experiments (Avalanche 1888a, 1888c, Kedzie 1888b). While the GAES had never before been plowed, the Brinks’ Farm had been cultivated for 3-4 years (Kedzie 1888b). On May 15, 1888, Professor Beal planted 2-10 cuttings or seedlings each of 35 species of central Russian trees and shrubs from Iowa Agricultural College in close rows on two acres in the southwest comer (Beal 1888, 1889a, Anonymous 1913). Two days later, the north five acres were plowed and seeded in l-rod2 (272.25 ftz) plots of 19 species of potential forage, such as lupine, clovers, timothy, vetch, Kentucky blue grass, and rye (Avalanche 1888c, Kedzie 1888b). The last acre in the southeast corner was fertilized with plaster and sown in spurry and vetch (Avalanche 1888c). Most of the grasses did not germinate or survive the Summer, so Beal sowed more seed from 7 forbs and 92 grasses in September (Beal 1889a). He also planted a Niagara grape vine from E.M. Roffee (Clyde, NY) in 1888, but the vine could not withstand the Winter and late frosts of Grayling (Beal 1889a). Beal (1890) reported the successful growth of several species of grasses and alsike clover at the Brinks’ Farm and contemplated plowing, fertilizing, and seeding with the “most promising clovers and grasses” the following Spring. He also reported that some of the Russian trees and shrubs seemed “to promise more for this soil and climate than most of our natives,” while others died or grew little 21 (Beal 1889a). However, no mention was made of the experiments at Brinks’ Farm after 1890. A restaurant now operates on this site. RESEARCH AT OTHER SUBSTATIONS IN THE PINE BARRENS Given the demand by settlers of northern Michigan, Professor Beal was directed to locate, rent, and conduct experiments at four more substations to improve the agricultural condition of the Pine Barrens (Beal 1888b). These substations were to be 5- 10 acres each and "subordinate to Grayling" (Willits 1888). Although more travel would be required to manage all five substations, Beal agreed that better results could be obtained by using multiple locations representing the breadth of Pine Barrens productivity (Beal 1888b). In fact. the eventual substation at Oscoda represented the poorest of the "jack-pine plains," followed by Grayling, then Walton and Baldwin, and finally Harrison with the best land (Beal 1889b). While the condition of the soil indicated its potential productivity. Beal also noted that the number of native plant species was smallest at Oscoda and largest at Harrison (Beal 188%). At each of the four substations, Beal established similar experiments with plots containing the same species of seed, either as individual species or mixtures within each plat, including mammoth clover, red clover, alsike clover. sweet clover, tall oat grass, orchard grass, perennial rye grass, June grass, meadow foxtail, millet, Hungarian grass, meadow fescue, spurry. timothy, alfalfa (luceme), and field peas. Although Beal conducted agricultural experiments at all of the substations. he intended to plant trees only at Harrison and Oscoda (Willits 1888). The poorest of jack-pine plains was located in northeast lower Michigan. Beal approached Mr. James Barlow, a farmer producing grains and vegetables on drained swampland on the south side of the AuSable River and l/.)-mile northwest of Oscoda's railway station (Beal 1888b). Barlow was so convinced that the "Jack-pine land" was unproductive that he deeded 10 acres of such property upon which Beal could conduct his experiments. After fencing the deeded property, 5 acres were cleared, plowed, harrowed, and sown with seed on May 15, 1888 (Beal 1888b) (Figure 1-10). In April 1889, Beal topdressed a portion of each plat with fine barnyard manure, which improved growth in every species by July, and another portion near the center of each plat with superphosphate (Beal 1889a). In that same Spring, Beal plowed and harrowed another acre of the deeded property, taking care to leave standing jack pines alone, and he planted small (6 in tall) seedlings of white pine, red pine, Norway spruce, box elder, and locust and sowed seeds of pitch pine (Beal 1889a). The source for the stock and seeds is unknown. The trees, too small by Beal's account, were planted in hoed rows spaced 4 ft apart and running east to west. No other records were located that describe the specific activities at the Oscoda substation. At some point during the 20th century, the land containing the substation was incorporated into the Huron-Manistee National Forest as part of the River Road National Forest Scenic Byway (Telewski 1998). Although the plantation was harvested in the early 19805, several mature red pine trees, probably left as seed sources, remain within 4-ft rows of stumps (Figure l-1 1), and black locust and pitch pine grow within the sapling layer. The plantation is easily located near the trailhead to the Eagle Run Trail System. 23 The 10-acre Walton substation of fenced jack-pine plains was rented from Abram F. Philips and was located about l/2-mile northwest of Walton and adjacent to the north side of the Grand Rapids & Indiana Railroad (Beal 1888, 1889a). Five acres of this "new land" in the southeast corner were rented for agricultural experiments. Similar to Oscoda, this rented land was platted, plowed, harrowed, sown with seeds, and rolled on the eleventh and at the end of May 1888 (Beal 1888, 1889a) (Figure 1-12). In October, Beal visited this substation with Hon. C.W. Garfield, a member of the State Board of Agriculture; Garfield agreed that the Pine Barrens "rank as poor" and that farmers would be most interested in the best land while it remains "cheap and abundant" (Beal 1889a). In the following April, Beal spread barnyard manure over 20 ft2 on the east end of each plat and covered a similar area near the middle of each plat with "Homestead superphosphate” (Beal 1889b). As at other substations, Beal reported that manure helped all species but especially the grasses, while superphosphate helped the clovers most (Beal 1889a). To test when plowing and seeding were most effective, Beal shallowly (5 in deep) plowed about l/3-acre of the "new land" in July 1889, fertilized with superphosphate (300 lbs/acre) and slaked lime, harrowed, and seeded the area with a mix of three clovers, tall oat grass, orchard grass, and timothy (Beal 1890). This mixture of clovers and grasses had proven to be the most productive to date for improving the constitution of the soil. Although no details were reported about this experiment, Beal did note that Summer plowing followed by Spring backsetting and seeding seemed to work better than Spring plowing and seeding because woody species did not have sufficient stored “protoplasm” to recover from disruption during the Summer (Beal 24 1890). No other records were found that describe the specific activities at the Walton substation, which has not been relocated. The Baldwin substation was located on 8 acres of jack-pine plains rented from TV. Childs and located "some 40 rods" (660 ft) south of the "village school-house" and adjacent to the Chicago & West Michigan Railway (Beal 1888b). The land, which had been first plowed in Spring 1888, was platted in an experimental design similar to Oscoda, harrowed, rolled, and sown with seeds on May 9, 1888. A brief visit on July 30 showed growth "a little better than those at Oscoda." On June 29, 1889, Beal noted that the plants in the plats with harrowing only (Plat 1) did not survive (Beal 1889a). In that visit, he applied plaster and a little manure from the village stables on portions of each plat. In July, Beal shallowly (5") plowed 1/3-acre and, in the following Spring, backset. fertilized, sowed seed, and harrowed similar to what he did at Walton. This substation was not mentioned after 1890 and also has not been relocated. Supposedly representing the best of the jack-pine plains, the Harrison substation was originally located on 5 acres of rented land from B.B. Pixley (Beal 1888b). This land had been cropped for 6 years without manure in rutabagas, muskmelons and watermelons, oats, beans, corn or beans, buckwheat or millet, and light oats, respectively. On the seventh and end of May 1888, the property was plowed, harrowed, sown with seeds, and rolled in a fashion similar to that at Oscoda. An August visit showed less promise in the plants than at other stations where the "newly broken land" contained fewer weeds, such as lamb's quarter, pigeon grass, sorrel. and wild morning glory (Beal 1888b). Beal also seeded some "new land" in that Spring that "all caught first rate" (Beal 188%). However, Beal was not satisfied with Pixley's fence maintenance practices, so he abandoned his experiments here but concluded that fertilizers would be necessary to grow crops (Beal 1889a). Instead, Beal accepted the deed for 10 acres of jack-pine land from W.H. and FA. Wilson (Beal 1888b). This property was located '/2- mile north of Harrison near the fairground and railroad. After fencing the property, from which some sizable red pine had been formerly cut, Beal plowed the south 5 acres but not before grubbing the most southern 2 acres in Fall 1888 (Beal 1888, 1889a). In the following Spring, the plowed area was rolled, harrowed, and seeded in plats with clovers, grasses, and alfalfa. A "liberal dressing of fairly well rotted bam-yard manure" was spread over 20 ft2 of each plat. These seeds reportedly did well but did not receive superphosphate according to Beal's plan. Although Beal stated that trees would be planted on the Wilson property in Spring 1889 (Beal 1888b), he did not mention any trees again in the record. However, a l-acre plantation of 18 mature red pines dating back to at least the 18905 (F. Telewski, unpublished data) grow in 4-foot center rows on the site. located on private property just north of Budd Lake and adjacent to the abandoned railroad right-of-way (Telewski 1998) (Figure 1-13). Beal apparently planted at least red pine at this substation. The record of activity for all four substations ends before the 20th century, thus the substations can be assumed to have been abandoned by that time. Beal (1890) last reported that he planned to plow under the crops in June 1891 and to seed in a mixture of 26 the seeds that showed the most promise: mammoth clover, red clover, alsike clover, tall oatgrass, orchard grass, timothy, and a "few other promising sorts not in the market." He concluded that a mixture rather than single species produced the best crop. Beal was also interested in developing methods to kill regenerating woody species in the agricultural field. Cutting "oak grubs" and other sprouts at the very base in July and August prevented their return, and mowing sweet fern and bracken fern close to the ground would kill these "very common and persistent" plants (Beal 1890). GRAYLING BEAL PLANTATION, HARTWICK PINES STATE PARK The Silvicultural work at the GAES was the first designed experiment to test the effects of soil preparation on the growth of a variety of native and exotic trees in North America. However, trees had been planted for the express purpose of reforestation since the early 18003 on the East Coast and for experimental purposes since the 18603 in the Midwest. In 1820, Zachariah Allen planted chestnut, oaks, hickories, and locust on 40 acres of a previously pastured hillside near Smithfield, Rhode Island (Sargent 1876). The seeds were planted in furrows 10 feet apart and carefully followed for at least 57 years. Around 1840, white pine seedlings dug from nearby woods were planted in furrows to reforest sand barrens in Massachusetts (Sargent 1886). In 1846, Richard S. Fay planted at least 400,000 imported and native trees over 200 acres of stony hillside near Lynn, Massachusetts, and, seven years later, J.S. Fay planted 35,000 imported trees, many native trees, and tree seed on 125 acres near Woods Hole, Massachusetts (Sargent 1876). After two decades, the brothers found Scots pine, European larch, and Corsican pine to 27 be most successful. In 1871, Bumet Landreth began planting his 7,000 privately-owned acres in Virginia with white pine, chestnut, catalpa, and other species (Rodgers 1991). While these previous efforts were made by private individuals with land and wealth, academic institutions did not become active in forest planting experiments until around 1867 when Thomas Jonathan Burrill investigated the “proper treatment of soils” for forest plantings at the Illinois Industrial University (now University of Illinois) (Rodgers 1991). In 1872, the Arnold Arboretum of Harvard College was established with Charles Sprague Sargent its first director to grow native and exotic trees (Rodgers 1991). In 1874, the Massachusetts Agricultural (State) College planted larches, Scots pine, and Austrian pine on a barren hillside (Rodgers 1991). In the early 18803, Joseph Lancaster Budd was experimenting with introduced tree species at the Iowa Agricultural College, and Kansas Agricultural College had two experiment stations that tested different trees to that State’s climate and soil (Rodgers 1991). Similar to its neighbors, University of Nebraska was interested in shelterbelt plantings and experimented with tree plantings well before the Montreal Congress of 1882 (Rodgers 1991). Finally, William Brown of Ontario Agricultural College at Guelph experimented with introduced species at its Experimental Farm in the early 18803 (Rodgers 1991). Although the plantings at GAES were certainly not the earliest experiments with native and exotic trees, they did occur earlier than the more famous Biltmore (1895, Schenck 1955), Harvard Forest (1907, Spurr 1907), and Fort Valley (C oconino) Experiment Station (1908, Gaines and Shaw 1959) experimental plantings. Furthermore, Beal did appear to be the first to explicitly design an experiment to test the effects of soil preparation on the survival and growth of native and exotic trees in North America. 28 The value of the early Silvicultural and agricultural efforts at the GAES was not recognized until later in the 20th century. The Public Domain Commission purchased the original 80 acres from the State Board of Agriculture in 1917 (Daw 1968). The following year, Filibert Roth (1919) visited the Plantation and recommended that the entire property be left undisturbed and used as an “object lesson” of a “forest and park” for visitors. However, the north 30 acres of orchards and woods were obtained by the Northeastern Michigan Development Bureau in an exchange in 1920 (Daw 1968) and now support a restaurant and wood products company. The south 20 acres of former agricultural test plots were obtained by the HA. Young Lumber Company in another exchange in 1964 (Daw 1968); this property was sold in August 1983 to Georgia-Pacific Corporation, who transferred ownership to Alro Steel Corporation in May 1998 (S. Seifert, Crawford County Equalization Department, personal communication). By 1968, the Michigan Department of Conservation (MDC) was interested in the value of the remaining 30 acres of the former GAES, now a part of the AuSable State Forest. Ted E. Daw (1968, Chief, Forestry Division, MDC) concluded that the plantation should be designated as an historical attraction and used in the Conservation Training School because the research provided the basis for the land use and reforestation policies for the State. The Plantation gained public recognition through a Michigan Forest Association (MFA) booklet showcasing Michigan's prominent trees (Moore and Botti 1976). However, the Plantation’s existence became threatened when Grayling’s City Manager, Jerry Morford, wanted to expand the City’s Industrial Park into the 30 acres still owned by the MDNR and containing the 5-acre Plantation. In a letter to Theodore 29 Tucker (Chief, Lands Division, MDNR) Morford (1977a) expressed interest in purchasing the 30 acres stating that the land offers “little current recreational or other value to the State of Michigan.” After consulting the Forestry Division (Tucker 1977a), Tucker replied that the City would need to demonstrate an “urgent need” to convert “the oldest (1888) recorded forest plantation in Michigan” to industrial park use (Tucker 1977b). Meanwhile, a devastating wind storm on 4 September 1977 blew down a number of the larger conifers in the Plantation (K. Gardiner, personal communication) resulting in a salvage cut approved by Weldon Montgomery (1977, Forest Management Division (FMD), MDNR) that Fall. Some of the logs were offered to MSU’s Forestry Club for the construction of a cabin but were not accepted (Morford 1978c). Many of the tipped stumps showing the salvage cut remain in the Plantation to this day. Citing the wind storm damage, lack of site identification, and lack of scientific value of the Plantation, Morford (1978a) lobbied Commissioner Harry Whiteley to convince the Natural Resources Commission to accept a 6-acre City-owned property located on the AuSable River in exchange for the 30 acres containing the Plantation. The State was initially interested in this offer of river frontage (Tucker 1978a) and began the process to consummate the even-trade exchange (Tucker 1978b, Schafer 1978). Support for the exchange grew within the Department as field staff were able to investigate the site. Noting that the 6-acre property where the East Branch and Main Stream of the AuSable River join is considered a “kids only” trout fishing location, Gary Schnicke (1978, District Fisheries Biologist, MDNR) recommended that the State acquire the property, especially considering the City’s lack of “protection or enhancement of natural 30 resources.” Raymond Perez (1978, Wildlife Habitat Biologist, MDNR) agreed that the protection of riparian wetland, which constitutes most of the 6 acres, for wildlife production should be considered and thus the land exchange approved; Perez did not assess the Plantation property. Although noting that the river property has no easement or legal description and is not contiguous with the existing Fish Hatchery, William Tarr (1978) recommended the land exchange with the City making up the difference in the far greater value of the State-owned 30 acres. He also pointed out that the City did not appear to have a great need for expanding into the Plantation property since only 6 of the Industrial Park’s l8 parcels had been sold to date. Representing the District, A.J. Coates (1978, District Forest Supervisor, MDNR) communicated to Robert Borak (Regional Forest Supervisor, MDNR) that the Plantation’s value is “negligible” but the river property should be protected against development. C. Troy Yoder (1978a, Regional Director, MDNR), on the other hand, agreed to the exchange if the City would include additional property and an easement to the river property but was opposed to the offer as submitted since the property was already protected from development by existing regulations. Fortunately, a City well is located on this additional property, so Morford was not interested in losing the easement (Yoder 1978b). However, some in the MDNR were concerned over the potential loss of the Beal Plantation. Weldon Montgomery contacted and requested Janet Kreger (1978, Regional Preservation Coordinator, Michigan History Division (MHD), Michigan Department of State) to assess the historical importance of the Plantation. In late March 1978, Kreger investigated the Plantation with Mac Collins (Architectural Coordinator. MHD) and 31 William Tarr (Area Forester, MDNR). She concluded that the Plantation should be protected and listed on the State Register of Historical Sites and that a State-owned buffer should be maintained around the Plantation (Kreger 1978). The President of the Michigan Forest Association (MFA), Wesley Manley (1978a), also caught wind of the proposed land exchange and expressed to Howard Tanner (Director. Natural Resources Commission) that the Beal Plantation represents significant research related to forestry in Michigan and should be preserved. Henry Webster (1978, Chief, FMD, MDNR) invited MFA to contribute “specific suggestions and active help” as well as bring MSU into the process of commemorating the research at the Plantation. Manley (1978b) promised to produce such a report as soon as an MFA committee had investigated the site. Conflict among different interests within the Department was brewing. C.D. Harris (1978, Chief, Bureau of Renewable Resource Management) complained to Wayne Tody (Deputy Director) that the F MD was dragging its feet on the proposed land exchange and brought to light that a FMD employee had notified the MHD and MFA about the proposal. He felt the “trees on the tract are not of any special value” and sought Tody’s approval to proceed with the land exchange. C.D. Harris had apparently not attended the November meeting where a one-acre reserve was proposed to be maintained for the Plantation, as only one acre was recognized to have survived to this time, and the other 29 acres would be considered in a land exchange (Morford 1978b); furthermore, this consideration was not recorded in the joint status report (Tucker and Webster 1978) as a result of the meeting attended by Jerry Morford, Walter Nowak (Manager, Grayling Chamber of Commerce), C.D. Harris, Theodore Tucker, Henry Webster, C. Troy Yoder, b) to and Weldon Montgomery. However, right before the MFA tour of the Plantation in December 1978, Weldon Montgomery phoned Jerry Morford to inform him that he was supporting retention of 5 acres for the Plantation and wanted to get MFA to support his proposal (Morford 1978b). While Morford (1978b) was certainly not satisfied with this proposal, MFA passed a resolution urging the FMD to retain the 5 acres (Clark 1979). Meanwhile, due to understaffing problems, the MHD postponed the processing of the application for the Plantation in the State Register of Historic Sites (Eckert 1979), and the Upper Great Lakes Regional Commission supported Grayling’s proposal to convert the 30 acres to industrial park (Rehberg 1979). However, Grayling Township, in which the 30-acre parcel is located, stated that the parcel is zoned Commercial as a buffer between Industrial and Residential property and should be placed on the market in an open bid process if the State chose to dispose of the property (Fowler 1979). By May 1979, the State was prepared to exchange all but 5 acres containing the Plantation with properties purchased by the City of Grayling based on the State’s selection (Schafer 1979). Concerned that the State’s offer did not include the 6-aere river property as the even exchange, Walt Nowak (1979) again lobbied Commissioner Harry Whitely to apply pressure to the MDNR. Even more brazen and close to the deadline for the land exchange agreement, Jerry Morford (1979) insisted on a meeting with the Natural Resources Commission to discuss the MDNR’s denial of the river property and full 30 acres as an exchange and inflation of assessed value for the 30 acres. In a stalwart yet refined response to Morford, Tucker (1980) reiterated the MDNR’s case: the river property was not deemed acceptable for exchange consideration; the Beal Plantation is of 33 historic significance but can be contained within a 5-acre reserve; lands purchased by the City for exchange must have the same appraised value as the 25 acres; the key decision has been on Grayling’s desk for over 7 months; and the only other alternative would be to dispose of the 25 acres by public sale procedure, as Grayling Township had suggested. Nine days later, Jerry Morford (1980) signed the proposed Land Exchange Agreement but still contended that the additional 5 acres were needed for industrial expansion. By February 1981, the State had selected private land8 to be purchased by the City (Harmes 1981), and the exchange was consummated on 4 May 1981 (Laylin 1981). However, Jerry Morford would not relent in his pursuit of the remaining 5 acres. In early July 1982, he requested the Department of Forestry (FOR) at MSU to evaluate the Beal Plantation with regard to its scientific values and to make suggestions as to alternative uses of the property. On 16 July, Dr. Victor Rudolph (Associate Chairman. FOR) and Jan Hacker (Graduate Assistant, FOR) made an on-site review of the Plantation with regard to its surrounding land use and prepared a report (Rudolph and Hacker 1982), which was submitted to Morford and the MDNR (Rudolph 1982). In the report, the authors concluded that the Plantation “no longer contains any scientific value to MSU, the DNR or the residents of the area” but does have some historical value. They suggested alternatives ranging from retention with no management to development and maintenance as a park to total conversion to industrial use. 8 That part of the NW '/4 NE '/4 lying E’ly ofthe AuSable River, T26N, R1W, Section 32. South Branch Township, Crawford County (Harmes 1981). 