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Historical Perspective c: Managing wildlife resources is largely a problem in land and water use. What can be done at present often is indicated by past events, insofar as these can be interpreted reliably. The coming of Caucasian man to North America vastly altered the complex of pristine plant-animal communities. It appears reasonable to assume that most such communi- ties represented a stage in a developmental series of types trending toward stability. The progression of changes was monitored by climate, conditioned by the substrate, and often modified or suspended by the influences of animals or aboriginal man. Although the dynamics of natural ecosystems are too little understood, biological investigations since 1920 have revealed relationships sufficiently consistent for guid- ance in many aspects of resource use. It is axiomatic that every currently living thing evolved as part of a working association of plants and animals (see Taylor, 19491. Thus, no species can now reproduce and survive in self-dependence outside such a community, each having specific relationships that limit the condi- tions under which it can exist. Whether it is broadly tolerant or highly specialized is likely to be reflected in its present distribution and how it has responded to alterations in environment. PRIMITIVE CONDITIONS IN NORTH AMERICA The record of faunal and floral events on this continent falls far short of a desirable level of detail, but major changes are fairly well known.

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2 Land Use and Wildlife Resources Pre-Columbian vegetation patterns probably approximated those mapped by Kuchler (1964) as "potential natural vegetation." Then, as now, any large area so characterized might encompass any or all stages in the sere. Testimony to the ultimate ascendancy of geological erosion is im- plicit in such fo~ations as the ancient stubs of the Appalachians, mas- sive river deltas, extensive loess deposits, and the slope of land from the foot of the Rockies eastward. Yet the stabilizing influence of vege- tation is also manifest, and not alone in the attainment of a self- perpetuating stage. At each step of succession there is a countering and buffering of the degrading action of water, wind, heat, cold, and animal life. In terms of the general thrift and coverage of vegetation, it is hardly to be doubted that in the late fifteenth century the land surface of this continent was undergoing a slower rate of change than at any time after the white man's enterprise became effective. Marsh (1864) clearly perceived these relationships more than a century ago. Aftermath of Glaciation As compared with much of the earth's surface, North America is largely a young land. The final phases of continental and alpine glacia- tion in mid-latitudes were only 8 to 10 thousand years in the past. After recession of the ice, further climatic perturbations induced ex- tensive biogeographic advances and retreats that left species and com- munity isolates as witness to their passage (Sears, 1948; Deevey, 1949; Smith, 19571. Of special significance was extension of the "prairie peninsula" eastward presumably during a dry period of the Hypsi- thermal Interval (Gleason, 1922; Transeau, 1935; Deevey and Flint, 1957), with grassland relicts remaining to mark the way, and even a heath-adapted prairie chicken established on the Atlantic coastal bar- rens (Gross, 19281. As pointed out by Sears (1942), some of the evi- dences of past xerothermic conditions may survive from interglacial, even Tertiary, climatic episodes. A major faunal occurrence of postglacial times was extinction of many species of North American large mammals-camels, llamas, horses, bison, muskoxen, pronghorns, stag-moose, peccaries, mammoths, mast- odons, ground sloths, giant beavers, and certain carnivores (Flint, 1957; Martin, 1958-19671. The disappearance of this megafauna has not been satisfactorily explained, but it was in progress during the period of roughly 12,000 to 5,000 B.C., when early men were numerous and active. Well-developed hunting cultures were widely distributed across

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Historical Perspective the continent (Wormington, 1962; Muller-Beck, 1966) and finely crafted clovis, Folsom, and subsequent types of projectile points were being used against mammoths, bison, and lesser beasts. Relative to further events, Martin (1963) surmised that 3 Following extinction of the large mammals the early hunters probably suffered economic depression and a population crash. Under a climate similar to the present and with the existing biotic zones in place, the early hunters were obliged to begin their 7,000-year experiment with native plants, leading in the altithermal to in- creasingly skillful techniques of harvesting and gathering, to the domestication of certain weedy camp-followers, and, within the last 1,000 years, to the widespread adoption of flood plain agriculture. Many clues along the trail remain to be de- tected by pollen analysis and other paleoecological methods. Certain it is that the beginnings of intensified land use in the New World interrupted long-term biotic adjustments. In no sense were the natural changes complete, nor was the situation static. The lag in such processes being what it is, there is reason to suppose that the mercurial North American climate was shifting sufficiently often to maintain a state of flux in associations of living things. Nonetheless, bioclimatic types were recognizable for large areas, which, under various defini- tions, have been described as life zones (Merriam, 1898), blames (Clements and Shelford, 1939), biotic associations or provinces (Vestal, 1914; Dice, 1943), biochores (Dansereau, 1957), or plant formations (Holdridge, 19479. Questions of te'~inology need not confuse the present discussion. That plant-animal associations commonly exhibit a "continuum" of variation from one locality to another is inherent in the distribution and complexity of environmental factors. Disturbance Communities As noted in the foregoing, conditions of the immediate prehistoric period were largely an expression of postglacial vegetation successions induced by climatic change. Associated with, or added to, this predis- posing influence were disturbance factors responsible for recurring cycles of developmental plant-animal communities. Weather extremes- high winds, torrential rains, floods, drought, exceptional heat or cold- often brought about drastic localized modifications of the biota, espe- cially through the setting back of vegetation to early seral stages. That animal and human activity produces similar effects is well known. Any review of habitat requirements in birds and mammals attests that relatively few species are characteristic of climax vegetation (see

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4 Land Use and W ildl if e R esou roes Shelford, 19631. Certain rodents are capable of living in fully developed grassland or tundra-e.g., voles and lemmings-or in terminal forest types. Among forest species, the white-footed mouse and eastern gray squirrel may be mentioned. A few birds (woodpeckers, titmouse, chickadee) are present the year round in stabilized forests such as beech-maple or oak-hickory. Of all North American hoofed animals, probably the barren-ground caribou can most logically be considered a climax species (Leopold and Darling, 1953: 54), although it wanders over a variety of vegetation types, and especially into the sparse boreal forests for wintering. Most of the wildlife species of major interest to man depend on a stage of vegetation below the climax, or they make use of several suc- cessional stages, or they migrate from one vegetation zone to another.* This undoubtedly was an important key to distribution in primitive times and to great changes that have taken place in recent centuries. To generalize on a widely varied relationship, it is true that many rodents and lagomorphs, both seed- and leaf-eaters, are largely depen- dent on pioneer herbaceous types of vegetation. Particularly in grass- lands, they respond to disturbance factors that induce the spread of fortes and annual grasses (see review by Bond, 1945), and by their own burrowing and other activities they may help to maintain such condi- tions, the prairie dog being an outstanding example (Koford, 19583. That there are also more subtle, long-term, cumulative effects impor- tant in the development of soils-loosening, mixing, aeration, weather- ing-is hardly to be doubted (Grinnell, 19231. In each forested region of the continent there are numerous species of thicket-inhabiting or edge-dwelling birds and mammals. They spread and increase whenever the woodland canopy is opened and a subsere initiated. They depend on a transitory condition, and they will disap- pear with the habitat in the course of successional change. Some of the game birds well exemplify this situation. In forested areas, nearly every kind of grouse and the various races of wild turkey require that a part of the range be in herbaceous vegetation-grassy and weedy openings where the young can sun and dust and obtain their "starter" diet of insects (cf. Edminster, 19544. The brushy fringes provide many kinds of fleshy fruits, and different vegetation types may fulfill other essen *The term "climax" will be used most often in this report to designate a terminal stage in the development of vegetation under given climatic conditions. This use recognizes that in some areas such a stage may not be clear and agreed-upon, and no universally accepted definition of climax, especially as applied to ecosystems, is available (see Tansley, 1935; Clements, 1936; Whittaker, 1953).