34 In the meantime, Janet Kreger (1983) restarted the process to nominate the Plantation to the State Register of Historic Sites, and Dean Sandell (1983, MDNR) prepared a development plan for the Plantation complete with parking, walking trail, and interpretive signs. A property survey ordered by Michael Moore (1983a) revealed that approximately one chain (66 ft) width of the Plantation was located on property owned by Georgia-Pacific Company (formerly owned by H.A. Young Lumber Company) to the south (Borak 1983). Taking advantage of Morford’s interest in gaining more land for the Industrial Park, Moore (1983b) requested Grayling to acquire this strip from Georgia- Pacific for another three-way exchange of land at the north end of the 5-acre parcel containing the Plantation. Meanwhile, the City of Grayling had overcome the Commercial zoning by the Township by annexing the property sometime before 1983 (Morford 1983). Although Morford (1984a) was more interested in gaining all 5 acres, he agreed to acquire a 60-ft strip from Georgia-Pacific in exchange for a l90-ft, or 2.9- acre, strip on the north side of the remaining 5-acre Plantation parcel (Morford 1984b), which Moore (1984) supported. Four years later, the MDNR was still waiting to acquire the acre owned by Georgia-Pacific and to develop Sandell’s vision of an interpretive facility at the Plantation (Botti 1988). Jerry Morford (1993) applied another surge of pressure to the MDNR for the entire 5 acres, probably as a result of an expansion planned by Monarch Millwork, Inc., north of the Plantation property (Hees 1993). Support for retaining ownership (McMillan 1993) and incorporating the Plantation into the Hartwick Pines State Park interpretive program (McMillan 1993, Dilts 1994) was high within the Department, but a compromise 35 was sought between the City, MDNR. and Monarch Millwork (Reuschel 1994a). As a result of an on-site meeting in late March 1994, Reuschel (1994b) agreed to a three-way exchange of an 80—ft swath of land on the north side of the 5-acre Plantation property for an 80-ft swath on the north side of Georgia-Pacific’s property to be acquired by Grayling. Four months later, the deal was negated as Georgia-Pacific expressed no interest in negotiating over their one-acre parcel due to financial stress at this plant and Monarch Millwork had begun expanding on their own property (Reuschel 1994c). The final push for protection of the Beal Plantation came in 1996 when Roger Rasmussen and Daniel Sikarskie. both from the Huron Pines Resource Conservation & Development Council (Huron Pines RC&D), were given the go-ahead from Gerald Thiede (State Forester, F MD, MDNR) to organize a Beal Plantation Committee (R. Rasmussen, personal communication). Persons and agencies who might be interested in preserving and interpreting this portion of Michigan’s early forest history were invited to an on-site meeting on 15 October 1996 (McMillan and Rasmussen 1996). Eleven persons, including Dr. Frank Telewski (MSU), attended this first planning meeting and determined that a property survey was needed, administrative authority between F MD or Parks and Recreation Division (PRD) should be decided, Ron Nagel (PRD) would develop an interpretive plan, Frank Telewski would complete an inventory of the Plantation, funding would be needed to implement the interpretive plan, and publicity was needed. The MDNR approved of the Committee’s direction and left the decision of which Division to administer the site up to the local Division managers (Reuschel 1996). The survey showed that “at least two rows of the largest trees are on Georgia-Pacific property” (Rasmussen 1997), so the Committee requested the donation of the l90-ft wide swath of Georgia-Pacific property to the State (Rasmussen 1998). By June 1998, Georgia-Pacific had donated the 2.7-acre parcel (Thiel and Sikarskie 1999). The Beal Plantation Committee gained the needed momentum with the land donation to begin planning the interpretive facilities. In August 1998, I joined the project and began an historical and ecological survey of the Plantation as well as participated in the planning process for the facility. This rustic yet moderately accessible site would feature a walking trail with interpretive signs and be linked to Hartwick Pines and North Higgins Lake State Parks by an auto tour. While persons from a number of organizations contributed to the Beal Plantation project (Table 1-2), Huron Pines RC&D provided the key coordination and leadership to keep the project moving. An over 1 100-ft handicap-accessible crushed gravel trail and crushed gravel parking lot was designed by Frank Telewski, Ann Stephens (PRD, MDNR), and me and installed by youths from the Shawono Center (Grayling, MI). As the trail winds through the Plantation, four interpretive nodes with wheelchair pullouts describe the site history (Figure 1-14) and role of the FMD in multiple use management of forest resources. The interpretive signs were written and designed by Earl Wolf (Office of Information Services, MDNR), Frank Telewski, Ann Stephens. and me. Construction materials were obtained by Huron Pines RC&D through grants from the Michigan Department of Agriculture and The Weyerhauser Company Foundation. A.J.D. Forest Products Company donated and installed three sturdy wooden benches along the trail. The interpretive facility is administered by the FMD and sponsored by 37 Kirtland Community College under the Michigan Adopt-A-Forest Program. On 7 October 1999, the Beal Plantation was formally dedicated as the birthplace of reforestation in Michigan. The Beal Plantation is an important historical and auto tour stop for forest visitors between the Hartwick Pines Logging Museum and Interpretive Center and the old Forest Nursery Site at North Higgins Lake State Park (Thiel and Sikarskie 1999). Visitors to the Plantation are impressed by the low impact on the site integrity, diversity and size of trees, and excellent interpretation of Michigan’s early forest history. Table 1-2. List of participants in the Beal Plantation Committee, which was formed in Fall 1996 to protect and recognize the historical importance of the Grayling Beal Plantation. Coordination was provided by Huron Pines Resource Conservation and Development Council. Representative(s) Organization Roger Rasmussen Daniel Sikarskie Dr. FriiikTéléWéki"W ” " ” ' ” Jason Kilgore Ron Nagel Ann Stephens “meMlenan .. Susan Thiel Denise Kemp ’1 Jerry Meyer Lynn Porritt- McConnell Dave Stephenson 1. ~ Paul Callm Herb Burkett Huron Pines Resource Conservation and Development Council W"). ‘BearBotanicar‘ear‘d‘e‘a‘aha‘caapu’g‘xrsorema”be’partae'm““‘ ., of Botany and Plant Pathology, Michigan State UnIverSIty .. Parks and Recreation Divisi6n, Michigan Department of Natural Resources Forest Management Division. Michigan Department of Natural Resources Natural SCIenCC Department Kirtland Communlt) COIIL Cgé: ' " ' Grayling Regional Chamber of Commerce A. J. D Forest Products Company ” ‘ Weverhaeuser Company .. .. ‘ - ‘ Michigan Department ongrIculture . . . . Charlie Guenther ’ i I Private Citizen 38 Ago: 36:63 Eat :85 gas .285 .9532 was wmEB< Dubai—5 35m 53:22 Co $3.58 samumouozm docs—=2 33c 95% :2: 208 2t: .32 380 E =2§m EoEtumxm _EB_=6_&< wE—xEO 2: .6.“ 5:82 05 on 32.5 Fem 336.35 5:? .N-_ onE 40 6.530230 BEBE: 6% 83:02 bioficb 88m 53:22 .wo $2.58 camcwouonm .30? £5 .wo 5w: 05 3 vuamznfimo on E? 53853 05 CoEoQ “83:58 05 9.: mixes 5:853 ozCo ammo Hmquoc coo—S m5: namcwouofi 2:. due Eon—qua Eat 3:053. xao 32:8 98 uni x8.“ postman FEB “.298 mm? .cawEEE .3550 38380 E 523m EoEtoaxm 2: Low war—mo? 9 hotn— US: mg .m-_ 2sz .fi nix/3o) 3.2.. 9:2. 5.2. .353 «9.43. $3.56.. 3 a.» A)...” gm 3:» 5 15.53.? Mo _. .335...» 5.x . __ 5...... J 41 8:280:00 Eur—085 a. mo>Eou< b85>ED 88w :amE22 mo 8258 :mmaouosm .m-_ 2:me E :08 on 8?. =8 5:?» .8: 05¢ v2 28 can. 8:0.“ cunmEmg 05 802 .mEEB 30.653 Ea wage—o can. 05 8 5.5: mac—02 :omgm EoEtoaxm 05 ,«o 5:8 385 82:2 2: Spa :87: m9: :meSonm £5. .mEoEtogxo .8332th 8m 3.835 EB .3393 .3520 20>» 3.5m om F8538 05 can €8un 225 was Bum—Sc 2: we mean ow 505:8 2:. .v; 2sz 42 Road (Industrial Parkway) North Field & Orchard (20 acres) Wooded Area Control W \ k \ Harrow Pl____ow 6— 660 ’ —-> South Field (20 acres) 43 2 Figure l-5. Layout for the 80-acre Grayling Agricultural Experiment Station as reconstructed from various sources. The Silvicultural experiment contained the l-acre Plow Treatment, 2-acre Harrow Treatment, and 0.4-acre Control. The end widths and length of the Harrow Treatment are known, but the actual orientation of the north and south boundaries is unknown; since several surviving trees border the Plow Treatment in a line, the location of the north boundary (dashed) is less known. The large arrow represents the approximate location and direction for the photograph shown in Figure l—3. Amman: Rom Set $55.58 235 .8—3 cocoom 520m .355: 2: 80¢ way was :52: 2: 223 £255 2: .5 $58 803538 2: 5 555:8 2: .3 E Ewsofi 053 5835 Ea :mm .355th Bfiztfi o>c SE F536 :2: 8.5 523 .85 28-25 m :o :38 was $508 scam 526% 505555 _E=::otw< wczbuo 2: .5 855 cm 5:8 05 Sm 2385 23:08:5me .0; Ezwi .MH UM 05 Cu m5 0“— O E ME QM 51H .55 w 55 B .m£ com .5 82m as. we 9 a: 8m :3 e5. .5: 8m 8 ENE .83. m5 2 a: com :3 .M .w H5: we 0 Z 'Imxos MOPBE’W anasag mopeaw $5913 3&1 [eiuuajad 'anosad [[91 'SSBJD pmrpxo 'ssmf) anlg 'A)1 'sself) uepeBunH 'surdn'] monak 'Kmowu. qeaqlvopng 'a/(H seed Plaid 'pxmsnw am] M 'qaia A 35913 uepeBunH IBAOID amsw 'JaAoD peg PHEJIV binds 'JaAoD [powwow .Z 53.5.53 8: :5 338.52.“ m: $5323: 33 £58 2: I 3.3 .H 3:: 3:» R. 3558 Stakmfim «5.x 3: Rs 8.53 cm 5:8 55 I $95336 .5 Emmk~5§5tm§xm 44 .83: 035! Set 58:5» 053m .23 5:8 05 8» mm 0E9. 05 05>» BEEN“ 05 Bu 8958 632.22: mag 05 he :ocfim EuEtuaxm _a._3_=utw< wszzfio EEO 3.6m on 5.5: 05 Sm cwuou 5585me K.— 2:3; .2803 £32 SHE .83»? .32 wan .esumwm BM .Emm .Emm .30.. thm daemon 30682 .bham .Emm .80.. mBXm 633m .mmwoam .Eamm .380 Saws: due .KXN .38...» Rafi .Eum 3:3 .38. $8 5320 8928 ..$>oU 8538 .635 «Saxon .39 wan gag—U 98 $80 PE .555 505532 9:5... 2.33 98 ago—U Ex .30.. wnXu .AOH “am can $.80 vamp—BO .§w> warn—m .Esmm .30.. wan doe Em Ea ESE: .Fsmm .mfiiusm $9 wmxmxomxa mm a % . . z . w 6% .Ea . 2 OH wwm “Ed 5055. «dam u m $5.. mmXH 30m .3 3m .23ch 262 m an egg doe Rm 93 >595 .23 $595 .39 we: do m cm A on: .93 . «w main .m 0.. .Su m .u 5 L. A 5 > . m v mem 6me 6me .32 .35 6%" .08“ dwww dar— u0 in $3» 855 tokmfiuuum .m2u< 35.35%. 3mm £82 45 .2280on 78:82: a. 32:02 biota: 8me 53:22 no 3258 :nEwSonm .38 C ozwom 3 38: mm 693% 32:02 98 05a BE? KEEP. 2: 202 .82 «mawsax ,Eoc 35:52me 20 05 Set :ozflcflm 05 .«o EoEEEH >65 05.3 can “mo? 2: 25 32> .w; Ezwi 46 .555ch 8me 53:22 imam—uh .m Mo @258 games: 4an E5 mo ammo Eve 3523:? 20 2: E :oEEoQ 2m 320538; :32 E85 docfiocowfl v.8 we can .333 223 .oEa Eon.“ can 38m 3 BEEEou E EEBMEQ 2: E Em: .3333th So 2: 2E? .ucsemxumn of E 382% big on :8 82% maioz Ea moEn E: was 823 .5382; 2:. .28. E 53 mm .cocficmE Gum wEEEO 2F d; 22%; 47 THE EXPERIMENTAL PLATS. Where a plat is marked as containing nine sorts there are the following: Orchard grass, Perennial rye grass, June grass, Meadow foxtail, Red clover, Mammoth clover, Alsike clover, Timothy, Meadow fescue. N.——— S. No plowing, no harrowing; nine sorts, mixed. Sown May 15. This plat only harrowed, not plowed; nine mixed sorts. Sown May 15. Blank. Field Peas. Sown May 15. Spring rye. Sown May 15. Nine sorts, mixed. Sown May 15. Perennial Rye Grass. " ' . S ' . , Mammoth Clover, mixed, Perennial R) e Grass own Ma) 15 Perennial Rye Grass, Orchard Grass, Italian Rye Grass. Sown May 15. Mammoth Clover, Orchard Grass. Sown May '15. Orchard Grass, mixed, Mammoth Clover. Sown May '15. June Grass. Sown May '15. Alsike Clover. Sown May '15. Meadown Foxtail. Sown May 15. Alfalfa, or Lucerne, with a little Mammoth Clover on one side. Sown May 15. Hungarian Grass. Sown May 15. Timothy. Sown May 15. Millet. Sown May 15. Meadow Fescue. Sown May 15. Sweet Clover. Sown May 15. Spurry. Sown May 15. Red Clover. Sown May 15. Figure 1-10. Experimental design for the agricultural plots at the Oscoda (losco County) substation, which contained 10 acres deeded by Mr. James Barlow. Each plat was 3 rods wide by 10 rods long (8167.5 ft2 or 0.1875 acre). Figure redrawn from Beal (l888b). 48 Saw—03:3 29m EwEEE Ewan—o... .n— .«o Amer—=8 Emewoaonm awoken 3:282 83:82-55: .Eoumhm :8... Si 035 05 2 325g 2: “a 3302 Boa £03835 £880 2: Eat wEEmEE moEn tom ._ _-_ 2sz 49 26. - Hungarian. 25. — Rye. Sown about 27.- Blank. Plowed not Sown about May May 20. Sown to sown. 20. Sown to red red clover late in clover late in Aug. August. 23. - Millet. Sown about 24.—-Meadow fescue. May 20. Sown to 22.-Alsf(a)l:a. b tM' ’ Sown about May 20. red clover late in 20 n a ou d) August. ' 21.-Alsike clover. Sown 20.--]une grass. Sown 19. — Eight sorts. Sown about May 20. May 11. May 11. 9. — Left wild. 18. —Timothy. Sown 17. — Italian rye grass. 'l6.-Spurry. Sown May May 11. Sown May 11. 11. 13.-Field peas. Sown 15. —Perennial rye grass. 14.— Meadow foxtail, May 11. Sown to Sown May 11. Sown May ’1']. orchard grass late in August. 12. — Mammoth clover. 1 1. — Red clover. Sown 10. — Sweet clover. Sown May 11. May 11. Sown May 11.. 5. —- Unplowed, not har- 8.—Orchard grass. 7. -Tall oat grass. 6. —Orchard grass. rowed. Seven Sown May 11. Sown May 11. Sown May 11. sorts. Sown May 11. 3. —Tall oat grass and 2. — Mammoth clover ’1 ' _ Unplowed, but har- 4.—Orchard grass. rowed. Seven Sown Ma 11 orchard grass. and orchard grass. rts So M , y ° Sown May11. Sown May 11. :2 ' wn ay Figure 1-12. Experimental design for the agricultural plots at the Walton (Grand Traverse County) substation on 5 acres rented from Abram F. Philips. Each plat was 3 rods wide by 10 rods long (8167.5 ft2 or 0.1875 acre). Figure redrawn from Beal (188%). 50 s... , a .. .Mwuaaufl .3. .9,” I r m. WV. Figure l-l3. Red pines remaining from the Harrison Substation, now located on private property just north of Budd Lake. Photograph courtesy of F. Telewski. Michigan State University. 51 .b_m5>_=3 38m $3222 dam—E ._. . - w .3 =83 ifiwoaonm 23m 35m 85m x2353 of no 555 5 26: 50:85.; .95 wEREO 05 8 ES u>cEEEE 2:53 2: 8 x83 Bantam E _ 2: E {Eli P .1. it. 5211.5 .: a AREA-xi... tit!» $119...» zllilx..li...ctt!:!x§ 1 51.7.91. . it. artilii 5.5.1.: 5.6:: till-{ti , fiffiiiiillil. l 3.1 (1:... a: [limititti III. 1.1 x1331... til-‘1. itsiiisrsitxllu; itililtuytl... I. It?! . til-(iii)... iliyti‘lttéttiti v.3 5.1.2. tiltiiiiitsfi f3§§t11! .8 4. so 52 CHAPTER 2 RESULTS FROM A LONG-TERM REF ORESTATION EXPERIMENT IN THE PINE BARRENS OF NORTHERN MICHIGAN ABSTRACT The establishment, growth, and regeneration of 3 locally native, 22 regionally native, and 16 nonnative (exotic to North America) tree species were measured in a common garden experiment established in 1888 on the pine barrens of northern Lower Michigan. Species with the greatest survival of original stems were Pinus resinosa Ait. (35%), Picea abies (L.) Karst. (24%), and Pinus strobus L. (7%); several stems of Juniperus virginiana L., Picea glauca (Moench) A. Voss, Pinus rigida Miller, and Thuja occidentalis L. also survived. Regeneration was dominated by Pinus strobus, Pinus resinosa, Picea abies, and Pinus sylvestris L. in the plantation and by Pinus strobus, unplanted P. banksiana Lamb, P. sylvestris, and unplanted Quercus spp. in the old field next to the plantation. No originally planted hardwoods survived, but regeneration from Prunus serotina Ehrh., Populus alba L., Robinia pseudoacacia L., F raxinus americana L., Acer rubrum L., and A. platanoides L. was present in the plantation and/or old field next to the plantation. Site index values reflect the low productivity of the site: Picea abies, 28; P. glauca. 29; Pinus resinosa, 44; and P. strobus, 45. 53 The nonnative Pinus .sylvestris and Picea abies performed as well as or better than species native to the pine barrens. Pinus .sylvestris is successfully spreading out from the original plantation and competing with the native early successional species P. banksiana and P. strobus. Within the plantation, Picea abies survived and produced as much regeneration as the native dominant Pinus strobus. The successful colonization by nonnative Pinus .sylvestris and Picea abies indicates that both have the potential to become invasive in the pine barren ecosystem. OBJECTIVES The objective of this chapter is to report the current community composition and structure of the Grayling Beal Plantation, which was established as an experiment in 1888 by Professor Beal to determine which trees species would survive and regenerate in the Pine Barrens of northern Michigan. This chapter can provide ecological data useful to investigators interested in the survival and growth of a variety of native and exotic trees species in this pine barren ecosystem. INTRODUCTION Reforestation of anthropogenically affected landscapes is recognized as important for sustainable land management by government (United Nations 1995, United Nations Division for Sustainable Development 2000), academia (Keddy and Drummond 1996), and industry alike (American Forest and Paper Association 2001). The question of which 54 native or nonnative species to use for reforestation was addressed early in Europe (Pontey 1828) as well as in the United States (Sargent 1876) and continues to be an issue for land managers. The ability of a species to survive and successfully regenerate determines the species assemblage of a reforested area. Furthermore, tree species not native to the reforested area could affect the ultimate composition, structure, disturbance regime, and functional attributes of the community (Richardson and Higgins 1998). The pine barren ecosystem is an often overlooked system that has been drastically affected by human disturbance. Traditionally, a barren was described as an unproductive community lacking canopy trees and dominated by grasses, forbs, shrubs, and scattered stands of trees (Curtis 1959, Anderson and Bowles 1999). Barrens are generally regulated by edaphic conditions and natural disturbances, especially fire, resulting in a particular flora, often containing a number of endemic species. For example, the shale barrens of the Appalachian region form on highly weathered shale outcrops that experience active erosion and extremely warm midday temperatures (up to 60° C) and support an open canopy forest of Pinus virginiana, Quercus prinus, Q. rubra, Q. ilicifolia, and Juniperus virginiana; at least 18 vascular plant species are shale barren endemics (Braunschweig et a1. 1999). Eastern serpentine outcrops also produce barrens dominated by xerophytic graminoids and herbs and fire-tolerant P. virginiana. P. rigida, and J. virginiana, species that can tolerate the low calcium to magnesium ratio, low water availability, and high heavy metal (e. g., nickel and chromium) concentrations (Tyndall and Hull 1999). Perhaps the most studied pine barren ecosystem in North America is the barrens of the New Jersey Pine Plains (Forman 1998). Dominated by stunted P. rigida 55 and Q. marilandica, these barrens are underlain by deep, acidic, and highly leached sand and are structured by short (<20 yr) fire return intervals (Gibson et a1. 1999). The pine barren ecosystem of the northern Great Lakes region ofNorth America occurs on post-glacial outwash plains, sand lake plains, and sandy riverine terraces. The development and composition of forests on these glacial features prior to European settlement have been summarized by Curtis (1971), Whitney (1986, 1987), Mokma and Vance (1989), Graumlich and Davis (1993), Barrett et a1. (1995), Schaetzl and Brown (1996), Radeloff et a1. (1999), and Zhang et a1. (2000). In Michigan, the pine barren forests were dominated by Pinus banksiana (Roth 1905) with elements of P. resinosa, P. strobus, Quercus ellipsoidalis, Prunus serotina, and Populus spp. (Whitney 1986, 1987, Comer 1996). After the merchantable timber was logged in the second half of the 19th century and numerous wildfires spread across northern Michigan, the pine barrens regenerated to forests of Pinus, remained stump prairies (Barrett and Schaetzl 1998), or were planted to Pinus resinosa, P. strobus, P. banskiana, or P. sylvestris during the 193 Os (Curtis 1971, Whitney 1987). Historical and experimental research has shown that edaphic factors and disturbance. especially fire, are important for maintenance and restoration of the pine barren ecosystem (Simard and Blank 1982, Abrams and Dickmann 1984, Whitney 1986, Host et a1. 