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Historical Perspective tial functions in the life history. A similar dependence is seen in many small mammals, including mice and ground squirrels, the snowshoe hare of boreal forests, and southward many races of the cottontail. Among larger species, the deer, elk, and moose are in part browsers upon the brush ranges of secondary successions. 5 Modifications of vegetation have far-reaching effects in the life com- munity. Responses occur from step to step in the energy-exchange linkage originating in the primary production of green plants. When key food supplies-such as ground-cover~ng types of herbage-flourish, the lower consumer levels (prey animals) are nourished and increased. These, through their augmented numbers, pass on to the carnivores the benefits of favorable change. Through this universal mechanism the herbivores exert a primary control on their dependent predators. When basic conditions are altered, at least for a time, the community may take on a new aspect arid composition. Animal Influences In every native ecosystem the animal component has played its part in annual and long-term dynamics. Some of the effects on vegetation and site were widespread and conspicuous, although relatively few can be described reliably. On the vast majority of wooded watersheds, innumerable beaver dams were constantly renewed to produce a cycle of deadenings, portals, meadows, and early-growth forests. The floodings and succeeding stages became habitat for waterfowl and other aquatic life, edge-inhabiting birds and mammals, and, in the North, moose and woodland caribou. The extent of wetlands in primitive times is not likely to be fully appreciated by those accustomed only to today's scenery. Although most early observers simply took for granted what they saw, a few left valuable descriptions of conditions that were changing rapidly even a century ago. Hubbard (1887, 362 et seq.), first state geologist of Michi- gan, described the lowlands between Lake Erie and Saginaw Bay as one of the great beaver-trapping grounds. To a great extent level, it is intersected by numerous water courses, which have but moderate flow. At the head-waters and small inlets of these streams the beaver established his colonies. Here he dammed the streams setting back the water over the flat lands, and creating ponds, in which were his habitations.... The trees were killed, the land converted into a chain of ponds and marshes, with interven- ing dry ridges. In time, by nature's recuperative process-the annual growth and decay of grasses and aquatic plants-these filled with muck or peat, with occasional

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6 Land Use and Wildlife Resources deposits of bog lime, and the ponds and swales became dry again.... In a semi- circle of twelve miles around Detroit, having the river for base, and embracing about 100,000 acres, fully one-fifth part consists of marshy tracts or prairies tsic] which had their origin in the work of the beaver. A little further west, nearly one whole township, in Wayne County, is of this character. Although the total ecological influence of beavers in aboriginal North America probably cannot be estimated today, it must have been great for a creature that ranged from south of the Rio Grande to Hudson Bay and from western Alaska to Labrador. According to Seton (1929, vol. IV, pt. 2, p. 446), the only major areas in the United States lacking beaver were the coastal region of the Southeast, including Florida, and the largely waterless deserts of the Southwest. He states that the early beaver population of the Adirondacks was estimated at 1 million, or about 60 to the square mile, and that a more recent esti- mate for Algonquin Park in Ontario was 50 per square mile. His ex- trapolation of a minimum population of 60 million in North America appears conservative. The extensive dewatering of the land surface that has resulted from killing off of beaver, deforestation, drainage, and natural and acceler- ated filling is one of the most significant changes affecting wildlife since the discovery of America. Disappearance of springs, siltation of stream- beds, and drying up of watercourses in summer have been especially well documented in Wisconsin (Scott and Hoveland, 19511. In degree, such changes were to be expected, but the extremes of conditions often are associated with overgrazing and destructive methods of agriculture, which will be terminated inevitably, either because they destroy pro- ductivity or as a result of land-use improvements. In forested areas a part of recent trends toward restoration of a more natural hydrology is the countrywide re-establishment and management of beavers. The bison provides another example. There is little doubt that this animal was an important influence in determining the floral composi- tion of the Great Plains.* Trailing and dust wallowing provided the bare soil conditions conducive to early successions of fortes and certain grasses (see Roe, 1951: 101; Weaver and Albertson, 1956: 104, 3701. More especially, the extensive coverage of short grasses on the high plains was largely a result of sustained grazing by the bands and herds that shifted about in response to forage conditions. This activity fa ~Here we have an example of the problems encountered in strictly defining climax. Said Shelford (1931): ". . . if the bison held some of the mixed prairie in a short grass stage, then short grass is the bioecological climax and its proper bioecological designation is Bouteloua- Bison, even though the climax with bison excluded is something different."

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Historical Perspective vored the more xeric buffalograss (Buchloe dactyloides) and gramas (Bouteloua) over the mid-grasses, which Weaver and Albertson (1956) regard as part of the "mixed prairie" climax of the high plains. Larson (1940) reviewed the early findings on this situation and discussed its conceptual semantics. Relative to the number of bison and associated species, he remarked: A buffalo requires about the same amount and kind of feed as a cow, so it is evi dent that if Seton's estimate of 20 million or Clements' and Shelford's estimate of 30 million buffalo in early days on the plains is anywhere near correct these ani- mals were sufficiently numerous, along with the 4 to 8 million antelope and exten- sive herds tsic] of elk, deer, and other wild animals, to hold the drier portions of the plains in a short grass stage. 7 It is probable that another creature, largely restricted to the eastern deciduous forest region, had important and widespread effects on vege- tation and associated animal life in pre-Columbian times. That the pas- senger pigeon was inconceivably abundant is attested by all original descriptions. After a review of various estimates, Schorger ( 1 955) "guessed" that there were three billion, and possibly five billion, in the primitive continental population. The clue to ecological impacts of the pigeon lay in its association in flocks numbering many millions. In such masses it nested, fed, roosted, and migrated. The great nestings in the Lake States region covered thousands of acres of forest, loading some trees to the breaking point. A nesting in Benzie County, Michigan, in 1874 was described as "50 square miles of pigeons" (Hubbard, 1887: 3089. The results of such a concentration were similar to those described by Audubon (quoted from Ornith ological Biography, 1831-1 839, by Mershon, 1 9 07: 3 3 for one of the nighttime roosts in Kentucky: The dung lay several inches deep, covering the whole extent of the roosting-place, like a bed of snow. Many trees two feet in diameter, I observed, were broken off at no great distance from the ground; and the branches of many of the largest and tallest had given way, as if the forest had been swept by a tornado.... The Pigeons, arriving by thousands, alighted everywhere, one above another, until solid masses as large as hogsheads were formed on the branches all around. Here and there the perches gave way under the weight with a crash, and, falling to the ground, de- stroyed hundreds of the birds beneath, . . . Whether on the wintering grounds of southern states or nesting in the North, the great flocks required huge quantities of food within flying distance, which might be 50 miles (Schorger, 19551. A wide

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8 Land Use and Wildlife Resources variety of tree fruits and berries were taken, but mainstays were the large mast crops of the forest, especially beech, oak, and chestnut. A visitation by millions of pigeons took the entire harvest and left little for squirrels, bears, deer, raccoons, and other wilderness mast- feeders. Ecological effects of the passenger pigeon were drastic and often lasting (especially forest damage and overfertilization), but they were localized. The pigeons were sustained by the environment because of the erratic migration habits of the species. One-time (or perhaps periodic) concentrations occurred in a given area in response to tem- porarily favorable conditions (especially large food crops). Then the birds passed on, not to return in force for some years, during which the range recovered. Similar adaptations are seen in other gregarious species, notably the migratory barren-ground caribou in its use of the slow-growing lichen ranges of the Arctic (Darling, 1 95 61. A recognition of some of the more obvious animal influences on the North American environment in no way implies that they were un- usual. In every natural community such relationships are prevalent, whether or not they are a part of our biological insight. F. Ire Of all the factors that disturb the earth's vegetation, fire is undoubt- edly the one that has the greatest effect. It is the inevitable concomi- tant of xeric conditions, and hence is an important means by which plant communities are determined in dry climates. It is evident that the nature of soils is closely tied in with these processes, in terms of both soil development and the predisposition of sites to burning. Shantz ( 1954) noted that "The grasslands are of two types: (A) Those determined largely by climatic conditions and on pedocal soils, un- leached and with dry subsoils, and (B) those which have replaced forests destroyed by cutting or by fire or both, or maintained against forest development by conditions favoring fire." The American tall-grass "prairies" were an outstanding example of the latter type-a disturbance climax (disclimax of Weaver and Clements, 1938: 86 et seq.), or fire subclimax. After the introduction of heavy grazing by domestic livestock and the breaking of the prairies for faming, the extensive fires of early days were no longer possible. In the North an immediate result was widespread overgrowth of un- cultivated tall-grass habitats by woodland (Gleason, 1913; McComb and Loomis, 1944; Curtis, 19599. In southern grasslands an evident