1988, Host and Pregitzer 1992, Vora 1993, Little 1998, Pregitzer and Saunders 1999). However, forest management practices such as fire suppression and monotype plantations have resulted in the destruction or conversion of all but 7% of presettlement barrens in the Great Lakes region (Whitney 1987. Temple 1995) 56 The effects of management and natural regeneration on the pine barrens are thus well documented, but no studies exist on the survival and successional outcome of a mixture of planted tree species in this ecosystem. The former Grayling Agricultural Experiment Station (GAES) in northern lower Michigan provides a unique opportunity to address this issue. In 1887, Michigan’s legislature directed the Independent Forestry Commission to survey the condition of the State’s remaining forests following extensive logging, rampant wildfires, settlement. and agriculture (Reynolds l888a, Telewski 1998). In the following year, the US. Congress passed the Hatch Act, which provided $15000 to every land grant institution to conduct experiments in agriculture and cognate sciences (Beal 1915). Using the Hatch funds, Michigan Agricultural College (now Michigan State University) established the GAES to determine how to reforest and increase agricultural productivity in the pine barrens of northern lower Michigan. As part of this effort, Professor William J. Beal established a test planting of 41 native and normative tree species to determine which would survive and reproduce in this ecosystem (Beal 1888, 1889a). The results of this test plantation from 1888 have not been thoroughly evaluated until now. The purpose of this study was to complete an historic Silvicultural experiment, viz. determine which tree species became established and their success at regenerating at the former GAES in northern lower Michigan. Given the environmental limitations (e.g., low pH, nutrients, and water holding capacity) to successful tree establishment in this ecosystem, I predicted that species native to the pine barrens would be better adapted to survive and regenerate than nonnative species. 57 METHODS Study site The investigation took place between August 1998 and October 2000 at the Grayling Beal Plantation (Hartwick Pines State Park annex) in Crawford C ounty, Michigan (44°39‘N, 84°42’W, elev. 348 m; Figure 2-1). This area became deglaciated about 12500 years ago (Weirlein 1998) and is located geomorphologically within the Grayling Outwash Plain (Albert 1994) and fioristically within the Great Lakes Pine Forest (Kiichler 1964). The soils at the plantation are a coarse, loose, strongly acidic, and excessively drained sand of the Grayling series (Typic, frigid Udipsamment) (Mokma and Vance 1989, Werlein 1998). Government Land Office Survey records show that presettlement forests on this soil type were largely composed of Pinus banksiana (Whitney 1986). Prior to planting in 1888. the site had experienced frequent fires leaving mostly P. banks'iana. scattered P. resinosa, and scrubby Quercus spp. and Prunus spp. (Kedzie 1888b), but no record or evidence of fire exists at this site since planting. The climate of Crawford County is characterized as humid continental (Whitney 1986). The mean annual precipitation (1951-1980) is 81 1 mm with 62% of the precipitation falling during April-September (Figure 2-2). The mean annual temperature is 6.3° C with a mean summer temperature of 18.60 C and a frostfree growing season of 110 days (Weirlein 1998). The growing season is generally cool and short with only 2068 growing degree days (Weirlein 1998). 58 Beal’s experiment at the plantation was designed to test the effects of site preparation (plowing or harrowing) on survivorship, growth, and reproduction of 40 native and normative tree species (Beal 1888, Anonymous 1913) (Appendix I). In May 1888, Beal planted an arbitrary mix of 5260 hardwood and conifer seedlings and cuttings on 4-foot (1.2 111) centers in rows running west-east. The plowed treatment was 1 acre (0.40 ha), the harrowed treatment was 2 acres (0.81 ha), and the control (no site preparation) was 0.4 acre (0.16 ha) in size. In the following year, the harrowed treatment was plowed under, and an unknown number of Pinus strobus and Picea abies seedlings were planted in the plowed treatment (Anonymous 1913). Beal (1889a) also planted seeds of Pinus rigida in a couple locations in early 1889. Unfortunately, Beal’s original notes (Anonymous 1913) are not sufficiently detailed to determine the original number of trees planted in each treatment, thus the site preparation treatments were not compared. Subsequent management, alteration, and monitoring (see Chapter 1) of the plantation have been minimal prior to the present. Survivorship and growth Using historical notes, I located, flagged, and measured all surviving planted trees. Survivorship was resolved simply by the presence or absence of stems purportedly planted in 1888-89 as determined by a stem’s relative size and location within an apparent row. Each surviving stem was given a unique alphanumeric aluminum tag, and height and diameter at breast height (dbh; breast height = 1.37 m) were measured using an electronic hypsometer (Vertex Forestor) and diameter tape, respectively. 59 The allometric aboveground biomass and site index were estimated for selected surviving Pinaceae only, including Picea abies, Picea glauca, Pinus resinosa, and Pinus strobus. Standard allometric biomass equations available for forest stands in the geographically nearest region were used to calculate biomass (Table 2-1) for each tree and site index (Table 2-2) for each species. Table 2-1. Allometric biomass equations used to estimate total aboveground biomass (AB), including foliage and branches, of individual surviving Pinaccae at Grayling Agricultural Experiment Station (Ter- Mikaelian and Korzukhin I997). Diameter Species Biomass Equation (Kg) Range (cm) Region Source Picea abies AB = 0.2722*dbh"2.1040 12-44 New York Jokela et ul. 1986 Picea glauca AB = O.0777*dbhA2.4720 1-33 Minnesota Harding and Grigal I985 Pinus resinosa AB = 0.0778*dbh’\2.4 I 71 3-46 Upper Great Lakes Perala and Alban 1994 Pinus strobus AB = 0.0755*dbh"2.3833 5-26 Upper Great Lakes Perala and Alban I994 Table 2-2. Equations used to estimate site index for surviving Pinaccue based on mean height at Grayling Agricultural Experiment Station (Carmean er al. 1989). Species Site Index (SI) Equation Year Range Region Source Picea abies SI = (0.0259)(ht)’\( l .2496)( I -expA Planting—60 yr Wisconsin Wilde et a1. (-0.0021)(1 12 yr))"(l .7841 )(ht"(—0. 1088)) I965 Picea glauca SI = (0.0380)(ht)’\(1.5142)( l—exp" 2()-—l30 yr Minnesota Carmean and (-0.0124)(1 l4 yr))"(-6.484)(ht"(-0.355)) Hahn I981 Pinus resinosa SI = (0.52919)(ht)( 1 -exp" 20--—120 yr Minnesota Gevorkiantz (-0.0198)(l I4 yr))”‘(-I .3892) I957a Pinus strobus SI = (0.5086)(ht)(l-exp" 20- 120 yr Northern Gevorkiantz (-0.024)(l I4 yr))A(-l .8942) Wisconsin 1957b To measure average radial growth. trees of the four selected Pinaccae and regenerating Pinus .sylvcstris were cored perpendicular to the stem lean to avoid reaction wood with 5.15 mm diameter increment borers. Cores were obtained at approximately 0.3 m from the ground to obtain as many growth-rings as possible. Increment cores were prepared, cross-dated, and measured using standard dendrochronological techniques 60 (Stokes and Smiley 1967). Ring-width measurements were analyzed to detect anomalies within each time series and to obtain descriptive statistics for each species with the program COFECHA Version 6.06P (Holmes 2000). Demography The current stand structure reflects the survival and regeneration of the originally planted trees. Since the trees were planted non-randomly in rows parallel to the treatments, I divided the plantation into four equal widths perpendicular to the planting rows and randomly selected two 2 m wide belt transects within each quadrant (Figure 2- 1). The remaining portion of the old agricultural field associated with the original GAES and abandoned by 1900 was also sampled to determine which woody species were migrating into open areas as opposed to establishing under the plantation cover. Each of the eight belt transects was approximately 143 m long and covered 0.02 ha. thus approximately 8.4% of the 5-acre (2.02 ha) area encompassing the plantation was sampled. The position of all woody plants (except Vaccinium spp. and Comptonia peregrina) with stems within the belt transects was recorded relative to the transect tape. The height, linear crown cover within the belt transect, and dbh for each stem were measured. The number of stems for each species within a belt transect was grouped by diameter size class and by location (plantation versus old field), standardized to a hectare basis, and grouped across all belt transects. Frequency was calculated as the number of 2 61 m long plots in which a species was present within the 2 m wide belt transects. To capture the relative contribution of each species in the plantation community, an Importance Value (IV) was calculated based on the sum of relative frequency, relative density, and relative [height*cover] by transect. The index [height*cover] was used instead of other units of coverage (e.g.. basal area) to include stems less than 1.37 m tall and to incorporate the effect of light interception; [height*cover] creates an apprOpriate dimensional occupation of space for each stem. However, since understory trees are more likely to have relatively horizontal rather than vertical canopies to optimize light capture (Oliver and Larson 1996), this index may disproportionately weight cover from understory stems. A Success Index was calculated as the IV for a species divided by the number of seedlings originally planted to compare relative success of each species based on original presence. .S'tatistical Analyses Means and standard deviations for the height, dbh, and biomass of each species were calculated in Systat Version 9.01 (SPSS Science Inc. 1998), while the mean ring- width and standard deviation were calculated in the program RESPO Version 6.06P (Holmes 2000). Differences in growth among originally planted species of select Pinaceae were tested using species as the fixed effect in a one-way analysis of variance for each growth category (height, DBH. biomass, and ring-width) in Systat Version 9.01 (SPSS Science Inc. 1998). 62 RESULTS Survivorship and growth Of 41 tree species originally planted as seedlings, cuttings. or seeds in 1888-1889, none of the original stems from the 32 hardwood species survived, but seven of nine conifer species had stems surviving to 2000 (Appendix 1). Of these seven species, only Pinus resinosa (34.5% survival) and P. strobus (8.4% survival) are considered to be native to the Great Lakes pine barrens. Picea abies (26% survival) is exotic to North America, Pinus rigida (2 stems from an unknown number of seeds) is native to eastern North America, and Picea glauca (4% survival), Juniperus virginiana (3% survival). and Thuja occidentalis (1% survival) are native to this region but not to this forest type (Appendix 1). Two of the conifer species that did not survive were Larix decidua and Pinus sylvestris; the latter is present as regeneration in the plantation and old field. Several stems of Acer rubrum. Fruximts spp., Juniperus virginicma. and Thu/a occidentalis are present as stump sprouts within the plantation, and Popular alba, Przmus serotina, and Robinia pseudoacacia are present as root sprouts in the plantation and old field. Survivorship among species was not statistically compared because trees were not planted randomly within each treatment and because postplanting activities selectively removed trees from particular species, as described in Methods. Of the surviving Pinaceue, those species native to these pine barrens demonstrated greater mean growth than all other Pinaceae examined in this study (Table 2-3). Pinus strobus attained the greatest mean height and dbh and had a mean ring-width similar to that of Pinus resinosa. but its aboveground biomass was similar to that of the exotic Picea abies. Based on its small surviving population, Picea glauca, which is native to the region but not to the pine barrens, expressed stunted growth relative to the native Pinaceae in this study. Growth of the two native species was more similar to each other and greater than the normative species (Table 2-3). Table 2-3. Mean size and growth statistics [i1 SD] for originally planted and surviving Pinaceae at the former Grayling Agricultural Experiment Station, Crawford County, MI. Species with the same letter do not differ significantly (Tukey HSD, p<0.05). Trees Height DBH Aboveground Site Trees/ Series Ring- Species (n) (m) (cm) Biomass (Kg) Index Cores (n) Length (yr) Width (mm) Picea abies 24 19.5 i 35.9 i 545 i 292a 28 II/ 25 92.3 :t 8 1.79 :t 0.5ab 3.7a 9.2a Picea glauca 2 18.3 d: 26.6 i 266 i 112a 29 2 / 4 74.5 i 28 1.35 :t 0.1a 2.5abc 4.7a Pinus resinosa 69 21.8 i 40.9 i 636 i 217a 44 48 / 97 82.6 i: 14 2.18 i 0.5c 3.3b 6.2b Pinus strobus 37 23.8 i 42.0 :1: 580 :t 219a 45 10 / 21 89.7 i 13 2.17 i 0.7bc 3.7c 6.7b Demography A total of 2301 stems from 20 woody species (except Vaccinium spp. and Comptonia peregrina) were documented within the belt transects. Approximately 72% and 92% of all woody stems in the plantation and old field, respectively, were seedlings (less than 1.37 m tall). Species originally planted at the GAES accounted for 38% of all woody stems and 24% of stems greater than 5 cm dbh in the plantation (Appendix II), and accounted for 13% of all woody stems and 53% of stems greater than 5cm dbh in the old field (Appendix III). 64 Only Pinus strobus (plantation and old field), P. sylvestris (old field), and Quercus rubra-group (plantation) (Figure 2-3) exhibited the reverse-J shape diameter distribution characteristic of old growth stands (Oliver and Larson 1996). The other species showed no presence in the plantation or old field (e. g., P. rigida), presence as juvenile trees only (e.g., Populus alba in old field), presence as large trees only (e. g., Q. ellipsoidalis in old field), or presence as a relatively uniform size class (e.g., Acer rubrum in plantation). Of the originally planted Pinaceae, only Pinus strobus had increasing numbers of trees in the smaller size classes for the plantation and old field. Picea abies experienced a reduction in the smallest size class (0-4.9 cm dbh) in both communities, while Pinus resinosa increased in density in the same size class in the plantation. Pinus sylvestris had a relatively flat distribution in the plantation but increased in abundance in the smaller classes in the old field. The majority of the non-seedling component of both the plantation and old field was of species native to these pine barrens (Figure 2-4). Pinus strobus clearly had the highest number of saplings (<20 cm dbh) and large trees (>20 cm dbh) in the plantation, followed by the subdominants P. resinosa. P. .sylvestris, Q. ruhra-group', P. banksiana. and Picea abies, in order of stem prevalence. The relative density of large trees in the old field was partitioned among the native species Pinus banksiana and Q. ellipsoidalis and the exotic P. sylvestris. The advanced regeneration stratum in the old field was ' Due to the presence of adult Quercus ruhra, Q. ellipsoidalis, and possible hybrids between the two related species (Overlease 1964, Tomlinson et al. 2000), seedlings of the Quercus section of Lobatae were lumped as Q. rubra-group. 65 dominated by P. banksiana, P. strobus, and P. .sylvestris. Species native to these pine barrens accounted for the majority of the total Importance Values (Tables 2-4,5, Figure 2-5). In the plantation, 78% of the total IV was from species native to the jack pine barrens. and the top four species accounted for 64% of the total IV. The exotics Picea abies and Pinus sylvestris accounted for 9% of the total IV, while the remaining 14 species contributed less than 5% each to the total IV. In the old field, 68% of the total IV was from species native to the pine barrens (Table 2-5). The large number (672, or 82% of all old field stems) of Quercus rubra-group seedlings from a recent mast year (personal observation) contributed heavily to the extraordinarily high IV for this species. The exotics Picea abies, Pinus sylvestris, and Populus alba accounted for 21% of the total IV in the old field. The Success Index (SI) indicates the relative success of each species as a function of the known number of stems originally planted in the plantation and its IV as measured in 2000. In the plantation, planted species native to the pine barrens (Table 2-4; 81:3 5% of summed SI for all species) were less successful than the other species (Sl=65%). Picea abies, Pinus strobus, and Prunus serotina were most successful, followed closely by Pinus resinosa. In the old field, species exotic to North America (Table 2-5; SI=71% of summed SI for all species) were more successful than planted species native to the region (SI=13%) and pine barrens (SI=16%). The exotic P. .sylvestris had an S1 over five times higher than the next species, the exotic Picea abies. 66 Table 2-4. Rank in dominance by Importance Value of woody species and their Success Index in the plantation at the Grayling Agricultural Experiment Station, Crawford County, MI. Status indicates whether the species is considered to be Native (N) to these pine barrens, native to the Region (R) but not the forest type, or Exotic (E) to North America. Number planted based on Beal’s notes for 1888 (Anonymous 1913). Despite the strong presence of Pinus strobus, note the similar relative success of Picea abies and Przmus serotina. Number Importance Success Rank Status Planted Species Value Index I N 500 Pinus strobus 80.99 0.162 2 N 0 Quercus rubra-group 64.24 * 3 N 200 Pinus resinosa 29.33 0.147 4 N 0 Quercus alba 17.36 * 5 E 100 Picea abies 16.42 0.164 6 R 100 Prunus serotina 16.20 0.162 7 N 0 Amelanchier sp. 15.42 * 8 N O Pinus banksiana 10.24 * 9 R 100 Acer rubrum 10.22 0.102 10 E 100 l’inus sylvestris 9.96 0. 100 I l N 0 Quercus ellipsoidalis 4.74 * 12 N 0 Populus tremu/oides 4.62 * 13 N 0 Przmus virginiana 3.61 * 14 R 1000 Rohinia pseudoaeacia 3.49 0.003 15 R 200 Fraxinus sp. 3.09 0.015 16 R 0 Prunus sp. 2.80 0.000 17 R 100 F raxinus americana 2.78 0.028 18 R 0 Prumrs pensylvaniea 1.98 * 19 R seeds Pinus rigida 1.40 * 20 R 100 .lzmiperus virginiana 1.13 0.01 l *These species were not planted or, in the case of Pinus rigida, planted by seed. 67 Table 2-5. Rank in dominance by Importance Value of woody species and their Success Index in the old agricultural field south of the plantation at the Grayling Agricultural Experiment Station, Crawford County, MI. Status indicates whether the species is considered to be Native (N) to these pine barrens, native to the Region (R) but not the forest type. or Exotic (E) to North America. Number planted in plantation based on Beal’s notes from 1888 (Anonymous 1913). Although the recent Quercus mast year (personal observation) gives much weight to the group’s importance value. the exotic Pinus sylvestris is as prevalent as its native analog P. banksiana. Number Importance Success Rank Status Planted Species Value Index 1 N 0 Quercus rubra-group 104.00 * 2 N 0 Pinus hanksiana 49.03 * 3 E 100 Pinus sylvestris 45.1 1 0.451 4 N 500 Pinus strobus 24.57 0.049 5 N 200 Pinus resinosa 15.66 0.078 6 R 1000 Rohim'a pseudoacaeia 13.73 0.014 7 R 0 Quereus alba l 1.28 * 8 E 400 Popu/us alba 9.30 0.023 9 E 100 Picea abies 8.83 0.088 10 R 100 Prunus serotina 6.77 0.068 I 1 N 0 Quercus ellipsoidalis 5.17 * 12 N 0 Amelanchier sp. 4.54 * 13 R 100 Acer rubrum 2.01 0.020 *These species were not planted. DISCUSSION Within the original plantation, the species native to these pine barrens had greater survival rates, radial growth, height. and regeneration rates than all other species. Of 41 hardwood and conifer species originally planted, stems from only seven conifer species survived, with Pinus resinosa, Picea abies, and P. strobus showing the highest survival rates (Appendix I). Site index was 44-45 for the native Pinus resinosa and P. strobus, 29 for the regionally native Picea glauca, and 28 for the nonnative P. abies. Relative to the number of individuals originally planted, P. abies, Pinus strobus, and Prunus serotina showed the greatest overall success based on total number of stems (including 68 regeneration), ground coverage, and frequency. However, P. strobus had the highest relative density of both saplings and large trees in the plantation. The old field community adjacent to and south of the plantation was composed of a different set of species than the plantation community. Pinus .sylvestris and the unplanted natives P. banksiana and Quercus ellipsoidalis dominated the overstory, while P. strobus, P. banksiana. and P. .sylvestris composed most of the advanced regeneration. However, the seedling stratum was composed almost entirely of Q. rubra-group. Relative to the number of individuals originally planted in the plantation. P. sylvestris was the dominant in the old field based on total number of stems (including regeneration), ground coverage, and frequency. Survivorship and Growth Survival, establishment, and growth of the planted seedlings, cuttings, and seeds depended upon a number of factors at the GAES. Most of the largest site preparation treatment (harrowing) was plowed under the year after planting, resulting in the loss of an unknown number of individuals (Anonymous 1913). This area may have been replanted, but no records exist to this effect. Assuming “the same lot” (Beal 1888b) were truly planted in each treatment. then the removal of these individuals should have had no effect on relative survival but would have an effect on the eventual plantation composition by providing preexisting trees (i.e., Pinus banksicma, Quercus spp.) an opportunity to mature in this area of the plantation. An additional unknown number of 69 seedlings of Pinus sp. (presumably P. strobus) and Picea abies were planted in the south side of the plantation to replace individuals accidentally removed while cultivating (Anonymous 1913), which could have given these species an advantage in numbers. Other human-related factors affecting the plantation include removal of holiday trees by local people and loss of large-diameter Pinus resinosa and P. strobus due to windthrow in 1977 (Montgomery 1977. K. Gardiner. personal communication). Site index (SI) is an indicator of the potential productivity for a particular species on that site (Barnes et al. 1998). Besides the general problems association with older anamorphic SI curves reviewed by C armean et al. (1989), some curves are not long enough to include stands over 100 years old. Extrapolation, such as for Picea abies in this study, is required to derive the S], but errors are magnified in the process. SI values estimated for Picea spp. at GAES were lower than those derived for P. g1auca(Sl 40-47. Alban 1985) on similar soils southwest of Grayling. Higher SI were calculated for Pinus resinosa (SI 62-70, Alban 1985) at the same site southwest of Grayling and on sandy soils (SI 62-69, Alban et al. 1987) in Minnesota and Wisconsin. In stands on well- drained sands and closer in age to that at GAES, SI values for P. resinosa and P. strobus were similar (Shetron 1975). Overall. the GAES site is on the low end for potential productivity for these four Pinaceae. especially for the species not native to the pine barrens. Environmental factors play an obvious role in determining the survival. establishment, and growth of trees at the GAES. Of the 41 planted species, many are not 70 adapted to the droughty. infertile, cold conditions. Some individuals probably never established due to high susceptibility to transplant shock, such as Picea glauca. which grows better in boreal environments with higher relative soil moisture (Burns and Honkala 1990a). Other species have not naturalized because of the short growing season in northern Michigan, such as Catalpa .S'peeiosa, Celtis 0ecidenta/i.s', G/editsia triacanthos, Gymnocladus a’ioica, and Juniperus virginiana (Burns and Honkala 1990b, Voss 1972, 1985, 1996). On the other hand, species such as Salix spp., Fraxinus nigra. F. pennsylvaniea, and Thu/a occidentalis are not drought resistant and thus cannot survive in the excessively drained pine barrens (Barnes and Wagner 1981, Burns and Honkala 1990a, Voss 1996). Some species may not be able to extract nutrients from the low-fertility soils, while others, such as Pinus .sylvestris, have evolved efficient mechanisms for cation uptake in such soils (Burns and Honkala 1990a). Although allelochemical interactions with these tree species are not recognized in the literature. lichens and plants common at the plantation, such as bracken fern, blueberry, and Cladina lichens, repress growth and reproduction of some Pinaeeae (Burns and Honkala 1990a, Dolling 1996,.1aderlund et al. 1997. Dolling 1999). Once the planted individuals established, herbivores and pathogens could have had deleterious effects on growth and possibly survival of both conifers (e. g., blister rusts, insects) and hardwoods (e.g., white- tailed deer). A multitude of factors, probably involving edaphic and climatic controls, prevented the successful establishment of 37 of the 41 species originally planted at the GAES. 71 Growth and eventual reproduction ofestablished individuals planted at the GAES were limited by edaphic and climatic conditions. The excessively drained sands are very strongly acid (pH 3.7-4.7) and contain very little (1%) organic matter in the B Horizon (Mokma and Vance 1989). Organic matter had only 110 years to accumulate since the last series of fires ran through the area prior to planting (Kedzie 1888b). Since organic acids are important for chelating aqueous cations like iron and calcium, cation exchange capacity (2.2-4.2 meq/ 100g) and water holding capacity are very low (Mokma and Vance 1989). In addition, the growing season is short with good chances for frost as late as early June and as early as early September, and most of the precipitation falls outside of the growing season (Werlein 1998). Consequently, growth at the GAES is limited by droughty, infertile sand in a short frostfree growing season. Biotic controls could have limited the growth of the less competitive individuals planted at the GAES. All individuals were originally planted on four-foot centers (Beal 1888b), thus all had less than one meter of aboveground space to branch horizontally. Those species that invest in height growth (e.g., Pinus resinosa or P. strobus) or produce dense shade (e.g., Picea abies) would have the greatest opportunity to sequester growing space (sensu Oliver and Larson 1996). Belowground resource competition could also have limited the growth of some species. Pinus .sylvestris, for example, when planted with other native pines. is a better belowground resource competitor due to its development of a taproot and ability to sequester nutrients at low concentrations (Burns and Honkala 1990a). Conversely, facilitation through direct and indirect interactions rather than direct competition for resources could have promoted the establishment and growth of some species (Callaway and Walker 1997). The conceptual model of Holmgren et al. (1997) predicts that in xeric habitats, such as the pine barrens, moisture limitation is more important than light limitation. Consequently, as the intensity of abiotic stresses, such as soil moisture limitation, increases. competition should decrease and the importance of facilitatory interactions should increase (Callaway and Walker 1997). Reanalyzing the data of Bertness and Callaway (1994), Callaway and Walker (1997) detected an increase in the nurse plant effect, or sheltering of one species by another. as the abiotic stresses in the environment increased. F urthermore, in a sandy habitat, Kellman and Kading (1992) found increased seedling and sapling densities of Pinus resinosa and P. .s'trobus under canopies of mature Quercus ruhra than in open areas. Before the practice was widely used, Beal insightfully planted deciduous species as nurse trees for the conifers at GAES, as he did at other experimental plantations and arboreta (Telewski 1998). While Beal might have expected the nurse trees to provide shade, limit evapotranspirative water loss, and add organic matter to the soil, most of the hardwood seedlings did not survive their first year (Anonymous 1913) and thus did not provide this expected nurse effect. Additionally, fast-growing conifers (e. g., P. resinosa, P. strohus, and Picea abies) could have eliminated shade-intolerant species (e. g., Pinus sylvestris) within the plantation. thus opening up areas for less competitive species (e.g., Picea glauca, Juniperus virginiana, and Thu/"a occidental/is) (.s'em'u C onnell 1990). 73 Demography The composition and structure ofthe plantation and old field are a function of residual species at time of planting. the tree species and number planted in 1888-1889, stand dynamics, and establishment, growth, and reproduction of all tree species present. However, the initial establishment, or mortality, of Beal’s seedlings played the greatest role in determining which species would subsequently mature and reproduce within the plantation. Following seedling establishment, though, moisture limitation can be a selective force in community succession (Livingston 1905, Holmgren et al. 1997), but shade tolerance appears to be more influential in shaping this community (sensu Kobe et al. 1995). The intermediate shade-tolerant Pinus strobus contributes more to the plantation community in terms of mature trees, advanced regeneration, and seedlings, which can tolerate up to 80% shade (Burns and Honkala l990a), than any other species planted here. Although seedlings of the shade-intolerant Quereus rubra-group are now very common. saplings and adults are not common. While most of the Q. ruhra-group seedlings are probably a result of a recent mast year, some may be sprouts from older root stock that had previously produced unsuccessful shoots (Beal 1888a, Burns and Honkala l990b). Although Pinus resinosa, another native shade-intolerant species, has slow-growing seedlings (Burns and Honkala l990a), successful advanced regeneration was present in the plantation. Picea abies, an exotic shade-tolerant conifer, produced abundant seedlings and advanced regeneration within the plantation (Appendix II). 74 Although Pinus banksiana was initially present and P. .sylvestris was planted, these conifers contributed less to the plantation community than the aforementioned conifers. The seeds for both of these species require bare mineral soil for germination and both species are very shade intolerant (Burns and Honkala 1990a). By the time these trees were sexually mature, the previously bare mineral soil likely was covered by deciduous and conifer leaf litter, and the canopy was beginning to close due to the other taller, more shade-tolerant conifers. Consequently, any regeneration by P. sylvestris and P. banksiana within the plantation was excluded due to lack of germination sites. except in areas of unsuccessful P. strobus, P. resinosa, or Picea abies establishment (e.g., harrow treatment). The shaded understory of the plantation provided an opportunity for regeneration of shade-tolerant Pinus strobus and deciduous trees. A few individuals of A melanchier sp., a slow-growing, shade-tolerant shrub native to the pine barrens (Barnes and Wagner 1981), established and now contributes an abundant number of seedlings to the plantation floor. Prunus spp. seedlings are very common in the understory, as is the intermediately shade-tolerant Quercus alba (Barnes and Wagner 1981). These species are characteristic of hardwood forests that have evolved from fire-suppressed pine forests (Whitney 1986). The old agricultural field presents an artifact of the species originally present at the site combined with the invasive nature of two conifer species. Pinus banksiana, the namesake for the jack pine barrens, was predominant in this physiographic landtype, with 75 Quercus rubra-group also common (Whitaker 1986, 1987). Both species are adapted to relatively short crown fire return times (80 yr, Whitney 1986)2 through mechanisms such as cone serotiny in P. banksiana and stump sprouting in Quercus spp. Although no severe fires have occurred since 1888. this area had been used for growing cereal grains and pasture crops (Kedzie 1888b). The fallow land was colonized by those shade- intolerant species best adapted to growing in soil after a land-clearing disturbance: P. banksiana, Q. ruhra-group, and P. .Sj‘lt’c.8'lt‘i.8'. Q. ruhra-group seedlings were the most common (91%) seedling in the old field, but this was probably due to a recent mast year. Under the plantation‘s heterogeneous canopy cover. P. ban/csiana and P. sylvestris were able to survive and regenerate in the gaps and thus producing a noncontiguous seed source, while P. strobus produced a steady supply of seed throughout the plantation edge that could germinate in the old field. The relatively flat size class distribution for P. banksiana and P. resinosa and strongly sloped distribution for P. strohus in the old field suggest that suitable sites for regeneration of the shade intolerant pines are limiting and that the shade tolerant P. strobus will increase in abundance in the old field. Oliver and Larson’s (1996) stand development model for single-cohort mixed- species stands best explains some of the dynamics observed in both the plantation and old agricultural field. In this four-step model, a relatively even-aged cohort of trees is established (i.e., planted) following a large disturbance. Following establishment. growing space decreases such that competition for space resources increases. Those species with a competitive size advantage (e.g., Pinus strobus) or growth pattern (e.g.. 2 Less severe ground fires in presettlement had a shorter return time (20-40 yr. Heinselman 1981; 25 yr. Whitney 1986) in thejack pine forests ofthe Great Lakes region. 76 Picea abies) can usurp growing space and reduce the growth rate sufficiently to stunt or kill less competitive species (e. g., Picea glauea). In addition, other species not present at the stand initiation stage are also excluded from successfully establishing in the stand. As the overstory grows and begins producing its own regeneration, new shade- tolerant species (e. g.. Amelanchier spp. and Aeer rubrum) are able to colonize and establish in the understory. Only those species adapted to low light levels are able to survive in the understory of the densely planted stand, until gaps form in the overstory due to localized disturbances, such as windthrow or damage, insect defoliation and mortality, and senescence. Gap formation sets back the stages of stand development and may allow less shade-tolerant but faster growing species (e.g., P. .sylvestris) to attain canopy (Kobe et a1. 1995). The old growth stage is reached once the overstory trees begin dying in an irregular spatial pattern. Although neither area appears to have reached the old growth stage, a diverse understory of new shade-tolerant species and overstory regeneration has developed in the plantation. The old field, however, contains an overstory and understory of predominately shade-intolerant species (e.g., P. banksiana, P. .sylvestris, Q. rubra-group). suggesting that the old field is still in the stem exclusion stage. Although the shade-tolerant Pinus .s'trobus is dispersing from the plantation into the old field, this species is also quite light-tolerant and often acts as both a pioneer and long-lived successional species (Bums and Honkala 1990a). In essence, the dynamics of this planted forest community. as a function of survival, growth, and reproduction ofdifferent species in an initially uniform environment, are largely regulated by interspecific physiological differences. 77 Traditionally shade-tolerant species maximize survival and net growth by maintaining a slow rate of growth, lower rates of whole-plant respiration, lower tissue turnover, and increased whole-plant carbohydrate storage (Walters and Reich 1999). As a consequence, these species are able to survive for long periods of time in the shaded understory to eventually attain canopy status. Conversely, shade-intolerant species possess a high rate of growth at low and high light levels but have short leaf lifespans, high respiration rates. and high nutrient turnover rates (Gower et a1. 1993, Walters and Reich 1999), especially since nutrients can be difficult to sequester in this low fertility soil. Similarly, regeneration is affected by this interaction of light and moisture availability. These interspecific morphological and functional trade-offs provide the interplay for community succession. Invasive Potential o/‘Nonnative Species Normative species that perform as well or better than the locally adapted native species could become invasive in a particular ecosystem (Sax and Brown 2000). Based on the measures used in this study, two exotic Pinaeeae may be classified as invasive species in this pine barrens ecosystem. Although rigorous criteria to designate species as invasive are not available, the accepted criterion for an invasive Pinus is that the species regenerates naturally and recruits seedlings in natural or semi-natural vegetation at least 100 m from the parent plants (Richardson et al. 1994). Beyond the 40 m width of the old agricultural field at GAES, the area surrounding the plantation has been impacted by industrial development and unknown land use practices, thus the 100 m criterion cannot 78 be used here. However, the potential for spread of nonnative species can be assessed within the limits of this study. Within the forest cover of the plantation, the exotic Picea ahies may be considered an invasive species commensurate in performance with the native Pinus strobus. While planted Picea ahies did not exhibit growth as great as Pinus strobus, the exotic species occupied an equitable amount of space relative to the native pine (Table 2- 4). Regeneration within the plantation was not particularly high (Figure 2-4), but Picea abies effectively blocks light needed by other species for regeneration and can regenerate asexually by layering (Burns and Honkala 1990a) as observed at GAES. Picea abies was the second most successful planted species in the old agricultural field (Table 2-5), indicating that it has the ability to spread into unoccupied habitat. Picea abies could compete effectively with the native Pinaceae, especially Pinus strobus and P. resinosa, in this ecosystem. The exotic Pinus .sylvestris was highly successful in spreading from the main plantation to the old agricultural field. This species was over five times as successful compared to any other planted species in the old field; only Pinus banksiana and Quercus rubra-group, preexisting non-planted species, had greater importance values (Table 2-5). Within the plantation, Pinus .sylvestris was not common under closed canopy conditions but was common in areas with canopy gaps (personal observation). Thirty years after planting, P. sylvestris was considered to have surpassed all of the other species in growth and was the only species with regeneration by seed (Roth 1919). P. .sylvestris is able to 79 germinate in low nutrient conditions, acquire nutrients at low concentrations, grow in high light conditions, and produce copious seed at an early age (Burns and Honkala 1990a), making it a potentially successful invader of jack pine barrens. The interest in potentially invasive pines increases as the structure and functioning of native ecosystems, especially shrublands and grasslands, are altered (Richardson and Higgins 1998). Although no phylogenetic relationship between invasive I pines is known (Price 1998), the strongest correlates to invasiveness include small seed mass, short juvenile period, and a short interval between large seed crops (Rejmanek and Richardson 1996). However, the interaction of these life history traits and the environment in which invasive species colonize determines the eventual community composition. If growth to the seedling stage as a function of the environment were guaranteed, then survivorship and regeneration should be a better measure of the potential for a species to colonize a site. There is a need for predictive models for which pine species are invasive in different locations (Richardson and Higgins 1998). Given the results of this study, potentially invasive species should not be used for reforestation, and existing populations should be controlled to prevent their spread. Future Composition ofthe Plantation and Old Field Barring major disturbances. such as fire, logging, and windthrow, the plantation and old field at the GAES are expected to evolve similar to other fire-suppressed forests in the pine barren system (Radeloff et al. 1999). As the less shade-tolerant species are 80 excluded by the more shade-tolerant species, Pinus strobus and Picea ahies, which are already dominant species in the overstory and understory, are likely to become more important in the plantation, but. through gap dynamics and senescence, a hardwood- dominated community characterized by Quercus spp. and Acer rubrum may develop (Curtis 1971, Radeloff et al. 1999). Although delayed, the old agricultural field will likely undergo a similar succession. especially since overstory disturbance is absent there as well. Since the GAES is enclosed by development on three sides and is now annexed to the Hartwick Pines State Park, return to a natural disturbance regime that would maintain the species typical ofthis pine barren ecosystem is not expected. 81 .wcoEN .4 mo $2.58 aaE 3mm .Gae 5:80 .82 EEoUV 38% 2a 2553 «En Exam .3 BEEEoon EwwEEZ E 8:550 dob—c8 56:2 .303 25:58.: 552805 :8 some E53» $52 $8.32: E wow—Ea 26>» me§8 28 meEuom dawEEE .5580 EQBEU E coufim EuECoaxm EEGER< wEREO L258 05% :S% was cocmooq .TN EsmE 20¢ 20 _ a... _ 265m: .9550 AllstNl¢ :2 .358 22390 82 GRAYLING (347 m) 11.8 °c mm C 1 (44°39’N, 84°42’W) 833 mm ”00 [1961-1990] 4/ // fl -- 80 30.0 '. / 20.0. N / l/‘NL E40 10.0 . JV ‘10'0‘J FMAMJ JASOND Figure 2-2. Climate diagram (sensu Walter and Leith I967) based on 1961- l 990 data from the Grayling, Michigan, Station (COOP ID 20339l), National Climatic Data Center. Mean annual temperature is l 1.8C, and mean annual precipitation is 833 mm. The top line refers to average monthly precipitation (mm), and the bottom line refers to average monthly temperatures (°C), The solid bars along the horizontal axis refer to months with freezing conditions, and hatched bars refer to months with chance of frost (Weirlein I998). 83 10 1 Acer rubrum 2 1 3 3 . 0 d) .2 h l a 6 E on o 1‘4 4 I— 2 —o—Plantation g 2 1 —|:|—Old Field '5 z o _g_ m m. c» a? m. 0? m. c» m. m m V 0') V O) V O) V O5 V 05 g o 11') ‘7 '7 ‘1‘ ‘7‘ “2 ‘1’ ‘1 Y . O In O LO 0 L!) O to O u- ‘- N N ('0 (‘0 V V L0 Diameter size class (cm) 10 . Fraxmus sp. 9 i 3 8 o a: a: I- 3 6‘ m i "' 4 U- . 3 —o—Plantation 3 —u—Old Field E 2. a Z 0 + m. m. m. m. a! m. m. m m c». m V O) V 0') V 0') V CD V 05 V o u') ‘T ‘7 “3 9.‘ ”9 ‘1’ 3’ ‘7 ‘9 O L!) O In 0 LO 0 In C) ._ v— N N (‘0 (‘0 V V In Diameter size class (cm) Figure 2-3. Number of stems (> I .37 m tall, per hectare) for each shrub and tree species surveyed in the plantation (solid line) and old field (dashed line) at the Grayling Agricultural Experiment Station. Crawford County, MI. Numbers were converted to per hectare to standardize for the different area sampled in the plantation and old field. The values on the _}"-21XlS differ by species. 84 10 Juniperus virginiana —D—Old Field n .w t a t n B P 9.53: .2. much .0 39:32 m.vm-om $9.1m v m.mN-mN méméw 0.9-2. méTo— Diameter size class (cm) 100 Picea abies —n—Old Field —o—P|antation 0 0 0 8 6 4 Mu ! 2302. .2. new: *0 .3532 0 mdmém m.mN-mN m.vm-om mmeF l @670? , m.m-m mélo Diameter size class (cm) Figure 2-3 (cont‘d). 85 mdmém demV méYov —C]—Old Field —-o—- Plantation m.mm-mm m.mm-m~ Pinus banksiana H 3&8 \ $72 . m3? _ m3 m.v.o 20. O 0 0 0 0 0 8 6 4. 1 120 2800: .2. mach to 59:52 Diameter size class (cm) 100 ‘ m.vm-om m.mv.mv mévdv _.D_Old Field —0— Plantation m.mm-mm mémém m.mm-m~ m.— mémém Pinus resinosa QmeF m.v7ow 0.06 1 Am. .mso 0 2 2300; .3 39: .0 32:52 Diameter size class (cm) Figure 2-3 (cont’d). 86 Pinus rigida 2 S 8 u l o .C a s. m o 2 C 4; 3 1 —o—Plantation 3 2 —C|-—Old Field g i Z 0 l .43. 0’. m m. m. 02 c». m. m m c». m V O) V O) V 0') V O? V O) V o «3 '7 ‘7 “.I 0.1 0? «3 at s “3 O In 0 In 0 In 0 In 0 ‘— V— N N m 0") V V “7 Diameter size class (cm) 2000 _ 300 . Pinus strobus . 250 e 1600 g g u 2 ; 200 g ‘- 1'“- 1200 h A a 5 3 g 5 - 3 3 . 150 3 IL 35’ 5 :3. 2 "6 9, 800 3 —o—P|antation E 9' L g \ —c]—Old Field 10° 3 s s 400 2 l 50 Z 0 f 0 on a) a) a) a) 05 or 07 m m a) V O) V CD V CD V O) V 0" V o u') *7 ‘7 “.1 0.1 "2 °? at ‘i “P O In 0 In C In 0 In 0 v— 1— N N (V) 0') V V In Diameter size class (cm) Figure 2-3 (cont’d). Note the second y-axis for Pinus SII‘OINJS in the old field. 