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Historical Perspective trend was the thickening and spread of the mesquite savanna and other shrub types (Jackson, 1965; Lehmann, 1965; Box, 19679. Recognition of the close relationship of fire dynamics to many types of grassland is now quite general; Sauer (1950) goes so far as to say, "Suppression of fire results in gradual recolonization by woody species in every grassland known to me. I know of no basis for a climatic grassland climax, but only of a fire grass 'climax' for soils permitting deep rooting." In all regions of the United States where pines occur, fire plays a major role in the ecology of many species of Pinus, as well as certain other conifers (Hanson, 1 939; Garren, 1 943; Little, 1 953; Biswell, 19631. The practical utility of such knowledge in producing timber crops is evidenced by the fact that on national forests of the southern region in 1963 the U.S. Forest Service carried out prescribed forest management burning on 3 16,65 8 acres. In Florida burning can be con- trolled on some half million acres, of which an average of 83,000 acres a year are so treated (Riebold, 19641. Through eons of evolution, fire has played its part in the speciation of many kinds of plants and animals and in the development of biotic communities. As Komarek remarked (19641: The antiquity of fire seems apparent in that the most ancient of tree families, such as the conifers, and the apparently oldest genera of grasses, such as Aristida, Stipa, Andropogon, etc., have the greatest concentration of those genes responsible for resistance and adjustment to a "fire environment." In fact, it appears that during long periods of time fire type communities of plants and animals have covered vast areas of the earth's land surface. While this factor must have been ever-present in dry climates, it is likewise influential in areas of light soil and other xeric sites in temperate and relatively moist climates. For fires to be effective in shaping the composition and aspection of vegetation, periodic dry summers may be sufficient to inhibit the growth of thicket-forming deciduous woody plants and to promote the spread of fortes, grasses, and fire-resistant trees, many of which are characteristically thick-barked. The ecological effects of fire have been categorized by Hanson (1939) as follows: 1. Burning causes partial or complete destruction of plant and animal life, as well as dead vegetation cover. 2. Burning causes modification of atmospheric factors; light, wind, precipitation (rainfall interception).

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10 Land Use and Wildlife Resources 3. Direct effect of the temperature of the fire, upon the soil, as organic and mineral contents, structure, texture, animal and plant life in the soil. 4. Effect of destruction of plant cover upon consequent soil moisture, wind and water erosion, mineral content, biotic processes in soil, pH, etc. 5. Effect of fire upon consequent plant and animal succession, relation of fire to aiding establishment of early invaders as Lodgepole pine. 6. Effect of subterranean fires upon rock formation and topography. Lightning strikes, especially on trees or stubs, undoubtedly were a common natural cause of fires. Hanson cites sources indicating that lightning caused 41 percent of fires on and adjacent to California forests (obviously in a period when fires of human origin also were prevalent) and about 8 percent of forest fires in the nation as a whole. In his re- vealing study of "the natural history of lightning," Komarek (1964) demonstrated that this factor is so effective and widespread that the presence of burnable material is the only requisite for producing fires frequently enough "to have lasting effect on plant and animal com- munities." There can be little doubt that these effects did accumulate to a highly significant degree over large areas before the coming of men to this continent. An appraisal of early human influences on the biota of North America must necessarily give further emphasis to the subject of fire. While the Paleo-Indians may well have had a hand directly in the disappearance of large mammals, their most far-reaching cultural impacts undoubtedly were exerted through habitat change. The great significance of burning by primitive man was described by Martin (19581: For the ecologist and biogeographer one point remains clear. From the time of man's arrival we may assume a radical change in fire frequency. In the strict sense, theoretical climatic climax vegetation in savanna and grassland areas . . . and even in parts of the Eastern Deciduous Forest, cannot postdate man's arrival. In addi- tion to savannas many areas of temperate forest may have been greatly modified and subclimax, consolidation, or even pioneer species favored at the expense of those typical only of climax positions in plant succession. The antiquity of the human species in North America needs no par- ticular discussion here (see summary by Griffin, 19671. Firm evidence supports the presence of man on this continent for the last 15,000 years, a period that began more recently than the time of maximum Wisconsin glaciation, which was about 18,000 years ago. Thus, many millenia passed during which plant-animal communities reached dy- namic adjustment to intensified regimes of burning as a result of the human factor.