87 120 Number of trees per hectare 8 01 i; q; 0) O In 20; l O giefii at 2 \ gm. in '\ 0 s8\ “5 '\ O #24 \ 3 2 Figure 2-3 (cont’d). 10-149 15-19.9 ‘ Diameter size class (cm) 15-19.9 Pinus sylvestris 0’. v ‘7‘ o N 25-299 J O) V. ('0 O (‘0 Populus alba 20-24.9 Diameter size class (cm) 88 25-299 -34.9 30 -39.9 35 —o-— Plantation —-c]—Old Field m. c». 02 =1 2; at» O In 0 V V In —-9— Plantation -Cl— Old Field 4044.9 4549.9 .9 50-54 Populus tremuloides 201 —C]—O|d Field —0— Plantation 6 2 8 4 1 230m: .3 much “.0 33:52 mfimém mévév m.mm-mm mémém m.mN-mN m.vm-o~ www-m— Diameter size class (cm) _.|j—Old Field n .m t a t n m P mémém m.mm-mm Prunus serotina www-mw @670? 200 160 . 2 9.302.. can new: be 355:2 Diameter size class (cm) Figure 2-3 (cont’d). 89 Quercus alba —C|— Old Field —o-— Plantation 233: 3a 33.: “—0 39.52 mdvfiv m.mm-mm m.vm-om m.mN-mN *méméw @me— 0.3-09 m.m-m m.v..o Diameter size class (cm) Quercus ellipsoidalis 20 l I —{:|—Old Field —0— Plantation 6 2 8 4 at 9.53... 3a new: *0 33:52 mémém m.m~-mm mémém Diameter size class (cm) Figure 2-3 (cont’d). 90 Quercus rubra - group 200 1 runs: ill- mémém 0.91m». —-E]—Old Field —0— Plantation m.mm-mm mémém m.mm-m~ m.m-m méuo nw / ,, 0e 0 O 8 160 2 253: 3a 335 *0 39:52 Diameter size class (cm) 15--.— —C]— Old Field —0— Plantation Robinia pseudoacacia 0 40 1 20 3333: 3a 335 20 39.52 0 m.vm-om QmYmv m.v¢.ov Qmmbm mémém m.mN-mN m.vw-om m.mw-mw méTOF m.m-m m.v.o Diameter size class (cm) Figure 2-3 (cont’d). 91 www— E .538an 2: E .3525 bEEwto was 330% 2: :2: .3823: 81:83 2; .=2 £5on 28380 522333 2: («o :58 Eva 3523:? 20 can 5:851 :ozfim 5353me _EB_:otw< wEEEO EEwto 2: E 330% E 2% Eu omAv 83.: owes ES 2% Eu o~18 meEMm mo 35% 3:23— .v1m eswi no 0 O O F W. m m mg m u I. I. m 1. d n d d d S S S O U S n w. n n I: V J 9 n n l, O S U n S Ila S J O 9 n I 9 S J 10 S n U S 3. A 9 9 n q H J S a m A S n J W 6 fl. VIA. I. W m w m a w n N B s w u 2 Rio 0 m e Aw o S w m S a w u. w. y: e u. m a. o J m.. e U. m. me: me, 9 Flu. o B. m. a. J w m W W w W 0. w. n W S W a. u w w W... :0 m. 9 a S 9s u S: e e; 9 Se 9... .u a ,1 Ilflfldlifl .1 . 1 1 1 I I Io — or r \ om w . A o 0/ Nu f on B r A a m ov u B. \IO- .A ,. om m8: 8.3 - 2m: 20- , 853mm - 22“. 200 . 00 $9... 53.. - 52335 I wmgamm - 3:935 I s1 on 92 33— .33 .NS 3 mmo> 80¢ 03E 3:98 2.29 gum: rod was; 36on we cEEEEBSQ dz 5580 28380 528% ~555me HEB—3:9». wEREO SEEM 2: 8 Eu: @332th 20 9a song—BE or: E 86on $003 $25 9: moam> 8:959:— uo 2225320 .m-~ Ezwi «2522 £5.59 5.02 2 0398 $6QO :069 m5 9 025: $6me 9.5 m5 9 w>=mc 86me \ . Q\1 ‘ \ I ; w 1 \ . o V % om w. m. w. ov w 0 u E U m w 00 A al. N 7 a , 8 22“. EC- 7 coszmED , 93 CHAPTER 3 COMPARISON OF CLIMATE-GROWTH RELATIONSHIPS FOR PINACEAE IN THE PINE BARRENS OF NORTHERN MICHIGAN ABSTRACT Mean chronologies were constructed for living Picea abies (L.) Karst., Pinus resinosa Ait., P. strohus L., and P. .sylvestris L. at an historical mixed-species plantation and from an existing raw ring-width data set for old-growth P. resinosa in the pine barrens of northern lower Michigan. All ring-width series were standardized by fitting a negative exponential or negative linear regression, and mean chronologies were constructed as biweight robust means of the autoregressively modeled indices. Chronologies ranged from 85 to 202 years in length. Correlation function analysis of the ARSTAN chronologies to lS-month precipitation and temperature data showed all species exhibiting positive responses to April temperatures. Responses to variations in precipitation were generally negative in the early growing season but varied strongly by species and location. The native pines were more complacent to variations in precipitation but exhibited similar responses to variations in temperature as compared to the responses of the nonnative P. .sy/ves'tris'. The nonnative Picea abies responded more to variations in temperature from the previous growing season and in a different pattern to precipitation than all other species. Continuing work with Pinus .sylvestris and the 94 native pines may elucidate physiological mechanisms that explain patterns of successful colonization of this nonnative species in the pine barrens. OBJECTIVES The objective of this chapter is to determine the climatic limiting factors to secondary growth in surviving trees at the Grayling Beal Plantation in order to elucidate causes for differences in relative survival and growth in this pine barren ecosystem. The raw and standardized tree-ring data from this chapter will be contributed to the lntemational Tree-Ring Data Bank for use by investigators interested in dendroecological relationships in the Great Lakes region. INTRODUCTION Climate is an important factor defining the geographic range for a plant species (reviewed by Woodward 1987). While many species can survive within the normal range of climatic variation in a region, extreme events (Cook et al. 1987), such as early and late season frosts, extreme winter temperatures, and droughts, affect growth and success in regeneration and therefore restrict the long-term species pool (Woodward 1987). Dendrochronology is a useful procedure for identifying the climatic limiting factors to growth for a tree species (Fritts 1966, 1976, La Marche 1978, Fritts and Swetnam 1989) and in turn provides a measure for assessing the suitability of that species to its particular environment. I prOpose the use of dendrochronological techniques to assess the suitability of native and nonnative Pinaceac to growth in the pine barrens of northern Michigan. The pine barrens of the Midwest were formed upon the thick deposits of glacial outwash, largely composed of coarse soils. laid down about 12000 years ago during the Pleistocene glaciation (Whitney 1986, Weirlein 1998). This region is now covered by at least second-growth upland pine forests and barrens growing on sandy soils interspersed 1.“- s... 'hl" -- n with bogs and poor conifer swamps bordering former meltwater channels. The pine forests are dominated by Pinus strobus and P. resinosa, while the barrens are dominated ’ .‘T' by P. banksiana with scattered P. resinosa (Curtis 1971, Whitney 1986, 1987, Comer 1.; 1996). In addition to these native pines, the normative P. .sylvestris has been planted in the pine barrens for timber and Christmas tree production (Wright et al. 1976, Burns and Honkala 1990a), while the nonnative Picea abies was historically planted in this region but is no longer planted in the pine barrens. The sandy soils of the barrens contain low nutrients, organic matter, and water holding capacity (Mokma and Vance 1989). Due to the droughty, infertile soil and cold continental climate in the region (Whitney 1986), trees growing in the pine barrens may be particularly sensitive to variations in climate and may express climatic limitations to their geographic spread. The only dendroclimatic study on the pine barrens of the Midwest reported that P. resinosa was sensitive to April and June-July temperatures and June precipitation (Koop 1985). The purpose of this study is to develop correlation functions from the climatic signal in the high-frequency variation of radial growth in native and normative Pinaceac 96 in pine barrens. In general, I predicted that radial growth for all species would be limited by cool early and late growing season temperatures and dry summers. However, the native species should be more adapted to their local environment and thus would be more complacent to variations in climate as compared to the nonnative species, assuming their native range has a more favorable climate than here. Two stands of Pinaceae located in the pine barrens of northern lower Michigan were studied to address these predictions. METHODS Study Site This study took place on the Grayling Outwash Plain (Albert 1994), Crawford County, Michigan (44°N, 84°W; Figure 3-l). Two study sites were selected based on the availability of tree species and existing tree-ring data. The Grayling Beal Plantation (GBP) site is located southeast of the town of Grayling and contains a S-acre (2.0 ha) plantation of native and nonnative tree species. Seedlings. cuttings, and seed (Pinus rigida Mill. only) from 41 species were planted in 4-foot-centered rows in 1888-1889. Prior to planting, the site had experienced frequent fires leaving mostly P. banksiana and scattered P. resinosa, scrubby Quercus spp., and dwarf Prunus spp. (Kedzie 1888), but no record or evidence of fire exists at this site since planting. The mapped soil is a coarse, loose, strongly acidic, and excessively drained sand of the Grayling series (Typic, Frigid Udipsamment) (Mokma and Vance I989. Werlein 1998). Although the depth to the water table was not measured in this study. Weirlein (1998) notes the water table to 97 be > 6 ft (1.83 m) in this soil type. while a soil scientist at the Natural Resources Conservation Service stated the water table to be 2-3 m from the surface (M. Kroell. personal communication). Government Land Office Survey records show that pre- European settlement forests on this soil type were largely composed of Pinus banks'iana (Whitney 1986). The Hartwick Pines State Park (HRP) site is located just north of Grayling and contains a 40-acre (16.2 ha) remnant stand of old-growth P. resinosa and P. strobus (Koop 1985). This stand was bypassed during Michigan’s logging era of the late 19th century due to the depressed logging markets and the relatively small size of the stand (Koop 1985). Consequently, trees several hundred years old remain on this site. The stand is located along a ridge with 10-15 m relief (Koop 1985) and on soils mapped as a strongly acidic, excessively well-drained sand to loamy sand of the Kalkaska—Blue Lake Association (Typic F rigid Lamellic Haplorthod) (Koop 1985, Werlein 1998). Although Koop (1985) did not measure the depth to the water table, Weirlein (1998) notes the water table to be > 6 ft (1.83 m) in this soil type. Pre-European settlement forests on this soil type and topography were dominated by P. resinosa (Whitney 1986). The local climate is characterized as humid continental (Whitney 1986). The mean annual precipitation (1951-1980) is 811 mm with 62% of the precipitation falling during April-September (Figure 3-2). The mean annual temperature is 6.3° C with a mean summer temperature of 186° C and a frostfree growing season of 1 10 days 98 (Weirlein 1998). The growing season is generally cool and short with only 2068 growing degree days (Weirlein 1998). Sample Collection and Data Analysis Large diameter Picea abies. Pinus resinosa, P. slrobus, and P. .sylvestris at the GBP site were cored perpendicular to the stem lean to avoid reaction wood with 5.15 mm diameter increment borers (Haglof and Suunto) in 1998-2000. One full diameter core or two cores were obtained at approximately 0.3 m from the ground to obtain as many growth-rings as possible. Increment cores were prepared, cross-dated, and measured to 0.01 m using standard dendrochronological techniques (Stokes and Smiley 1967). Ring-width measurements within each time series were computer analyzed to detect anomalies (COFECHA Version 6.06P, Holmes 2000). Tree-ring data for P. resinosa from the HRP site were obtained from the lntemational Tree-Ring Data Bank (ITRDB, Koop and Grissino-Mayer 1988). Methods used to obtain cores, measure ring-widths, and check for quality control followed Stokes and Smiley (1967) and are fully described in Koop (1985). Mean raw ring-width was compared across species using the conservative Tukey HSD Multiple Comparison procedure (Systat Version 9.01, SPSS Science Inc. 1998). All series of raw data were converted to mean chronologies for climate response analysis using the same program settings to ensure consistent comparisons among species and sites. Each series was detrended with a single pass high-frequency filter, i.e., a 99 negative exponential or negative linear regression was fit to the series (ARSTAN Version 6.04P, Holmes 2000). This conservative technique removes exogenous variation and the age-related growth trend but retains the climatically induced high-frequency variation (Fritts 1976, Cook et al. 1990a). Three types of mean chronologies can be constructed from the detrended series: STANDARD, RESID, and ARSTAN (ARSTAN Version 6.04P, Holmes 2000). The STANDARD chronology is the biweight robust mean of the detrended series. While the use of a biweight robust mean enhances the common signal and minimizes the influence ofoutlier index values, which could be a result of endogenous disturbance events that are random in space and time (Cook et al. 1990b), the STANDARD chronology retains some autocorrelation in successive index values (i.e., autoregression) (Cook and Holmes 1997). The RESID chronology contains the biweight robust mean of the normalized residuals (>3 standard deviations) from univariate autoregressive modeling and results in a strong common signal without persistence (Cook and Holmes 1997). The ARSTAN chronology combines the pooled autoregression model from multivariate regression modeling with the residual chronology to produce a chronology expressing the greatest sensitivity to climatic variations (Cook and Holmes 1997). Since the objective of this study was to determine climatic responses, the ARSTAN chronology was developed (Appendix IV). Similar temporal response patterns among the ARSTAN chronologies constructed for each species were compared by simple correlation analysis (Pearson correlation. Bonferroni probability, Systat Version 9.01, SPSS Science Inc. 1998). 100 The relative importance of monthly precipitation and monthly mean temperature in influencing tree-ring growth for each species and site was determined by correlation and response function analysis (RESPO Version 6.06P, Holmes 2000). Correlation functions are the correlation coefficients between the predictors and predictands, while the response function is a result of a multivariate regression analysis on a set of principal components (PC s) calculated from climatic variables (Fritts 1976). PCs enter the stepwise regression if the cumulative multiple of eigenvalue is greater than one, thus the number of PCs retained varies by chronology (RESPO Version 6.06P, Holmes 2000). Although response function analysis has been shown to be highly effective at showing relationships between climate and growth (Fritts and Wu 1986), the response function can be more difficult to replicate and interpret (Blasing et al. 1984). However, providing the statistical constraints used in the analysis and making the tree-ring data available (e.g., ITRDB) allows other researchers to make comparative studies. In this study, the predictand was each ARSTAN chronology, and the predictors were15 sequential months (previous July through current September) of total precipitation and mean temperature variables for the period 1931-1988 for HRP and for the period 1931-1998 for GBP (National Climatic Data Center 2001, Appendix V). RESULTS Site master chronologies (Figure 3-3) were constructed for four species based on the standardized ring-width data summarized in Table 3-1. The GBP site yielded four master chronologies ranging from 85-105 years in length, and the HRP site yielded a single 202-year master chronology. Based on a signal strength threshold of 0.85 (Briffa 101 and Jones 1990), Pinus .sylvestris was the only species that would have required additional samples to fully replicate the chronologies from a subsample. The mean ring- widths for the native pines at GBP were larger than for the normative Pinaceae, while the old-growth P. resinosa at HRP had a significantly smaller mean ring-width. The signal- to-noise ratio (SNR), a measure of variance common within detrended series (Graumlich 1993), varied by species but was more a reflection of sample size (Briffa and Jones 1990). The mean sensitivity was highest ”for Picea abies followed by Pinus strobus, while P. resinosa and P. .s'ylvestris expressed less but similar amount of high-frequency variance. Graumlich (1993) found similar mean sensitivity in P. strobus and slightly higher mean sensitivity in P. resinosa. The amount of low-frequency variance as expressed by the standard deviation of the chronology was highest for P. resinosa at GBP. This chronology also retained the largest first-order autocorrelation coefficient, suggesting that some low-frequency variance remains in the standardized chronology. Since these conifers retain needles, a higher first-order autocorrelation is expected (Graumlich 1993). However, the chronology for Picea abies retained little low- frequency variance even at the first-order for autocorrelation. Although the correlations between ARSTAN chronologies were generally more significant within the genus Pinus than to Picea abies, the Pinus .s'trobus chronology was significantly (p<0.001) correlated to the Picea abies chronology (Table 3-2). The response functions of mean monthly temperature and monthly precipitation explained 28-52% of the variation in the ARSTAN chronologies examined in this study (Figures 3-4 through 8). A greater number of significant responses to variations in 102 temperature and precipitation were indicated by the correlation functions than by response functions, consistent with Fritts (1976). However, caution must be given regarding the interpretation of the direction of correlation coefficients or weights, in the case of response functions, since either a wider or narrower ring could be associated with an abnormally high or low climate parameter. In addition, the more conservative and interpretable correlation functions. rather than response functions, are considered in this study (Blasing er al. 1984). Table 3-1. Dendrochronological characteristics for the raw ring-width data and ARSTAN (ARS) mean chronology for each of the Pinaceae at Grayling Beal Plantation (GBP) and Hartwick Pines State Park (HRP). Mean ring-width values with different letters across species are significantly different (Tukey HSD, p<0.01). Signal-to-noise ratio (SNR) based on detrended series. Minimum sample size based on a signal strength (SS) threshold of 0.85 (Briffa and Jones 1990). All values calculated by ARSTAN (Holmes 2000) GBP GBP GBP GBP HRP Pic-ca Pinus Pinus Pinus Pinus Characteristics abies resinosa stmhus .s‘_t=lvestri.3' resinosa No. trees 1 | 22 10 10 28 No. cores 25 82 21 20 52 Total years 105 101 105 85 202 Mean ring-width (mm) 1.788b 2.177a 2.167ab 1.896ab 1.136c SNR (detrended) 3.697 9.818 5.949 1.131 8.452 Mean index (ARS) 1.0063 1.0446 0.9863 0.9603 0.9845 Mean sensitivity (ARS) 0.2569 0.1330 0.1810 0.1349 0.1462 Std. deviation (ARS) 0.251 1 0.3646 0.2232 0.2389 0.2045 Skewness (ARS) -0.3041 2.1610 -0.2052 0.4611 0.1966 Kurtosis (ARS) 0.7971 6.8889 0.2696 0.991 1 0.1009 Partial Autocorrelation (ARS) lSt order 0.1318 0.8021 0.5049 0.7399 0.6307 2"d order -0.0623 0.0304 -0012 1 0.1707 003% 3rd order -01 197 -0.0765 0.1053 0.0443 0.0142 Min sample size (no. trees) 10 13 6 38 10 103 Table 3-2. Pearson correlation coefficients (r) and significance (Bonferroni probability) between the ARSTAN master chronologies developed for each species at Grayling Beal Plantation (GBP) and Hartwick Pines State Park (HRP). All values calculated by Systat Version 9.01 (SPSS Science Inc. 1998). GBP G BP GBP GBP HRP Picea Pinus Pinus Pinus Pinus abies resinosa strohus .s:1’l1‘ustri.s' resinosa GBP - Picea abies 1.000 GBP - Pinus resinosa 0.234 1.000 GBP - Pinus slrobus 0.408‘" 0594““ 1.000 GBP - Pinus sylvestris 0.313 0583““ 0.429‘" 1.000 HRP - Pinus resinosa 0.162 0447"“ 0.335' 0458”” 1.000 77<0.05. "p<0.01. "'p«<0.00 1. ""Ir 0.0001 Significant responses by the nonnative Picea abies were limited to a positive correlation (r=0.28) to December temperatures of the previous growing season (Figure 3- 4). The other nonnnative, Pinus .sylveslris, responded positively to April temperatures (r=0.31) and September precipitation (r=0.25) and negatively to April (r=-0.30) and previous January precipitation (r=-0.34) (Figure 3-7). Similarly, the native P. strobus (Figure 3-6) and P. resinosa (Figure 3-5) at GBP exhibited positive associations with April temperatures (r=0.26, 0.24, respectively) but expressed no other significant response to variations in temperature or precipitation. P. resinosa at HRP responded negatively to July temperatures (r=-0.30) and precipitation in August (r=-0.29) and the previous September (r=-0.32) (Figure 3-8). Since the HRP chronology ended at 1988, the P. resinosa chronology at GBP was reanalyzed for the common period (1931-1987); I observed no difference in significant correlations from the full period (1931-1998). A post-hoe correlation analysis of March-April mean monthly snowfall and maximum snow depth (National Climatic Data Center 2001) to the STANDARD chronologies yielded no significant (p<0.05) correlations. 104 DISCUSSION This study examined the high frequency radial growth response of four species of Pinaceae to monthly variations in mean temperature and total precipitation on the pine barrens of northern lower Michigan. All of the species demonstrated sensitivity to climatic variations but to somewhat different signals and extents. The response functions from monthly temperature and precipitation explained 28-52% of the total variation in annual radial growth. The most common growth response was to abnormally high April temperatures, a correlation also observed by Koop (1985) and Graumlich (1993), who worked with Pinus resinosa and P. strobus in the upper Great Lakes region, suggesting that warmer early growing season temperatures may increase radial growth. No species showed significant correlations to variations in late growing season temperatures. In general, abnormally high precipitation during the growing season correlated to greater radial growth in all chronologies but P. resinosa at HRP. The Pinaceae native to less xeric environments appear to be less adapted to short- terrn fluctuations in the climate regime of the region, which is not expected to influence the responses as much as the species-to-species variation (Graumlich 1993). Mean sensitivity, an indication of the degree to which radial growth varies from year to year (Fritts 1976), was largest for Picea abies and Pinus strobus (Table 3-1), species that prefer more mesic soil conditions than the other Pinaceae growing at GBP (Whitney 1986, Schweingruber 1993, MacDonald et al. 1998). Allocation to secondary radial growth appears to be more sensitive to available resources, or photosynthates, in the 105 short-term for these two species. Since radial growth is affected by climate (Fritts 1976), greater mean sensitivity gives an indication of the strength of dependence on a limiting factor(s) having high-frequency variance. presumably climate. In addition. the chronologies for Picea abies and Pinus strabus exhibit lower first-order autocorrelation coefficients than the other species (Table 3-1), indicating lower persistence in growth in subsequent years (Graumlich 1993). While the chronologies for Pinus slrobus (Figure 3-6) and P. resinosa (Figure 3- 5) responded similarly to climatic variables, the Picea abies chronology (Figure 3-4) expressed more significantly negative responses to temperature in the previous growing season than any other chronology. Physiologically, an abnormally warm, dry Fall could result in an excess of root and stem respiration over gross photosynthesis, thereby reducing carbohydrate stores available for Spring growth (Fritts 1976). Alternatively, a abnormally cool, wet Fall could reduce water stress and thus allow for continued cambial growth later in the season than during normal years. In either case, radial growth in P. abies is more sensitive to variations in climate, especially in the previous growing season (Makinen et al. 2001), than the native Pinaceae considered here. Although species living at the edge of their range express the greatest sensitivity to limiting factors like climate (Fritts 1976), the geographic range for P. abies includes much colder to more mesic environments (Schweingruber 1993) than Grayling. However, the physical features (e.g., short needles and fastigiate shape) of the P. abies at GBP resemble the more northern races of the species (F. Telewski. personal communication). 106 Despite a low mean sensitivity. the nonnative Pinus sylvestris responded more to variations in precipitation (Figure 3-7) than any other species considered here. Increased radial growth was associated with higher late season precipitation, suggesting that P. sylvestris has a longer phenology of cambial activity than the other Pinaceae thereby increasing production of photosynthates at the end of the growing season in these years more efficiently than the other species. Furthermore, although all species at GBP expressed a negative association of radial growth with January and March/April precipitation, these correlations are significant only for P. .sylvestris. Abnormally high snowfall in January may maintain low soil temperatures through early Spring (Thomsen 2001), thus delaying bud break, leaf expansion, and cambial induction and reducing radial growth for the year. The negative April relationship may be better explained by less influx of solar radiation due to clouded skies than by water availability (F ritts 1976); unfortunately, these climate data are not available. Although the small sample size and low SNR for this species necessitates concern over the interpretation of these results, the broad ecological distribution (Burns and Honkala 1990a) of P. .s'ylvestris and its potential ability to take advantage of local climate conditions supports the notion that P. .sylveslris is a successful pioneer species (Gutierrez 1989). The ARSTAN master chronologies for Pinus resinosa at GBP and HRP were significantly (p<0.0001) correlated (Table 3-2), but their growth responses to variations in monthly precipitation and mean temperature were different. Both chronologies had similar mean sensitivities (Table 3-1) and indicated positive correlations to April temperatures. but the trees at HRP expressed negative correlations to July temperatures 107 and precipitation in August and the previous September (Figure 3-5.8). Using a different version of the HRP chronology (RESID versus ARSTAN) with only the current year‘s climatic parameters, Koop (1985) also found significant correlations to June-July (p<0.005) and April (p<0.05) temperature but found a slightly positive correlation to June precipitation (p>0.05). The difference in Summer precipitation responses could be most attributable to the use of a different number of climatic parameters in the analysis. Furthermore, while GBP is a relatively even-aged closed canopy forest on level ground, the crowns of the uneven-aged trees at HRP are more exposed to solar radiation and wind, and the roots are presumably further from the soil water since they grow along a ridge (Koop 1985). Consequently, these trees may suffer greater water stress through evaporative water loss and have less access to soil water than those trees at GBP. In addition, the pines at HRP are larger, thus have a greater overall respiratory demand, and older, thus are less physiologically robust (Kozlowski 1971). A differential response to variation in temperature and precipitation would thus be expected across the two sites. In general, the correlation and response function models showed that radial growth appears to be more limited by climate before and at the beginning of the growing season. The models explained 28—52% of the variation observed in the standardized radial growth data. Some of the unexplained variation could be due to the low waterholding capacity of the excessively drained soil. If all rain and melted snow drains through the soil before roots are able to absorb the water, then the trees could be contantly drought stresssed and may show greater sensitivity to fluctuations at a higher amount of precipitation. Since the plantation was planted within a small, homogeneous 108 plot, some of the unexplained variation should be attributed to genetic differences between individuals and within-stand (endogenous) disturbances rather than climate alone. Given the mixed-species planting, the branching architecture and foliage density of neighboring trees could increase the variation in radial growth observed between trees of the same species in the plantation. Wind stress is an endogenous disturbance that removes photosynthate sources and sinks of individual and neighboring trees, dehydrates foliage, and induces differential cambial development (Telewski 1995). Trees at the edge of the plantation suffer greater acute and chronic wind stress. Furthermore, gaps in the canopy created by windfall, such as after the Windstorm of 1977 (Montgomery 1977, K. Gardiner, personal communication), released some trees from competition for light, soil moisture, and nutrients, thus increasing overall variation in photosynthetic activity and cambial growth within a species. However, these trees also endured greater wind sway as a result of the disruption of the canopy boundary layer (Telewski 1995). Subsequent establishment of regeneration in the gaps would eventually moderate any initial release from competition for subcanopy resources. Finally, although evidence was not observed here, acute defoliation (Fritts and Swetnam 1989) and chronic pathogenesis could affect photosynthate production and subsequent radial growth leading to intraspecific variation. Nevertheless, the linear models derived in this study explain a substantial percentage of the high-frequency variation in radial growth for trees in the Midwestern United States (Fritts 1976). 109 mu“ 1 1 . .. . o . . . : 3F. £85m @815 230 mm=< 3:232 5:8:— unCo “5.5.8900 835 335 “858 9:: 03m .Gafl SE00 @3— :unouv @035 2d E05288 530:5 8 Sta mcotan oEa €53. 3 noEEEouoa cmmEEE E 3:550 dmeEE c9550 E8380 E 43m 25m moEm :35: can c2555 Rom wEESO ECO 22:82 2: 9:265 92: EcEmE max“: 320 ._.m 0.5er lIO GRAYLING (347 m) A 113 cc mm 40-0 . (44°39’N,84°42'W) A 833 mm "100 25-9 [1961-1990] /4 \ fl //14rfl \\ ’ — 80 l4 x / \ 0C A W\ 26.9 4 M \ - 60 Al \ s // \NP 9 / 20.0 . \ Le - 40 \N / ywyb H‘NK M y K 2“ “I 10.0 « V \ -20 l \ /V K / \ 45.2 ) . . - V 0 -42.8 ' -1001 Figure 3-2. Climate diagram (sensu Walter and Leith 1967) based on data from the Grayling. Michigan. Station (COOP ID 203391), National Climatic Data Center. The top line refers to average monthly precipitation (mm), the bottom line refers to average monthly temperatures (°C), solid bars refer to months with freezing conditions, and hatched bars refer to months with chance of frost for the period 1961-1990. 111 (JO Picea abies - GBP N on N A Index va1ues A 01 .0 on O 2.5 1.5 Index values 0.5 Pinus resinosa - HRP Index value .. I; ? <> :8 0 O O O O O O O O O O O O O) O t- N (‘0 V In (D N (D O) O w 0) O) O) O) O) O) O) O) O) O) O ‘— v- v- x- ‘— V" ‘- 1— ‘- v- 1- N Figure 3-3. ARSTAN ring-width chronologies constructed for conifers at the Grayling Beal Plantation (GBP) and Hartwick Pines State Park (HRP). 112 2.5 1.5 Pinus strobus - GBP [Vbqw IV) rAWAf\ 11. Index values 0.5 V‘l\l"’ r1111... wwvvwvm 2.5 1.5 Index values Pinus sylvestris - GBP 0.5 1890 Figure 3-3 (cont’d). 1900 1910 1920 1930 1940 1950 1960 1 113 1970 4 1980 . 1990 . 2000 0.4 Temperature *0 c .2 u E 0 O O I Correlation function a Response function -0.4 iaiaiaiaiaiai—i—i—i—i—i—i—i—i- _1 0 Q l— > 0 2 m [I D: >' Z —l (D O. < UJ < o. < 3 D 3 LU 22$8§gfimz- Z —’ (D O. D ‘3 fi 5 (>3 8 < w < a < 2 2 a w '7 < <0 O z D “ LL 2 < 2 P < (0 0.4 .Correlation function Precipitation a Response function Coefficient -0.4 o‘i oCi ad ad (:11 od 0. CL CL 0. a :1 CL :1 CL .1 z m o: o: >- z .1 o o. 3 g $ 5 c>L 8 < “J < o. < 3 2 3 w '5 < (D O O o " LL 2 < E P < <0 2 Figure 3-5. Correlation and response functions (22 PCs retained, R2=0.3380) for the ARSTAN chronology for Pinus resinosa (23 trees, 82 series) against the mean monthly temperature (T) and monthly precipitation (P) for the previous (p) and current growing seasons at the Grayling Beal Plantation. Significant (p<0.05) responses to a climatic variable are denoted by a *. 115 0.4 Temperature Coefficient .Correlation function a Response function -O.4 lalalalalal-at-l—l—l-l—l-l-l-t- ZCDCKOC>-Z—|0CL 33$558 0 Z in m a: >' Z —‘ (D CL 4: UJ 4: CL < 3 D 3 Lu 2 2 901 8 Cz> ‘5‘ e U- 2 < 2 w a < w 0.4 fireclpltatioq Coefficient .Correlation function a Response function * * _0 4 all If: 05. ad C‘Ll C‘LI (a Ca 0. CL 0. CL 0. CL CL CL a _| 2 (D m K >' Z —’ (9 CL 3 8 fl 5 g 8 g 31' <2: 0. < :3 9, 3 Lu .5 < (D 0 g D E < 2 '3 < (D Figure 3-7. Correlation and response functions (22 PCs retained, R2=0.4680) for the ARSTAN chronology for Pinus sylvestris (5 trees, 20 series) against the mean monthly temperature (T) and monthly precipitation (P) for the previous (p) and current growing seasons at the Grayling Beal Plantation. Significant (p<0.05) responses to a climatic variable are denoted by a *. 117 0.4 * Temperature 0.2 Coefficient O 'Correlation function a Response function -0.4 +3 la la la 13 i3 1- l- i- l— l- i- i- l- l- 2 no a: o: >- z A o o. '5 ‘3 fi 5 (>3 8 at, E < a < a 2 2 w P < <0 O z o 2 < 2 w 4 U) 0.4 .Correlation function Precipitation [3 Response function * 0.2 Coefficient O I' “if -o.2 -o.4 * 2mmm>Z—IOO. 3%&J5%8v woCoE< 552 2 Amv 0:95 .5 dab 528 2: so: :5 9: common 2: 8 02%: £5th 0:5 o—ofi. 2: 8 AZV o>zaz on 9 882250 E 86on 2: 55023 38065 258m 62m: cm.“ mm 360% some .8 33% can .cosfiocowfl .«o 385%? ._o A+Voocom2m .3323 £8.53 35 Ea 53:5: oak .32 .mm .32 co couflm EoEfiomxm _SE_=otm< 9:380 2: pm 5:853 05 E @853 bEEwto 860% com ANNE 595220 .3“: .momv 3:3: .8658 use 2.2328 20 05.3 33— .32 .NS_ 30> :32 mfifitw .39 E355 use .5320 4 m9 muov bub—05$ EovoE no 0:89:00 a m. a: mi... ZO~H=0£wto 0003 860% 00005:: 0:55 €02.20 b.0538 Ho: 0.03 860% 00025 .802 .30— .NS_ mmo>v wut0E< 552 9 Am: 0coxm E .005 30.8 05 Ho: :5 Ev :2w0m 05 9 0300: .3055 0:6 060% 05 8 A70 02002 0n 9 00002200 £ 360% 0:0 650:3 00806.: £58m .cocficmi 00.58 05 nwzefi 5.5: 9 5:8 @552 3002.0: :3 02>» E-N Ew6 5 00.3230 mEBm 3003 .«o 306 0N6 3 E0800 0:0 £0383: £0603 05 $505 038 $5. ZOEEHm HZmEEm—mvmm AEDHADUanN OZE>m mzmkm >QOOB m0 ZOHHDmaHmHQ = X—GZHA—m< 123 APPENDIX III 124 N—é sad 3.: 2:. cod 2.: :6 cmd u cw.— mné gfi mmdw # Each. u: =\.. — M o m N— b— um: .38. Efi 2-3 Elm “ . .u _ . w I ....w: . T .L‘ m m . _. ... ... L. ....z. W : . n ., ...“. .mr. {~‘1. 3 v5 . .WV . W ., _. .u w .. W H .7.: :.7. a (A. w... . . . . u : U .H w m M . m .. . F : . . J. ... 1 t .... .. . .+.. . . ) ...: .+. . m n é. ‘ ,. .a. 5...: .1. : . : . .. 2 . . . 1 . i .....:......?.:. . .l : . . .. . . .. z . ... 1 1.1-721! .. .r. .. . . . m 2-8 8-3 8-0m ‘, .4- ......— ‘A :..”...w..- .r om-m_ 2-0— gmévé—a—gvé on ENEM €—-§~o§—-§Eé~oi—-f—§ mé —~o—~ ,flflflflwwmw w _ 0° ._ a w . ”-.....f ...... EEK 52v :8: f: E 82: mw:::oum EnauuchzmeQEEQQ .33 .wmuxuzm. . . .... . . mebun ..wmzik . . .. .. ,. 5....-. - . .. w3u©§§:% . . ... -.wgmmi .wx:..Q -33: ,3.&% ......f,:-,..,, _...%.mw3u:&u§;w..w . .. . .. . ... . .. .. .. . .. ... :33? puvw .. . . .. 53$. .QUKENW. - ,. .. . . ,, ,. . .. .. .. .. . SEEQWNNVVE..wmu:¥.®- .. .. - . . $.85qu meN . ,. . mgzm€ziziz§mim§z§szfza¢ 863m mafim doom E 6:80:00 803 «SD .C 52251:: oomv BEE: xzmfiwto 20>» 360% 8927:: 233 68:3: b_m:_wto :o: 895 86on 392m Go? .32 .NBE mmo>v «23:? 5.52 8 Am: ocoxm :0 dab 328 2: Ho: :5 Q: :o_wom 05 8 033: .2223 0:3 :33 2: 3 A75 0332 on 8 35238 m_ 360% 05 552—3 3:865 mafim .:o_§:m_a 36:8 of .8 5:8 $3. Bo: E: 2: @385 5.5: 8 528 w:_:::._ 808:8: :3 023 E-m Ewfi :_ Bantam 355 @003 a: $20 36 3 28:0: :5 59:3: .862? 2: .365 03:: £5. ZOHH§O MmEMOm 3.5. OH =~ X—GZHAE< HZmUm $2um >000? m0 ZOEDmEHmHQ 125 APPENDIX IV 126 APPENDIX IV TREE-RING CHRONOLOGIES DEVELOPED FOR PINACEAE AT THE FORMER GRAYLTNG AGRICULTURAL EXPERIMENT STATION AND HARTWICK PINES STATE PARK Raw ring-width data from the Grayling Beal Plantation will be made available via the International Tree- Ring Data Bank (ITRDB), a searchable database maintained by the National Oceanic and Atmospheric Administration; the raw ring-width data from Hartwick Pines State Park (Koop and Grissino-Mayer 1988) were obtained from the 1TRDB. The ARSTAN chronologies (ARSTAN Version 6.04P, Holmes 2000) developed from the raw ring-width data and used to detect climate sensitivity are provided in this Appendix. Picea abies - Grayling Beal Plantation 0 1 2 3 4 5 6 7 8 9 1890 0.08404 1.43938 1.43107 1.39603 1900 1.29877 1.28073 1.43303 1.00533 0.92791 1.10313 1.04524 0.71119 1.04120 0.90107 1910 0.80832 1.14398 1.38590 1.02490 0.80075 1.21506 1.20357 1.14232 1.20835 0.69123 1920 1.25518 0.92842 1.16880 0.74388 0.92843 0.63992 0.77119 1.03799 0.89567 1.00723 1930 1.09741 0.84808 1.22751 0.87582 0.58508 0.81295 0.64786 0.91579 1.12318 1.15133 1940 1.27393 0.66483 1.37923 1.27051 0.83875 1.04109 1.03225 0.88432 0.72814 1.15563 1950 1.07731 1.25330 1.12759 1.56357 0.86753 0.76434 0.85231 1.12626 1.03124 0.79377 1960 0.86967 1.29974 1.01783 1.06073 0.67823 0.73008 0.58899 1.14891 1.28564 1.51652 1970 0.81062 0.59837 0.77447 1.12427 1.13591 0.9821 1 0.82367 0.58984 0.76845 0.76826 1980 1.13584 1.27332 1.14047 0.87961 1.21942 0.89828 1.44441 0.99455 0.68883 0.93927 1990 1.12904 0.98944 0.91218 1.34690 1.06058 0.94806 1.15200 0.78393 0.94272 Pinus resinosa - Grayling Beal Plantation 0 1 2 3 4 5 6 7 8 9 1890 1.96439 1900 2.43952 2.57670 2.43775 1.89389 1.55836 1.57289 1.43844 1.00529 1.01229 0.96009 1910 0.99742 1.04755 1.22553 1.04833 0.97902 0.98792 0.84050 0.82522 0.77350 0.60782 1920 0.81 195 0.84526 0.82151 0.77827 0.89237 0.90320 0.91594 0.95588 0.88049 0.95065 1930 0.80872 0.75125 0.95039 0.83940 0.79672 0.93428 0.70720 0.94590 1.1 1061 1.06583 1940 1.05481 0.79679 1.26030 1.26534 0.94859 1.17426 1.13688 0.97908 0.98163 0.95767 1950 1.02908 1.09990 0.94288 0.92992 0.84436 0.77385 0.89658 1.07589 1.05571 1.05853 1960 0.95119 1.03539 1.24394 1.69134 1.44797 1.52466 1.18901 1.11260 1.16094 1.21263 1970 1.17458 1.12511 1.00029 1.18271 0.97776 0.78475 0.84087 0.84479 0.94355 0.89828 1980 0.90556 0.95539 1.02122 0.93341 1.09181 1.24832 1.30859 1.10902 0.91935 0.68146 1990 0.90618 0.82812 0.68463 0.74560 0.73092 0.70646 0.78510 0.72450 0.75551 127 Pinus resinosa — Hartwick Pines State Park ‘3 0 l 2 3 4 5 6 7 8 9 1770 (180135 (189023 1.00532 1.04032 1.35573 1.58100 1.27730 1.18320 1780 1.16413 1.09316 1.29851 1.33232 1.01233 (181699 1.03759 1.31608 (180931 (168997 1790 (169684 (181209 (173329 (182969 (171525 (174720 (174387 (178854 (185826 (182905 1800 (185337 (187837 1.04492 (177197 (163461 1.00464 1.02451 (193756 (178565 (176367 1810 (196875 1.14908 1.02523 1.11763 (195427 1.19014 1.15041 1.2033 (181285 (191384 1820 (167091 (160759 (183712 (196692 1.16522 1.28095 (193528 1.27780 1.39378 1.19596 1830 1.49255 1.36366 1.18197 1.07623 1.14760 1.04115 1.14344 (189482 (192820 1.03678 1840 1.03419 1.10308 1.33710 1.27777 1.37157 1.00855 (187622 (160100 (194065 (181816 1850 (179852 1.04724 (188066 (193931 (199149 1.04379 (192962 1.12163 1.03779 (196854 1860 (185338 (199392 1.02879 1.06259 (174959 (187158 (183095 (185431 (183106 (199297 1870 1.19791 1.10292 1.06336 1.05123 (165261 (176531 (178546 (170955 1.03705 (188028 1880 (198525 (183336 (197855 1.05804 1.12951 1.08771 1.07586 (188413 (164360 1.07407 1890 (189244 (168602 (194217 (183039 (185930 (175908 1.00656 (179071 (184681 1.03289 1900 1.12470 (193173 1.24011 1.26588 1.13410 1.10846 1.13465 (187449 (179024 (155811 1910 (147048 (151999 (177439 (192303 (179313 (196687 1.13320 (197752 1.00834 (176867 1920 (165983 (160754 (180274 (178019 1.00425 1.19301 1.35123 1.28426 1.25195 1.27641 1930 1.15990 (199674 1.05718 (192099 (164686 (187552 (171880 (160635 (183287 (177387 1940 (171043 (153352 (191388 1.00130 (196624 1.35644 1.34883 1.18691 1.11838 1.02977 1950 1.10741 1.08628 (193235 1.08706 1.04000 (186840 (194688 1.14601 1.05523 1.17225 1960 (188602 1.06097 (198605 1.17927 1.25628 1.27471 (189417 (183301 (194660 1.10216 1970 1.07232 (196337 (193710 1.16352 1.12190 (193828 1.00023 (195172 1.01443 1.03363 1980 1.06145 (196071 1.04805 (180848 (188639 1.22218 1.59275 1.43102 Pinus strobus — Grayling Beal Plantation 0 1 2 3 4 5 6 7 8 9 1890 (147006 (132449 (152785 (160930 (196219 1.26140 1900 1.32293 1.15539 1.35371 (197023 1.06657 1.16925 1.17300 (174131 (199688 (179845 1910 (184897 1.15021 1.18017 1.02561 1.19055 1.14883 (187997 (183806 (166262 (160612 1920 1.06357 1.03448 1.14753 (190311 1.10337 (181206 1.16091 1.31000 1.18902 1.09005 1930 1.00445 (169963 (197187 (170046 (158167 (174338 (168655 1.07645 1.45702 1.35518 1940 1.21660 (159431 1.09276 1.02300 (174446 (194740 (189269 (181860 (198543 (190796 1950 (199231 (183313 (172812 (190514 (174097 (177739 (192651 (197177 (183811 1.00235 1960 1.11997 1.16372 1.15007 1.39816 1.15937 1.36622 (183201 (191373 1.30193 1.35612 1970 1.00831 (189316 (191929 1.11885 (198097 1.11458 1.02006 1.01925 1.19056 (197797 1980 (198136 1.07952 1.14706 (187322 1.24585 1.29811 1.52531 1.06311 1.07203 (197115 1990 (194348 (181095 (172952 (197358 (193017 (192581 (198763 (171287 (181873 Pinus sylvestris — Grayling Beal Plantation 0 1 2 3 4 5 6 7 x 9 1910 113048 1.36331 1.18722 (175155 (172833 1920 (162211 (185759 (168146 (160505 (184234 (166153 (174587 (184901 (196573 1.00229 1930 1.01666 (189568 (197998 (188687 (189105 1.00854 (196404 1.02466 1.19018 1.17288 1940 (194599 (177089 1.13511 1.04000 1.08709 1.29646 1.24909 (192583 (179617 (187743 1950 (187640 (183678 (171670 1.03068 1.01100 (196956 (198926 1.04042 1.04326 (197541 1960 (191611 1.10839 1.02196 1.24645 1.14150 1.15210 (192155 (187404 1.08965 1.16931 1970 1.09661 1.02875 (190765 1.05287 (192014 (176730 (166988 (176814 (186879 (184018 1980 (187727 1.07882 (197755 (189806 1.21369 1.72646 1.57623 1.51191 1.33216 1.26254 1990 1.28343 (183652 (164047 (152277 (152954 (169643 (151017 (147294 (151847 APPENDIX V APPENDIX V CLIMATE DATA Monthly precipitation and mean monthly temperature data used in the correlation analysis were obtained for the Grayling Station (COOP ID 203391) from the National Climatic Data Center (2001). a website maintained by the US. Department of Commerce. National Oceanic and Atmospheric Administration, and National Environmental Satellite, Data. and Information Service. Monthly precipitation (hundredths of inches) Year Jan Feb Mar April May June July Aug Sept Oct Nov Dec 1931 63 32 116 228 524 400 112 220 685 565 457 163 1932 147 144 48 146 506 377 573 492 79 581 134 227 1933 117 94 114 287 391 224 140 80 244 733 177 110 1934 58 20 84 139 126 311 192 206 822 348 258 92 1935 81 85 115 95 91 284 104 393 292 227 186 78 1936 115 75 61 219 397 106 209 343 414 458 161 87 1937 123 94 26 109 96 266 460 260 602 268 206 84 1938 193 206 174 48 348 228 451 525 477 128 201 180 1939 141 128 99 249 490 363 160 706 304 223 133 196 1940 105 58 84 172 441 329 302 685 427 247 441 127 1941 121 130 63 271 275 68 227 282 478 626 323 149 1942 150 25 383 113 527 354 588 128 540 216 227 293 1943 139 185 305 257 462 829 587 352 243 237 470 82 1944 89 104 239 264 315 743 331 194 413 141 317 115 1945 70 111 65 337 779 378 389 397 694 359 346 99 1946 206 139 136 129 327 493 235 222 363 89 296 168 1947 176 119 104 411 689 165 327 236 625 32 213 174 1948 60 80 261 481 201 731 591 72 121 181 537 166 1949 277 142 149 256 203 503 535 339 242 174 241 296 1950 377 220 243 355 182 444 639 429 325 306 201 193 1951 176 160 213 423 306 297 461 374 475 507 291 194 1952 227 86 241 225 148 418 744 398 209 50 538 233 1953 193 358 192 284 434 300 782 359 307 82 187 247 1954 179 144 156 592 357 866 115 120 457 442 142 114 1955 130 104 254 161 262 154 110 159 41 334 277 161 1956 51 96 42 403 223 371 563 746 250 131 287 166 1957 124 61 106 345 357 550 398 164 295 274 348 127 1958 79 57 48 62 89 219 168 296 397 275 313 91 1959 141 206 216 391 402 63 579 717 388 537 275 230 1960 163 167 176 271 589 235 388 216 282 216 425 52 1961 80 164 175 204 185 296 548 496 744 167 328 147 1962 195 204 112 79 297 274 279 359 306 257 80 114 1963 235 72 308 168 446 350 451 301 212 68 342 167 1964 105 36 221 271 417 55 483 217 465 153 260 136 1965 271 123 153 364 251 261 97 901 555 247 227 302 1966 137 92 239 294 69 161 451 183 300 300 589 268 1967 228 94 117 478 220 708 73 301 251 285 344 215 1968 109 199 57 152 332 512 352 264 389 325 205 254 1969 240 24 97 246 317 584 354 67 170 605 228 1(H 1970 143 79 187 160 342 250 383 178 672 250 397 172 1971 217 361 189 208 253 72 314 296 257 89 304 404 1972 90 136 254 163 202 149 365 536 310 282 139 446 1973 182 167 142 206 429 475 236 191 420 317 249 252 1974 307 147 140 422 328 492 254 270 308 214 130 170 1975 279 200 348 186 425 459 495 676 445 172 216 171 1976 221 250 453 145 399 274 207 136 201 194 127 137 1977 132 95 181 197 130 156 525 430 607 322 178 99 APPENDIX V — Continued Monthly precipitation (hundredths of inches) Year Jan Feb Mar April May .1 une July Aug Sept Oct Nov Dec 1978 132 39 87 107 318 263 282 390 724 298 157 182 1979 178 105 359 507 307 260 273 486 28 374 268 135 1980 142 52 96 406 242 624 222 218 456 269 165 187 1981 64 175 74 477 193 249 434 395 218 355 143 61 1982 308 43 259 254 147 288 849 268 415 380 342 387 1983 80 81 175 162 819 100 202 590 493 532 144 198 1984 119 78 212 154 317 505 567 441 491 358 I32 243 1985 144 255 245 199 353 187 334 592 595 379 345 166 1986 98 140 191 204 360 496 487 184 1251 316 85 96 1987 108 60 72 122 219 453 160 903 389 368 196 234 1988 122 120 206 252 162 221 526 536 318 