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Historical Perspective 11 In an incomplete review of this subject, Stewart ( 1951 ) found "more than 200 references to Indians setting fire to vegetation in aboriginal times, and these references cover all major geographic and cultural areas." Wherever plant cover would burn, it was burned repeatedly as a part of the cultural way of life. Some of the beneficial effects probably were evident. Fires opened up thick growth where game might be hunted more easily. They brought about early green-up and improved grazing on the prairie, making areas near villages more attractive to buffalo. Fires also were the means of producing large berry crops in some situa- tions, and they helped abate the mosquito nuisance in others. Scientific study now shows that after fires certain kinds of vegetation are more succulent and palatable, and higher in protein content, providing im- mediate benefits to browsing and grazing animals (Aldous, 1934; Lay, 1957). Basic changes brought about by repeated burning and resulting in the establishment and maintenance of fire disclimax communities have been described by Sauer (19561: Pyrophytes include woody monocotyledons, such as palms, which do not depend on a vulnerable cambium tissue, trees insulated by thick corky bark, trees and shrubs able to reproduce by sprouting, and plants with thick, hard-shelled seeds aided in germination by heat. Loss of organic matter on and in the soil may shift advantage to forms that germinate well in mineral soils, as the numerous conifers. Precocity is advantageous. The assemblages consequent upon fires are usually characterized by a reduced number of species, even by the dominance of few and single species. Minor elements in a natural flora, originally mainly confined to acci- dentally disturbed and exposed situations, such as windfalls and eroding slopes, have opened to them by recurrent burning the chance to spread and multiply. In most cases the shift is from mesophytic to less exacting, more xeric, forms, to those that do not require ample soil moisture and can tolerate at all times full ex- posure to the sun. In the long run the scales are tipped against the great, slowly maturing plants-the trees (a park land of mature trees may be the last stand of what was a complete woodland). Our eastern woodlands, at the time of white settlement, seem largely to have been in process of change to park lands. As noted previously, changes of this nature over large areas had given character to the vegetation and animal life long before the advent of man on the American scene. The human influence was one of intensifi- cation. At all levels of culture man depends largely on early ecological successions. With the passage of time he not only adapted to regional habitats of the continent; he also made large-scale environmental changes that, at least in some cases, helped to serve his needs.

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18 Land Use and Wildlife Resources and Indians, and population buildup westward was rapid. At that time more than 70 percent of Americans were living in rural communities. This percentage declined to 54.2 in 1910 and 48.6 in 1920, as the popu- lation passed 105 million (Edwards, 19401. Changes on the Land Under many types of grants and incentives, the private occupation, breaking, and clearing of huge areas had gone forward (Edwards, 19401: It was practically Impossible to have foretold in 1860 that within 30 years a half billion acres of the public domain would have been disposed of or reserved for governmental purposes. The land was considered valueless unless it was put under cultivation as rapidly as possible, and the various land acts did help to people the wilderness. To have opposed unregulated settlement would have been considered either a mad perversion or a reflection of selfish economic interest. This was the period of heedless wiping out of wildlife and wastage of forest resources that gave rise to new and more conservative trends of thinking. The conservation movement became recognizable during the Cleveland administration when land policies were revised (1891) to abate flagrant abuses and Congress first authorized the reservation of forested areas of the public domain for public purposes. The idea of using resources "for the greatest good of the greatest number" received its greatest stimulus under Pinchot and Roosevelt. The industrialization, mechanization, and specialization of American agriculture came rapidly after the Civil War, and the permanent land- use pattern was well established by the turn of the century. Potentially, nearly a third of the land area of the 48 contiguous states (about 600 million acres) is suited to some kind of agricultural cropping, but only about 400 million acres is actually so used (Barnes and Marschner, 19581. The widespread conversion of primitive habitats has posed the great problems, as well as many of the opportunities, in wildlife management. Lands primarily adapted to forestry have likewise undergone exten- sive change. Abounding abuses in the timberlands of the nation received initial federal recognition in 1876 with the appointment in the Depart- ment of Agriculture of an agent to study forest problems. This step led to creation of the Division of Forestry in 1881 and the Forest Service in 1905. By the end of the Roosevelt administration in 1908, some 148 million acres had been reserved for national watershed and timber pro- tection purposes. By that time the great bulk of virgin forests in the East and Midwest had been cut, and loggers were moving into the South.

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Historical Perspective 19 Vast areas had been converted to fire-ravaged barrens, scrub timber, and brush growth. As state and federal forestry programs expanded and became more professional, the cutovers were brought under protection and management. New crops of timber began to grow. Effects on Wildlife By the early 1 900's, faunal conditions in much of North America bore little resemblance to the communities of early times. What may be termed "wilderness wildlife" had been greatly reduced. The panther, wolf, elk, and wild turkey were largely gone from the eastern United States, and the grizzly, antelope, and mountain sheep were disappearing in the West. Deer, beaver, black bear, and many lesser species had be- come locally or regionally scarce or absent. In part, this was due to lack of adequate protection, as well as to widespread changes in habitat. In contrast, some species of birds and mammals obviously throve on the land-use changes and came to be known as farm wildlife. Bobwhite quail, cottontail rabbits, and the edge-inhabiting fox squirrel spread widely in the region formerly occupied by extensive hardwood forests. In the rail-fence era of generalized farming, they became the wildlife symbols of rural America. Other familiar creatures, including the wood- chuck, prairie deermouse, field sparrow, meadowlark, and killdeer, ex- tended their ranges into regions of new croptields. Over the Midwest, the raven, gray squirrel (especially the black phase), and gray fox were reduced in number with the forests, while the crow, fox squirrel, and red fox increased with the openings. Commonly it was said that one creature "drove out" another. There was a general extension northward of the range of the opossum. Effective control of forest fires became a reality in the second and third decades of the century. During this period many states were establishing agencies to administer and manage their woodlands, waters, and wildlife. In 1918 responsibility for migratory birds was assumed by the federal government, and resident wildlife came under increasingly efficient regulation by the states. As new brush-stage forests developed, whitetail deer spread northward into the Lake States and Canadian border country where moose and caribou had been the principal big game. Deer herds reached their maximum in Pennsylvania and Michi- gan during the thirties, and a decade later it was evident that western mule-deer and blacktails were on the increase. Upland game birds were prominent in reflecting the changes in vege- tation. When forests of northern Wisconsin and Michigan gave way to grassy burns and brushlands, the prairie chicken and sharptailed grouse spread eastward beyond Sault Ste. Marie and the Straits of Mackinac

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20 Land Use and Wildlife Resources (Schorger, 1944; Ammann, 19571. Conversely, by midcentury, with the planting of openings and regrowth of timber stands, these prairie grouse were on the decline in Michigan's Upper Peninsula. From coast to coast across the North, and southward in both eastern and western mountains, ruffed grouse spread and multiplied in the vigorous growth of cutovers, old fields, and edges. Western blue grouse responded similarly. In the central hardwoods of the East, restocking of ruffed grouse and eastern turkeys (and also deer) helped to re-establish these species where they were wiped out early in the century. In the Southwest, the Merriam and Rio Grande turkeys returned to brushlands where they had long been absent. The development of early-stage forests was an extremely favorable condition for beavers, and they were widely restored in the thirties and forties-to the point of becoming a nuisance as they moved into agri- cultural areas. Largely after 1940, a general increase and spread of car- nivores occurred in the eastern United States. Black bears became abun- dant from the Deep South to Canada. Coyotes extended their range through woodlands and semiwild country to the east coast, and rac- coons and foxes increased in numbers beyond anything previously observed. The extent to which this carnivore prosperity may be attri- buted to habitat changes, as against long-term trends of obscure origin, is by no means certain. These discussions have centered on striking examples of faunal change following the extensive alteration of North America as a wild- life environment. In actuality, practically the entire biota of the con- tinent was affected and is being affected further as human populations build up. A relatively recent factor in the situation is the conscious effort toward wildlife habitat management that has developed since the mid-thirties. LAND-USE PLANNING The "redoing" of the American landscape has largely been a history of opportunism and short-term self-interest on the part of individuals and short-term expediency on the part of government. No doubt, extensive trial and error alone would eventually bring about a durable and pro- ductive use pattern for the various regions. This was the manner in which early crop specialization developed. But errors are costly, and obviously a better approach utilizes the application of science and technology. Edwards (1940) noted that the "agricultural revolution" was accompanied by