567 621 157 1989 125 82 284 130 343 469 133 419 214 234 284 83 1990 327 113 247 181 420 472 293 323 416 442 282 154 1991 125 61 228 484 624 152 399 264 503 1048 202 182 1992 129 168 179 459 59 223 390 233 499 323 541 155 1993 165 71 43 484 344 676 243 601 466 255 226 84 1994 163 111 168 512 324 394 992 437 347 242 373 45 1995 243 37 189 454 199 274 408 458 212 527 476 231 1996 183 184 59 245 193 453 508 177 506 340 181 209 1997 209 212 197 63 367 102 345 444 204 227 149 38 1998 209 71 417 214 202 326 143 114 202 382 268 167 Mean monthly temperature (tenths of degree Fahrenheit) Year Jan Feb Mar April M ay .lune .1 uly Au g Sept Oct Nov Dec 1931 212 240 254 412 517 650 706 652 628 502 420 275 1932 288 222 220 364 536 644 666 674 585 473 315 246 1933 272 182 273 410 572 694 706 646 632 457 283 203 1934 214 80 210 384 584 670 700 636 588 478 387 204 1935 178 176 297 410 484 606 728 676 552 474 337 192 1936 158 86 301 358 584 618 714 670 606 450 302 270 1937 206 229 244 410 564 634 690 708 586 445 340 202 1938 174 226 346 444 545 641 694 706 560 512 376 254 1939 208 171 248 384 570 652 688 672 588 456 328 275 1940 145 196 208 356 507 626 670 636 573 454 310 245 1941 174 176 200 452 556 640 672 626 596 456 352 272 1942 172 154 285 460 522 620 636 626 534 436 306 168 1943 102 166 178 332 476 620 629 612 492 404 259 158 1944 202 159 182 318 526 604 618 618 574 440 378 198 1945 134 218 418 462 488 604 662 671 575 460 358 190 1946 204 176 398 433 507 621 670 631 597 518 377 230 1947 199 163 224 368 476 598 678 732 602 572 309 234 1948 136 174 257 410 510 624 679 673 621 465 398 1949 239 228 281 434 553 685 703 676 561 526 324 1950 228 192 229 339 543 645 657 626 573 520 326 2 2 2 1951 192 207 279 422 579 616 663 640 557 496 278 23 1952 227 223 259 457 543 668 705 660 605 433 376 285 1953 233 232 306 395 546 657 685 680 590 532 405 278 1954 174 284 257 440 505 670 669 658 583 492 380 227 1955 195 202 267 502 591 655 748 731 586 510 317 209 1956 188 214 243 395 522 658 642 661 551 536 368 266 1957 150 237 296 450 536 656 691 648 574 468 364 266 1958 213 166 310 448 537 591 674 669 583 508 371 159 1959 140 I50 266 419 600 658 687 720 619 458 293 284 131 APPENDIX V — Continued Mean monthly temperature (tenths of degree Fahrenheit) Year Jan Feb Mar April May .lune July Aug Sept Oct Nov Dec 1960 217 197 195 451 556 608 658 666 611 485 393 196 1961 163 243 329 402 522 629 678 658 637 506 361 238 1962 165 151 291 430 600 635 650 661 557 495 351 227 1963 113 108 294 458 521 662 688 628 574 573 404 186 1964 235 222 282 449 593 640 694 633 574 460 398 218 1965 170 192 236 394 590 622 647 649 580 473 361 290 1966 148 223 338 401 490 665 719 665 578 472 355 237 1967 233 154 288 438 491 669 656 629 576 462 309 253 1968 175 135 333 464 508 627 674 672 613 497 354 218 1969 199 201 253 438 537 584 675 681 587 445 328 209 1970 122 157 239 432 557 644 706 673 574 468 358 242 1971 152 183 243 389 521 678 654 645 618 557 350 266 1972 189 162 206 349 566 591 664 645 558 414 327 219 1973 220 161 372 414 498 645 668 686 558 504 350 222 1974 194 124 271 417 490 598 666 641 532 437 363 261 1975 194 206 235 346 583 622 662 649 532 478 402 224 1976 146 229 276 438 493 652 669 631 541 416 277 136 1977 99 158 335 450 601 612 692 609 579 435 336 200 1978 136 98 207 384 558 609 651 645 571 425 349 189 1979 106 72 289 385 497 614 656 604 583 450 349 252 1980 188 121 210 407 550 585 669 673 555 412 320 168 1981 118 220 305 440 514 627 666 652 542 414 338 234 1982 89 143 228 361 594 576 672 609 561 489 348 308 1983 199 235 303 380 463 617 716 680 582 446 344 156 1984 112 262 209 438 492 646 655 677 535 473 335 260 1985 146 165 286 445 552 584 651 627 580 451 316 174 1986 160 171 284 458 559 588 683 603 557 429 308 257 1987 189 201 310 439 558 656 699 637 574 413 356 267 1988 149 153 242 405 552 652 695 676 558 412 355 216 1989 216 130 213 386 520 608 679 626 546 453 290 101 1990 246 196 293 426 499 615 660 641 560 429 369 245 1991 136 219 295 447 590 663 670 667 544 460 313 238 1992 200 200 248 3737 522 591 609 602 541 426 308 241 1993 176 123 260 372 520 591 679 665 517 421 323 229 1994 59 80 268 399 512 644 660 621 583 477 361 282 1995 211 151 301 364 508 673 675 696 530 474 263 166 1996 118 148 215 348 487 626 635 654 582 450 280 239 1997 149 196 242 374 447 636 660 609 559 446 308 276 1998 214 294 291 427 596 622 663 669 602 480 366 279 LIST OF REFERENCES LIST OF REFERENCES Abrams, M.D., and D1 Dickmann. 1984. F loristic composition before and after prescribed fire on a jack pine clear-cut site in northern lower Michigan. Can J For Res 14:746-749. Abrams, M.D., D.G. Sprugel, and DJ. Dickmann. 1985. Multiple successional pathways on recently disturbed jack pine sites in Michigan. For Ecol Mgt 10:31-48. Albert, DA. 1994. Regional Landscape Ecosystems of Michigan, Minnesota, and Wisconsin: A Working Map and Classification (Fourth Revision). Gen. Tech. Rep. NC-178. St. Paul, MN: US Dept of Agriculture, Forest Service, North Central Forest Experiment Station. 25pp. American Forest and Paper Association. 2001. Sustainable Forestry Initiative (SFI)SM Program. http://www.afandpa.org/forestry/sfi_frame.html Anderson, RC, and M.L. Bowles. 1999. Deep-soil savannas and barrens of the Midwestern United States. Pages 155-170 in RC. Anderson, J .S. F ralish, and J .M. Baskin, editors. Savannas, Barrens, and Rock Outcrop Plant Communities of North America. Cambridge Univ Press, New York, NY, USA. 470pp. Anonymous. 1913. Notes copied from those taken by Dr. Beal and others at the dates indicated. William J. Beal Papers, Collection UA17.4, Box 891 , Folder 14, Michigan State University Archives and Historical Collections. East Lansing. MI, USA. Avalanche. 1888a. No title. April 26, 1888, Grayling, MI, USA.. Avalanche. 1888b. No title. May 17, 1888, Grayling, MI, USA. Avalanche. 1888c. Experiment Station. June 7, 1888, Grayling. MI. USA. Avalanche. 1888d. No title. July 5, 1888, Grayling, MI, USA. Barbour, M.G., J .H. Burk. W.D. Pitts, F.S. Gilliam, and M.W. Schwartz. 1999. Terrestrial Plant Ecology, third edition. Benjamin/Cummings, Menlo Park. CA, USA. 649pp. Barnes, B.V., and W.H. Wagner, Jr. 1981. Michigan Trees. Univ of Mich Press, Ann Arbor, MI, USA. 384pp. Barnes, B.V., D.R. Zak, S.R. Denton, and SH. Spurr. 1998. Forest Ecology. 4th edition. John Wiley & Sons, Inc.. New York, NY, USA. 774pp. 9‘ ‘9‘): xxh- 5‘ Barrett, L.R., J. Liebens, D.G. Brown, R.J. Schaetzl, P. Zuwerink, T.W. Gate, and D.S. Nolan. 1995. Relationships between soils and presettlement forests in Baraga County, Michigan. Am Midl Nat 134:264-285. Barrett, LR, and R.J. Schaetzl. 1998. Regressive pedogenesis following a century of deforestation: evidence for depodzolization. Soil Sci 163(6):482-497. Beal, W.J. 1878. Report of the Professor of Botany and Horticulture. Pages 64-84 in Sixteenth Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1877. W.S. George & Co., State Printers and Binders, Lansing, MI, USA. 643pp. Beal, W.J. 1888a. Observations on the succession of forests in northern Michigan. Pages 74-78 in Twenty-Seventh Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1888. Thorp and Godfrey, State Printers and Binders, Lansing, MI, USA. 596pp. Beal, W.J. 1888b. Report of the Botanist of the Experiment Station. Pages 171-193 in Twenty-Seventh Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1888. Thorp and Godfrey, State Printers and Binders, Lansing, MI, USA. 596pp. Beal, W.J. 1889a. Report of the Botanist of the Experiment Station. Pages 160-167 in Twenty-Eighth Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1889. Darius D. Thorp, State Printer and Binder, Lansing, MI, USA. 618pp. Beal, W.J. 1889b. Experiments and observations on the jack-pine plains, Bulletin No. 54 — Botanical Department. Pages 321-327 in Twenty-Eighth Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1889. Darius D. Thorp, State Printer and Binder, Lansing, MI, USA. 618pp. Beal, W.J. 1890. Report of the Botanist. Pages 89-100 in Twenty-Ninth Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1890. Robert Smith & Co., Lansing. MI, USA. 559pp. Beal, W.J. 1910. Report of the Department of Botany. Pages 84-99 in Forty-Ninth Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1910. Wynkoop Hallenbeck Crawford Co., Lansing, MI, USA. 614pp. Beal, W.J. 1915. History ofthe Michigan Agricultural College. Wynkoop Hallenbeck Crawford Co., Lansing, MI, USA. 519pp. Bergeron, Y., and M. Dubuc. 1989. Succession in the southern part ofthe Canadian boreal forest. Vegetatio 79:51-63. 135 m""1 Bertness, M.D., and RM. Callaway. 1994. Positive interactions in communities. Trends Ecol Evol 92191-193. Blasing, T.J., A.M. Solomon, and D.N. Duvick. 1984. Response functions revisited. Tree-Ring Bull 4421-15. Bogue, BE. 1903. Report of Forestry Department. Pages 42-46 in Forty-Second Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1903. Robert Smith Printing Co., State Printers and Binders, Lansing, MI, USA. 363pp. Borak, R.A., Regional For Mgr, Mich Dept Natl Res. 1983. Unpublished memorandum 1"“ to Michael D. Moore, Asst Div Chief, For Operations, For Mgt Div, Mich Dept Natl Res, dated 28 September. Beal Plantation File, For Mgt Div, Mich Dept Natl . Res, Lansing, MI, USA. " Botti, W.B., Program Leader, Timber Mgt, For Mgt Div, Mich Dept Natl Res. 1988. Unpublished memorandum to Donald B. Grant, Assistant Chief, For Mgt Div, Mich Dept Natl Res, dated 26 May. Beal Plantation File, For Mgt Div, Mich Dept 1 Natl Res, Lansing, MI, USA. Braunschweig, S.H., E.T. Nilsen, and T.F. Wieboldt. 1999. The Mid-Appalachian shale barrens. Pages 83-98 in RC. Anderson, J .S. Fralish, and J .M. Baskin, editors. Savannas, Barrens, and Rock Outcrop Plant Communities of North America. Cambridge Univ Press, New York, NY, USA. 470pp. F orman, R.T.T. 1998. Pine Barrens: Ecosystem and Landscape. Rutgers Univ Press, New Brunswick, NY, USA. 601pp. Briffa, K., and RD. Jones. 1990. Basic chronology statistics and assessment. Pages 137- 152 in ER. Cook and LA. Kairiukstis, editors. Methods of Dendrochronology: Applications in the Environmental Sciences. Kluwer Academic Publishers, Dordrecht, The Netherlands. 394pp. Burns, RM. and B.H. Honkala, tech. coords. 1990a. Silvics of North America: 1. Conifers. Agric Handbook 654. US Dept of Agric, For Serv, Washington, DC, USA. 681pp. Burns, RM. and B.H. Honkala, tech. coords. 1990b. Silvics of North America: 2. Hardwoods. Agric Handbook 654. US Dept of Agric, For Serv, Washington. DC, USA. 877pp. Callaway, R.M., and LR. Walker. 1997. Competition and facilitation: a synthetic approach to interactions in plant communities. Ecol 78(7): 1958-1967. 136 Carmean, W.H., and J .T. Hahn. 1981. Revised site index curves for balsam fir and white spruce in the Lake States. Res. Note NC-269. US Dept of Agric, For Serv, St. Paul, MN, USA. 4pp. Carmean, W.H., J .T. Hahn, and RD. Jacobs. 1989. Site index curves for forest species in the eastern United States. Gen Tech Rep NC-128. US Dept of Agric, For Serv, St. Paul, MN, USA. 142pp. Chittenden, AK. 1921. Report of the Forestry Section. Page 286 in Fifty-Ninth Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1921. Wynkoop Hallenbeck Crawford Co., State Printers, Lansing, MI, USA. 700pp. Chittenden, AK. 1922. Forest planting in Michigan, Special Bulletin 103. Pages 468-481 in Sixtieth Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1922. Wynkoop Hallenbeck Crawford Co., State Printers, Lansing, MI, USA. 636pp. Clark, B.G., President, Michigan Forest Association. 1979. Unpublished letter to Dr. Henry Webster, Chief, For Mgt Div, Mich Dept Natl Res, dated 25 January. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Clements, F .E. 1916. Plant Succession: An Analysis of the Development of Vegetation. Carnegie Inst Pub 242. Washington, DC, USA. 512pp. Coates, A.J., District Forest Supervisor, Mich Dept Natl Res. 1978. Unpublished memorandum to Robert A. Borak, Regional Forest Supervisor, Mich Dept Natl Res, dated 2 May. Subject: Possible Exchange - City of Grayling. Beal Plantation File, For Mgt Div, MI Dept Natl Res, Lansing, MI, USA. Coates, K.D., and P1 Burton. 1997. A gap-based approach for development of Silvicultural systems to address ecosystem management objectives. For Ecol M gt 99:337-354. Cohen, J .P. 1996. Natural community abstract for oak-pine barrens. Michigan Natural Features Inventory. Lansing. MI, USA. 6pp. Comer, R]. 1996. Natural community abstract for pine barrens. Michigan Natural Features Inventory, Lansing, MI, USA. 3pp. Connell, J .H. 1990. Apparent vs. “real” competition in plants. Pages 9-26 in J .8. Grace and D. Tilman, editors. Perspectives on plant competition. Academic Press, New York, NY, USA. 484pp. 137 Cook, E.R., A.H. Johnson, and T.J. Blasing. 1987. Forest decline: modeling the effect of climate in tree rings. Tree Phys 3:27-40. Cook, E.R., K.R. Briffa, SG. Shiyatov, and V. Mazepa. 1990a. Tree-ring standardization and growth-trend estimation. Pages 104-123 in ER. Cook and L.A. Kairiukstis, editors. Methods of Dendrochronology: Applications in the Environmental Sciences. Kluwer Academic Publishers, Dordrecht, The Netherlands. 394pp. Cook, E.R., SG. Shiyatov, and V. Mazepa. 1990b. Estimation of the mean chronology. Pages 123-132 in ER. Cook and L.A. Kairiukstis, editors. Methods of Dendrochronology: Applications in the Environmental Sciences. Kluwer Academic Publishers, Dordrecht, The Netherlands. 394pp. Cook, ER, and R.L. Holmes. 1997. ARSTAN: chronology development. Pages 75-87 in lntemational Tree-Ring Data Bank, H.D. Grissino-Mayer, R.L. Holmes, and HG Fritts, editors. Documentation to the lntemational Tree-Ring Data Bank Program Library Version 2.1. http://tree.ltrr.arizona.edu/software.html Cowles, H.C. 191 1. The causes of vegetative cycles. Bot Gaz 51:161-183. Crawford County Equalization Department. 1884. Entry for William Brinks. Tax Rolls for Crawford County, Grayling, MI, USA. Crawford County Equalization Department. 1886. Entry for William Brinks. Tax Rolls for Crawford County, Grayling, MI, USA. Crawford County Equalization Department. 1887. Entry for J .L. & S. Railroad Company. Tax Rolls for Crawford County. Grayling, MI, USA. Crawford County Equalization Department. 1888. Entry for Martha Brinks. Tax Rolls for Crawford County, Grayling. MI, USA. p.48. Crozier, AA. 1895. Notes at the Grayling Station, Nov. 13-14, 1895. William J. Beal Papers, Collection UA17.4, Box 891, Folder 14, Michigan State University Archives and Historical Collections. East Lansing, MI, USA. Curtis, J .T. 1971. The Vegetation of Wisconsin: An Ordination of Plant Communities. Univ Wisc Press, Madison, WI, USA. 657pp. Daw, TE. 1968. Unpublished letter to Ralph A. MacMullan. Director, Michigan Department of Conservation, dated 10 June. Subject: Beal Plantation, Crawford County. Lansing, MI, USA. Dempsey, D. 2001. Ruin & Recovery: Michigan‘s Rise as a Conservation Leader. Univ of Mich Press, Ann Arbor. MI, USA. 336pp. Dickmann, D.I., and L.A. Leefers. (In Press) The Forests of Michigan. Univ of Mich Press, Ann Arbor, MI, USA. Dilts, D.A., Reg. Park Supervisor, Parks and Rec Div, Mich Dept Natl Res. 1994. Unpublished email to W. Botti, dated 2 January. Subject: Beal Property, Grayling. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Dolling, A. 1996. Interference of bracken (Pteridium aquilinum L. Kuhn) with Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies L. Karst.) seedling establishment. For Ecol Mgt 88:227-235. Dolling, A. 1999. The vegetative spread of Pteridium aquilinum in a hemiboreal forest — invasion or revegetation? For Ecol Mgt 124:177-184. Eckert, K.B., Supervisor, Historic Sites Research Unit, Mich History Div, Mich Dept State. 1979. Unpublished letter to Weldon J. Montgomery, Dept Natl Res, dated '1 February. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Egler, F .E. 1954. Vegetation science concepts. 1. Initial Floristic Composition, a factor in old-field vegetation development. Vegetatio 4:412-417. Fastie, CL. 1995. Causes and ecosystem consequences of multiple pathways of primary succession at Glacier Bay, Alaska. Ecol 76:1899-1916. Fowler, B.J., Supervisor, Grayling Township. 1979. Unpublished letter to CE. Rademacher, Tax Lands and Service, Land Div, Mich Dept Natl Res, dated 10 May. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. F ritts, H.C. 1966. Growth-rings of trees: their correlation with climate. Science 154:973- 979. Fritts, H.C. 1976. Tree Rings and Climate. Academic Press, New York, NY. USA. 567pp. Fritts, HO, and X. Wu. 1986. A comparison between response-function analysis and other techniques. Tree-Ring Bull 46:31-46. F ritts, HO, and T.W. Swetnam. 1989. Dendroecology: a tool for evaluating variations in past and present forest environments. Adv Ecol Res 19:111-188. Gaines, E.M., and E.W. Shaw. 1959. Half a century of research ~ Fort Valley Experimental Forest 1908-1958. J For 57:629-633. 139 Garfield, CW. 1905. Prefatory Notes. Report of the Michigan Forestry Commission for the Years 1903-1904. Wynkoop Hallenbeck Crawford Co., State Printers, Lansing, MI, USA. Gevorkiantz, SR. 1957a. Site index curves for red pine in the Lake States. Tech Note 484. US Dept of Agric, For Serv, St. Paul, MN, USA. 2pp. Gevorkiantz, SR. 1957b. Site index curves for white pine in the Lake States. Tech Note 483. US Dept of Agric, For Serv, St. Paul, MN, USA. 2pp. Gibson, D.J., SL. Collins, and RE. Good. 1988. Ecosystem fragmentation of oak-pine forest in the New Jersey Pinelands. For Ecol Mgt 25:105-122. Gibson, D.J., R.A. Zampella, and AG. Windisch. 1999. New Jersey Pine Plains: the "true barrens" of the New Jersey Pine Barrens. Pages 52-66 in RC. Anderson, J .S. Fralish, and J .M. Baskin, editors. Savannas, Barrens, and Rock Outcrop Plant Communities of North America. Cambridge Univ Press, New York, NY, USA. 470pp. Gleason, HA. 1926. The individualistic concept ofthe plant association. Bull Torrey Bot Club 5327-26. Gleason, H.A., and A. Cronquist. 1991. Manual of Vascular Plants of Northeastern United States and Adjacent Canada, Second Edition. The New York Botanical Garden, Bronx, NY, USA. 910pp. Gower, S.T., P.B. Reich, and Y. Son. 1993. Canopy dynamics and aboveground production of five tree species with different leaf longevities. Tree Phys 12:327- 345. Graumlich, L1 1993. Response of tree growth to climatic variation in the mixed conifer and deciduous forest of the Upper Great Lakes region. Can J For Res 23: 133-143. Graumlich, L.J., and MB. Davis. 1993. Holocene variation in spatial scales of vegetation pattern in the uuper Great Lakes. Ecol 74(3):826-839. Grimes, J .P. 1979. Plant Strategies and Vegetation Processes. Wiley, New York, NY, USA. 222pp. Gutierrez, E. 1989. Dendroclimatological study of Pinus sylvestris L. in southern Catalonia (Spain). Tree-Ring Bulletin 46: 1-9. Harding, RB, and DE Grigal. 1985. Individual tree biomass estimations for plantation- grown white spruce in northern Minnesota. Can J For Res 15:73 8-739. 140 Harmes, R., Jr., Acting Chief, Lands Div, Mich Dept Natl Res. 1981. Unpublished letter to Jerry W. Morford. City Manager, Grayling, MI, dated 17 February. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Harris, C.D., Chief, Bureau of Renewable Res Mgt, Mich Dept Natl Res. 1978. Unpublished memorandum to Wayne H. Tody, Deputy Director, Mich Dept Natl Res, dated 9 November. Subject: Land Exchange - City of Grayling. Beal Plantation File, For Mgt Div, MI Dept Natl Res, Lansing, MI, USA. Harwood, RM. 1892. Report of the Agriculturist. Pages 175-177 in Twenty-First Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1892. Robert Smith & Co., Lansing, MI, USA. 607pp. —_:I '0 in 1 .‘l Hees, M.D., President, Monarch Millwork, Inc. Subject: Summary of facts regarding proposed Monarch expansion, dated 25 February. Beal Plantation F ile, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Holmes, R. 2000. Dendrochronology Program Library. 3 ftp://ftp.cricyt.edu.ar/users/dendro/. E Holmgren, M., M. Scheffer, and MA. Huston. 1997. The interplay of facilitation and competition in plant communities. Ecol 78:1966-1975 Host, G.E., K.S. Pregitzer, C.W. Ramm, D.P. Lusch, and D.T. CIeIand. 1988. Variation in overstory biomass among glacial landforms and ecological land units in northwestern Lower Michigan. Can J For Res 18(6): 659-668. Host, GE, and KS Pregitzer. 1992. Geomorphic influences on ground-flora and overstory composition in upland forests of northwestern lower Michigan. Can J For Res 22: 1547-1 555. Huston, M.A., and T.M. Smith. 1987. Plant succession: life history and competition. Amer Nat 130:168-198. Jaderlund, A., O]. Zackrisson, A. Dahlberg, and M. Nilsson. 1997.1nterference of Vaccim’um myrtillus on establishment, growth, and nutrition of Picea abies seedlings in a northern boreal site. Can J For Res 27:2017-2025. Jokela, E.J., K.P. Van Gurp, R.D. Briggs. and EH. White. 1986. Biomass estimation equations for Norway spruce in New York. Can J For Res 16:413-415. Keddy, P.A., and CG. Drummond. 1996. Ecological properties for the evaluation, management, and restoration of temperate deciduous forest ecosystems. Ecol Appl 6(3):748-762. 141 Kedzie, R.C. 1888a. Proposals for Material and Labor. The Avalanche, Grayling, MI, USA, 29 March. Kedzie, R.C. 1888b. The jack pine plains, Bulletin No. 37 - Chemical Department. Pages 207-211 in Twenty-Seventh Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1888. Thorp and Godfrey, State Printers and Binders, Lansing, MI, USA. 596pp. Kedzie, R.C. 1889a. Report of the Chemist of the Experiment Station. Pages 78—143 in Twenty-Eighth Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1889. Darius D. Thorp, State Printer and Binder, Lansing, MI, USA. 618pp. Kedzie, R.C. 1889b. The problem of the plains. Pages 393-401 in Twenty-Eighth Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan. 1889. Darius D. Thorp, State Printer and Binder, Lansing, MI, USA. 618pp. Kedzie, RC. 1890. Report of the Chemist of the Experiment Station. Pages 129-135 in Twenty-Ninth Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1890. Robert Smith & Co., Lansing, MI, USA. 559pp. Kedzie, RC. 1891. Report of the Chemical Department of the Experiment Station. Pages 74-75 in Thirtieth Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1891. Robert Smith & Co., Lansing, MI, USA. 689pp. Kellman, M., and M. Kading. 1992. Facilitation of tree seedling establishment in a sand dune succession. J Veg Science 3:679-688. Kobe, R.K., SW. Pacala, J.A. Silander Jr., and CD. Canham. 1995. Juvenile tree survivorship as a component of shade tolerance. Ecol Appl 5(2):517-532. Kobe, R.K. 1996. lntraspecific variation in sapling mortality and growth predicts geographic variation in forest composition. Ecol Mon 66(2): 181-201. Koop, D.L. 1985. Reconstruction of past climate in northern lower Michigan through the analysis of red pine (Pinus resinosa Ait.) tree rings. Masters thesis, University of Georgia, Athens, GA, USA 77pp. Koop, D.L, and H.D. Grissino-Mayer. 1988. Red pine at Hartwick Pines State Park, Michigan. lntemational Tree-Ring Data Bank, NOAA Paleoclimatology Program and World Data Center for Paleoclimatology, NOAA/NGDC Paleoclimatology Program, Boulder, CO, USA. http://www.ngdc.noaa.gov/paleo/ftp-treering.html. Kozlowski, T.T. 1971. Growth and Development of Trees, Volume 1: Seed Germination, Ontogeny, and Shoot Growth. Academic Press, New York, NY, USA. 443pp. 142 Kreger, J ., Mich Hist Div. 1978. Unpublished letter to File, Michigan Department of State, dated 30 May. Subject: F ieldtrip report on the Beal Plantation, Grayling, Crawford County. Lansing, MI, USA. Kreger, J .L., Coordinator, Mich Hist Div. Mich Dept State. 1983. Unpublished memorandum to Dean Sandell. dated 20 July. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Ktjchler, A.W. 1964. Potential Natural Vegetation of the Conterminous United States. American Geographical Society, Spec Publ No. 36. 1 l6pp. La Marche, V.C, Jr. 1978. Tree-ring evidence of past climatic variability. Nature 276(5686):334-338. Laylin, R.L., Asst Acquisition Supervisor, Lands Div, Mich Dept Natl Res. 1981. Unpublished letter to Jerry W. Morford, City Manager, Grayling, MI, dated 12 May. Beal Plantation File, For Mgt Div, Mich Dept Natl Res. Lansing, MI, USA. Lichter, J. 2000. Colonization constraints during primary succession on coastal Lake Michigan sand dunes. J Ecol 88:825-839. Little, S. 1998. Fire and plant succession in the New Jersey pine barrens. Pages 297-3 14 in R.T.T. Forman, editor. Pine Barrens: Ecosystem and Landscape. Rutgers Univ Press, New Brunswick, NJ, USA. 601pp. Livingston, BE. 1905. The relation of soils to natural vegetation in Roscommon and Crawford Counties, Michigan. Bot Gaz 39:22-41. MacDonald, G.M., L.C. Cwynar, and C. Whitlock. The late Quaternary dynamics of pines in northern North America. Pages 122-136 in D.M. Richardson, editor. The Ecology and Biogeography of Pinus. Cambridge Univ Press. Cambridge, UK. 527pp. Makinen, H., P. Nojd, and K. Mielikainen. 2001. Climatic signal in annual growth variation in damaged and healthy stands of Norway spruce [Picea abies (L.) Karst.] in southern Finland. Trees 15:177-185. Manley, W.L., President, Michigan Forest Association. 1978a. Unpublished letter to Howard A. Tanner, Director, Natl Res Comm, dated 25 April. Beal Plantation File, For Mgt Div, MI Dept Natl Res, Lansing, MI, USA. Manley, W.L., President, Michigan Forest Association. 1978b. Unpublished letter to Henry H. Webster, Chief, For Mgt Div, Mich Dept Natl Res, no date. received 21 July. Beal Plantation File, For Mgt Div, MI Dept Natl Res, Lansing, MI, USA. Matlack, GR. 1994. Plant species migration in a mixed-history forest landscape in eastern North America. Ecol 75(5): 1491-1502. McMillan, J., Area For Mgr, For Mgt Div, Mich Dept Natl Res. 1993. Unpublished memorandum to Bill Botti, Forester, For Mgt Div, Mich Dept Natl Res, dated 28 April. Beal Plantation F ile, For Mgt Div, Mich Dept Natl Res. Lansing, MI, USA. McMillan, J. Area Mgr, For Mgt Div, Mich Dept Natl Res, and R. Rasmussen, Chairman, Huron Pines RC&D Forestry Committee. 1996. Unpublished letter to Jack Pilon, Mich Dept Natl Res, dated 9 September. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI. USA. Mokma, D.L., and GP. Vance. 1989. Forest vegetation and origin of some spodic horizons, Michigan. Geoderma 43:311-324. Montgomery, W.J. 1977. Unpublished memorandum to William Tarr, Area Forester, AuSable State Forest, Mich Dept Natl Res, dated 26 October. Subject: Salvage of Blow Down in the Beal Plantation. Beal Plantation File, For Mgt Div, MI Dept Natl Res, Lansing. MI, USA. Moore, M.D., and W.B. Botti. 1976. Michigan’s Famous and Historic Trees. Mich For Assoc, Corunna, MI, USA. 28pp. Moore, M.D., Asst Div Chief, For Operations, For Mgt Div, Mich Dept Natl Res. 1983a. Unpublished memorandum to Robert A. Borak, Regional For Mgr, Mich Dept Natl Res, dated 5 July. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Moore, M.D., Asst Div Chief, For Operations, For Mgt Div, Mich Dept Natl Res. 1983b. Unpublished letter to Jerry Morford, City Manager, Grayling, MI, dated 20 December. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI. USA. Moore, M.D., Asst Div Chief, For Operations, For Mgt Div, Mich Dept Natl Res. 1984. Unpublished letter to Jerry Morford, City Manager, Grayling, M1, dated 16 May. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Morford, J .W., City Manager, Grayling, MI. I977a. Unpublished letter to Theodore Tucker, Chief, Lands Div, M1 Dept Natl Res, dated 24 January. Beal Plantation File, For Mgt Div, MI Dept Natl Res, Lansing, MI, USA. Morford, J .W., City Manager, Grayling, MI. 1977b. Unpublished letter to Theodore Tucker, Chief, Lands Div, Ml Dept Natl Res, dated 10 February. Beal Plantation File, For Mgt Div, MI Dept Natl Res, Lansing, MI, USA. 144 Morford, J .W., City Manager, Grayling, MI. 1977c. Unpublished letter to Theodore Tucker, Chief, Lands Div, Mich Dept Natl Res, dated 3 May. Beal Plantation F ile, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Morford, J .W., City Manager, Grayling. MI. 1978a. Unpublished letter to Harry H. Whitely, Natl Res Comm, dated 6 January. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Morford, J.W., City Manager, Grayling, MI. 1978b. Unpublished memorandum to file, dated 1 December. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Morford, J .W., City Manager, Grayling. MI. 1979. Unpublished letter to Jay A. Schafer, Easements-Exchanges, Lands Div, Mich Dept Natl Res, dated 7 December. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Morford, J.W., City Manager, Grayling, MI. 1980. Unpublished letter to Ted Tucker, Chief, Lands Div, Mich Dept Natl Res, dated 16 January. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Morford, J .W., City Manager, Grayling, MI. 1983. Unpublished letter to Robin Bertsch, For Res Development. Mich Dept Natl Res, dated 26 May. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Morford, J .W., City Manager, Grayling, Ml. 1984a. Unpublished letter to Roger Rehberg. Mich Business Ombudsman Office, Mich Dept Commerce, dated 12 April. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Morford, J .W., City Manager, Grayling, MI. 1984b. Unpublished letter to Michael More, Asst State Forester, For Mgt Div, Mich Dept Natl Res, annotated May. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Morford, J.W., City Manager, Grayling, MI. 1993. Unpublished letter Jim McMillan, For Mgt Div, Mich Dept Natl Res, dated 26 April. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing. MI, USA. Morford, S, Secretary, Forestry Club, Mich State Univ. 1978. Unpublished letter to Weldon Montgomery, For Mgt Div, Mich Dept Natl Res, dated 6 June. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. National Climatic Data Center. 2001. Monthly climate data for Grayling (COOP ID 203391). US Department of Commerce, National Oceanic and Atmospheric Administration, and National Environmental Satellite, Data and Information Service. http://lwf.ncdc.noaa.gov/oa/ncdc.html. 145 Nowak, W., Executive Director, Grayling Regional Chamber of Commerce. 1979. Unpublished letter to Harry Whitely, dated 28 November. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Oliver, OD, and BL. Larson. 1996. Forest Stand Dynamics, Update Edition. John Wiley & Sons, Inc. New York, NY, USA. 520pp. Otis, OH. 1931. Michigan Trees: A Handbook of the Native and Most Important Introduced Species, 9th edition. Regents of the Univ of Mich., Ann Arbor, MI. USA. 362pp. Overlease, W.R. 1964. A Study of Variation in the Red Oak Group in Michigan and Nearby States. Ph.D. Dissertation, Mich State Univ, East Lansing, MI, USA. 191pp. Palik, B]. and KS Pregitzer. 1993. The vertical development of early successional forests in northern Michigan, USA. J Ecol 81:271-285. Perala, D.A., and DH. Alban. 1994. Allometric biomass estimators for aspen-dominated ecosystems in the Upper Great Lakes. Res. Paper NC-314. St. Paul, MN: US Department of Agriculture, Forest Service, North Central Forest Experiment Station. 38pp. Perez, R., Wildlife Habitat Biologist, Mich Dept Natl Res. 1978. Unpublished memorandum to Thomas V. Havard, District Wildlife Biologist, Mich Dept Natl Res, dated 16 March. Subject: City of Grayling Land Exchange. Beal Plantation File, For Mgt Div, MI Dept Natl Res, Lansing, MI, USA. Pregitzer, KS, and SC. Saunders. 1999. Jack pine barrens of the northern Great Lakes region. Pages 343-361 in RC. Anderson, J .S. Fralish, and J .M. Baskin, editors. Savannas, Barrens. and Rock Outcrop Plant Communities of North America. Cambridge Univ Press, Cambridge, UK. 470pp. Price, R.A., A. Liston, and SH. Strauss. 1998. Phylogeny and systematics of Pinus. Pages 49-68 in D.M. Richardson, editor. The Ecology and Biogeography of Pinus. Cambridge Univ Press, Cambridge, UK. 527pp. Rasmussen, R., Huron Pines RC&D Council. 1997. Unpublished letter to Gerald J. Thiede, State Forester, Mich Dept Natl Res, dated 13 February. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Rasmussen, R., Huron Pines RC&D Council. 1998. Unpublished letter to George W. Daniel, Georgia Pacific Corporation, dated 6 April. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. 146 Radeloff, V.C., DJ. Mladenoff, H.S He, and M.S. Boyce. 1999. Forest landscape change in the northwestern Wisconsin Pine Barrens from pre-European settlement to the present. Can J For Res 29: 1649-1659. Rehberg, R.R., Regional Director, Upper Great Lakes Regional Comm. 1979. Unpublished letter to Jerry Morford, City Manager, Grayling. dated 11 January. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Rejmanek, M., and D.M. Richardson. 1996. What attributes make some plant species more invasive? Ecol 77:1655-1661. Reuschel, T., Sect Leader, State For Operations, For Mgt Div, Mich Dept Natl Res. -— 1994a. Unpublished memorandum to Jim McMillan, For Area Mgr, For Mgt Div, Mich Dept Natl Res, dated 4 March. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Reuschel, T., Sect Leader, State For Operations, For Mgt Div, Mich Dept Natl Res. 1994b. Unpublished letter to Jerry Morford, City Manager, Grayling, MI, dated 5 April. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Reuschel, T., Sect Leader, State For Operations, For Mgt Div, Mich Dept Natl Res. 1994c. Unpublished memorandum to Bill Botti, For Mgt Div, Mich Dept Natl Res, dated 22 August. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Reuschel, T., Sect Leader, State For Operations, For Mgt Div, Mich Dept Natl Res. 1996. Unpublished letter to Roger M. Rasmussen, Chairman, Huron Pines RC&D Forestry Committee, dated 18 November. Beal Plantation File, For Mgt Div, Mich Dept Natl Res. Lansing. MI, USA. Reynolds, H.G. 1888a. The law providing for a State Forestry Commission, Public Acts of Michigan, 1887 - Act No. 259. Page 70 in Twenty—Seventh Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1888. Thorp and Godfrey, State Printers and Binders, Lansing, MI, USA. 596pp. Reynolds, H.G. 1888b. Questions sent to supervisors and their replies. Pages 102-108 in Twenty-Seventh Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1888. Thorp and Godfrey, State Printers and Binders, Lansing, MI, USA. 596pp. Reynolds, H.G. 1888c. Recommendations. Page 128 in Twenty-Seventh Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1888. Thorp and Godfrey, State Printers and Binders, Lansing, MI, USA. 596pp. Reynolds, H.G. 1888d. Proceedings of the Forestry Convention held in Grand Rapids, Michigan, January 26 and 27, 1888. Bulletin No. 32 of the Michigan Agricultural 147 College. Pages 305-3 77 in Twenty-Seventh Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1888. Thorp and Godfrey, State Printers and Binders, Lansing, MI, USA. 596pp. Richardson, D.M., Williams, P.A., and R.J. Hobbs. 1994. Pine invasions in the Southern Hemisphere: determinants of spread and invadability. J Biogeography 21 :51 l- 527. Richardson, D.M., and SI. Higgins. 1998. Pines as invaders in the southern hemisphere. Pages 450-473 in D.M. Richardson, editor. The Ecology and Biogeography of Pinus. Cambridge Univ Press, Cambridge, UK. 527pp. Rodgers, AD. 111. 1991. Bernhard Eduard F emow: A Storey ofNorth American Forestry. Forest History Society, Durham, NC. USA. 623pp. Roth, F. 1905. What the state should do and why it should do it now. Report to the Michigan Forestry Commission 1903-4280—96. Roth, F. 1919. Communication from Mr. Filibert Roth, Professor of Forestry at University of Michigan, telling of his visit to the Forest Reserves. Pages 104-1 1 l in Biennial Report of the Public Domain Commission from July 1, 1916, to June 30, 1918. Fort Wayne Printing Co., Fort Wayne, IN, USA. 134pp. Rudolph, V.J., Associate Chairman, Dept of For, Mich State Univ. 1982. Unpublished letter to Jerry Morford, City Manager, Grayling, and Dr. Henry Webster, Chief, For Mgt Div, Mich Dept of Natl Res, dated 6 October. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Rudolph, V.J., Associate Chairman, and J .J . Hacker, Graduate Assistant, Dept of For, Mich State Univ. 1982. The Beal Plantation at Grayling: Alternatives for Its Future, dated 24 September. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Sandell, D. 1983. Beal Plantation. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Sargent, CS 1876. A few suggestions on tree-planting. Pages 7-29 in Massachusetts Society for Promoting Agriculture. Prizes for Arboriculture. Alfred Mudge & Son, Printers, Boston, MA, USA. 29pp. Sargent, CS 1878. Notes on Trees and Tree—Planting. Reprinted from the 25th Annual Report of the Secretary of the Board of Agriculture. Rand, Avery, & Co., Printers to the Commonwealth, Boston, MA, USA. 22pp. 148 Schafer, J.A., Easements-Exchanges, Lands Div, Mich Dept Natl Res. 1978. Unpublished letter to Jerry W. Morford, City Manager, Grayling, MI, dated 10 February. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Schafer, J .A., Easements-Exchanges, Lands Div, Mich Dept Natl Res. 1979. Unpublished letter to Jerry Morford, City Manager, Grayling, MI, dated 22 May. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing. MI, USA. Schaetzl, R.J., and D.G. Brown. 1996. Forest associations and soil drainage classes in presettlement Baraga County, Michigan. The Great Lakes Geographer 3(2):57-74. Schenck, CA. 1955. The Biltmore Story: Recollections of the Beginning of Forestry in the United States. American Forest History Foundation, Minnesota Historical Society, St. Paul. MN, USA. 224pp. Schnicke, G.T., District Fisheries Biologist. Mich Dept Natl Res. 1978. Unpublished memorandum to A.J. Coates, District Forest Supervisor, AuSable State Forest, Mich Dept Natl Res, dated 21 March. Subject: Proposed exchange - City of Grayling. Beal Plantation File, For Mgt Div, MI Dept Natl Res, Lansing, MI, USA. Schweingruber, EH. 1993. Trees and Wood in Dendrochronology: Morphological, Anatomical, and Tree-Ring Analytical Characteristics of Trees Frequently Used in Dendrochronology. Springer-Verlag, Berlin, BRD. 402pp. Simard, A.J., and R.W. Blank. 1982. Fire history ofa Michigan jack pine forest. Mich Acad 15:59-71. Sodders, B. 1997. Michigan on Fire. Thunder Bay Press, San Diego, CA, USA. 390pp. SPSS Science, Inc. 1998. Systat Version 9.01. Chicago, IL, USA. Spurr, SH. 1956. Plantation success in the Harvard Forest as related to planting site and cleaning, 1907-1947. J For 54:577-579. Stokes, M., and T. Smiley. 1968. An Introduction to Tree-Ring Dating. The University of Arizona Press, Tucson. AZ, USA. 73pp. Taft, LR. 1892. Report of the Horticulturist. Pages 177-180 in Twenty-First Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1892. Robert Smith & Co., Lansing, MI, USA. 607pp. Tarr, W.H., Area Forester, AuSable State Forest, Mich Dept Natl Res. 1978. Unpublished memorandum to A.J. Coates, District Forest Supervisor. AuSable State Forest, Mich Dept Natl Res, dated 11 April. Subject: Possible Exchange with the City of 149 Grayling. Beal Plantation File, For Mgt Div, MI Dept Natl Res, Lansing, MI, USA. Telewski, F.W. 1995. Wind-induced physiological and developmental responses in trees. Pages 237-263 in M.P. Coutts and J. Grace, editors. Wind and Trees. Cambridge University Press, Cambridge, UK. 485pp. Telewski, F.W. 1998. The beginning of an artificial forestry in mid-19th century Michigan: the contributions of W.J. Beal to silviculture. The Mich Bot 37(2):35- 58. Temple, SA. 1995. Biodiversity, landscape-scale management and the ecological importance of the pine barrens community. Page 2 in BA. Borgerding, G.A. Bartelt, and W.M. McCown, editors. The Future of Pine Barrens in Northwest Wisconsin: A Workshop Summary. Wisc Dept of Nat Res PUBL-RS-913-94, Madison, WI, USA. 105pp. Thiel, S, Unit Mgr, Mich Dept Natl Res, and D. Sikarskie, Coordinator, Huron Pines RC&D. 1996. Draft press release, dated 23 September. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Thomsen, G. 2001. Response to winter precipitation in ring-width chronologies of Pinus sylvestris L. from the northwestern Siberian Plain, Russia. Tree—Ring Research 57(1):]5-29. Tilman, D. 1988. Plant Strategies and the Structure and Dynamics of Plant Communities. Princeton Univ Press, Princeton, NJ, USA. 376pp. TMR. 1994. Unpublished manuscript by an unknown author, dated 3 March. Tomlinson, P.T., R.J. Jensen, and J .F. Hancock. 2000. Do whole tree silvic characteristics indicate hybridization in red oak (Quercus section Lobatae)? Am Midl Nat 143:154-168. Tucker, T.R., Chief, Lands Div, Mich Dept Natl Res. 1977a. Unpublished letter to Jerry W. Morford, City Manager, Grayling, MI, dated 10 February. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Tucker, T.R., Chief, Lands Div, Mich Dept Natl Res. 1977b. Unpublished letter to Jerry W. Morford, City Manager, Grayling, MI, dated 16 May. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Tucker, T.R., Chief, Lands Div, Mich Dept Natl Res. 1978a. Unpublished letter to Jerry W. Morford, City Manager, Grayling, MI, dated 23 January. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. 150 Tucker, T.R., Chief, Lands Div, Mich Dept Natl Res. 1978b. Unpublished letter to Jerry W. Morford, City Manager, Grayling, MI, dated 6 February. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Tucker, T.R., Chief, Lands Div, and H.H. Webster, Chief, For Mgt Div, Mich Dept Natl Res. 1978. Unpublished draft memo to CD. Harris, Chief, Bureau of Renewable Res Mgt, Mich Dept Natl Res. no date. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Tucker, T.R., Chief, Lands Div, For Mgt Div, Mich Dept Natl Res. 1980. Unpublished letter to Jerry Morford, City Manager, Grayling, MI, dated 7 January. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Tyndall, R.W., and J .C . Hull. 1999. Vegetation, flora, and plant physiological ecology of serpentine barrens of eastern North America. Pages 67-82 in RC. Anderson, J .S. Fralish, and J .M. Baskin, editors. Savannas, Barrens, and Rock Outcrop Plant Communities of North America. Cambridge Univ Press, New York, NY, USA. 470pp. United Nations. 1995. Santiago Declaration: Statement on criteria and indicators for the conservation and sustainable management of temperate and boreal forests. Issued in Santiago, Chile, on 3 February 1995. United Nations Division for Sustainable Development. 2000. Mandate of the Intergovernmental Panel on Forests. http://www.un.org/esa/sustdev/forestsmandate.htm Verheyen, K., and M. Hermy. 2001. The relative importance ofdispersal limitation of vascular plants in secondary forest succession in Muizen Forest, Belgium. J Ecol 89:829-840. Vora, RS 1993. Moquah Barrens: pine barrens restoration experiment initiated in Chequamegon National Forest. Rest Mgt Notes 11(1):39-44. Voss, E.G. 1972. Michigan Flora: Part I. Gymnosperms and Monocots. Bull. Cranbrook Inst. Sci. 55 and Univ Mich Herb, USA. 488pp. Voss, E.G. 1985. Michigan Flora: Part II. Dicots (Saururaceae-Cornaceae). Bull. Cranbrook Inst. Sci. 59 and Univ Mich Herb, USA. 724pp. Voss, E.G. 1996. Michigan Flora: Part III. Dicots (Pyrolaceae-Compositae). Bull Cranbrook Inst Sci 61 and Univ Mich Herb, USA. 622pp. Voss. E.G., and GE. Crow. 1976. Across Michigan by covered wagon: a botanical expedition in 1888. The Mich Botanist 15(1):3-70. 151 Walter, H., and H. Lieth. 1967. Klimadiagramm-Weltatlas (World Atlas of Climate Diagrams). G. Fischer Jena, Germany. Walters, M.B., and RB. Reich. 1999. Research review: low-light carbon balance and shade tolerance in the seedlings of woody plants: do winter deciduous and broad- leaved evergreen species differ? New Phytol 1431143-154. Webster, H.H., Chief, For Mgt Div, Mich Dept Natl Res. 1978. Unpublished letter to Wesley Manley, President, Mich Forest Assoc, dated 14 June. Beal Plantation File, For Mgt Div, Mich Dept Natl Res, Lansing, MI, USA. Werlein, J .O. 1998. Soil Survey of Crawford County, Michigan. US Dept of Agric, Nat Res Cons Serv, Washington, DC, USA. 274pp. Whitney, G.G. 1986. Relation of Michigan’s presettlement pine forests to substrate and disturbance history. Ecol 67(6):1548-1559. Whitney, G.G. 1987. An ecological history of the Great Lakes forests of Michigan. J Ecol 75:667-684. Wilde, SA., J .G. Iyer, C. Tanser, W.L. Trautmann, and KC. Watterston. 1965. Growth of Wisconsin coniferous plantations in relation to soils. Res Bull 262, Univ Wisc, Madison, WI, USA. 81pp. Willits, E. 1888. Report of the Director of the Experiment Station. Pages 138-146 in Twenty-Seventh Annual Report of the Secretary of the State Board of Agriculture of the State of Michigan, 1888. Thorp and Godfrey, State Printers and Binders, Lansing, MI, USA. 596pp. Woodward, F1 1987. Climate and Plant Distribution. Cambridge Univ Press. Cambridge, MA, USA. 174pp. Yoder, C.T., Regional Director, Mich Dept Natl Res. 1978a. Unpublished memorandum to C .D. Harris, Chief, Bureau of Renewable Res Mgt, Mich Dept Natl Res, dated 6 June. Subject: Land Exchange for City of Grayling. Beal Plantation F ile, For Mgt Div, MI Dept Natl Res. Lansing, MI, USA. Yoder, C.T., Regional Director, Mich Dept Natl Res. 1978b. Unpublished memorandum to C .D. Harris, Chief, Bureau of Renewable Res Mgt, Mich Dept Natl Res, dated 16 June. Subject: Land Exchange - City of Grayling. Beal Plantation File, For Mgt Div, MI Dept Natl Res, Lansing, MI, USA. Zhang, Q., K.S Pregitzer, and DD. Reed. 2000. Historical changes in the forests of the Luce District of the Upper Peninsula of Michigan. Am Midl Nat 143294-1 10.