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Historical Perspective a quickening of the tendency for certain agricultural crops and commodities to dominate in the regions naturally suited to their production. In its Yearbooks for 1921 through 1925 the Department of Agriculture provided a notable series of articles which include historical descriptions, both textual and graphic, of the westward movement and current location of the agricultural crops and products of the United States. 21 Our commonest miscalculation has always been to overuse the land, especially during intervals of exceptionally favorable weather. Even the disastrous drought years of the thirties did not result in complete ad- justment of cropping demands to soil-climate limitations Chepil (1957) described the situation 20 years later: Large acreages suited only to permanent grass or forests are still devoted to culti- vated crops. In the Great Plains alone about 14 million acres not suited for perma- nent cultivation were cultivated in 1955. Much of this land offers low returns and is subject to severe erosion even in average years. The overuse momentum of the last century was widely evident as farmers attempted to cultivate areas cleared by lumbering and fires. The most knowledgeable agriculturists recognized the quality difference between "pine land" and "hardwood land." But pioneer settlers com- monly had little familiarity with the soil they wished to till, and their only sources of guidance were the representations of speculators or state land agents. In the three upper Lake States-Michigan, Wisconsin, and Minnesota-more than 38 million acres of the public domain were granted to the states (Lee and Wooten, 1958), and "these States quickly disposed of the land in order to get settlers on the land and to get cash for public improvements." Many a homestead was hewn out of the forest edge where, in a later generation, crumbling buildings would bear witness to the tragedy of misled optimism. Through tax reversions, "The Lake States began to see much land moving back into State ownership in the 1 920's. They suddenly became the unwilling owners of millions of acres of cutover land and abandoned farms because the private owners could no longer make a living . . ." (Lee and Wooten, 19581. The almost universal tendency toward overgrazing by the world's nonnomadic pastoralists has been discussed by Darling (19561. The market-grazing economy on western ranges is a convincing example. When, in 1936, Secretary of Agriculture Wallace sent to the President a report on the first comprehensive study of public and private grazing lands (U.S. Forest Service, 1936), he commented that "the general public knows less of the range resource, and as a result has been and is less concerned about its condition and conservation. than any of our

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22 Land Use and Wildlife Resources other important natural resources." The report noted that nearly 40 percent of the land area of the 48 states is available for livestock graz- ing, about half being privately owned. Through neglect and overstock- ing, "Range depletion on the public domain and grazing districts aver- ages 67 percent, on private, Indian, and State and county lands about half, and on national forests about 30 percent." Passage of the Taylor Grazing Act in 1934, followed by this historic Forest Service report, initiated a new era in which better practices gradually became established. As the errors of guesswork became manifest and studies of soil and its cover made headway, it became clear that farsighted government action would be necessary to avoid the continuation of economic and social disruption through unwise land use. In the ultimate resolution of factors, a durable husbandry must demand only the kind and quantity of products that can be yielded in perpetuity. As Shantz (1954) stated the case: Ecology should establish the proper balance of priorities of production. This planning should be based on the natural resource of climate, topography, soil, plant cover, animal population, and finally on the needs of the dependent human society.... Few lands can be given over to a single crop or to local or temporary needs. In many states the need was recognized for determining the produc- tion potential of lands and the reorientation of uses. Land economics surveys were carried out and in some cases zoning regulations were adopted (see Smith, 19261. By 1948, 34 states had enabling legislation on zoning, and in eight states there were counties with specific rural zoning restrictions. Among leading states in this field, Wisconsin used zoning ordinances in combination with the State Forest Crop Law to close some 5 million acres of forest land to agricultural development (Ruess et al., 19481. The limited-use capability of certain sites and soils was the basis for allocating them to extensive, as opposed to intensive, cropping. On new land-utilization maps these submarginal tracts were assigned to forestry and recreation. Many such lands would be incor- porated into fish and wildlife restoration areas, especially under pro- grams spurred by the availability of federal aid funds provided in acts of 1937 and 1950. Largely in response to destructive cutting, forest fires, and the tax reversion process, New York's justly famous Forest Preserve was established by an act of 1885, the provisions of which became a part of the state constitution (Lee and Wooten, 19581. The Adirondack and

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Historical Perspective 23 Catskill Forest Preserve, nearly 2.5 million acres, was placed in protec- tive custody of the state for public uses and ". . . shall be forever kept as wild forest lands. They shall not be leased, sold, or exchanged, or be taken by any corporation, public or private, nor shall the timber thereon be sold, removed or destroyed...." The primary benefits from this area were to be watershed protection and recreation-certainly a conservative appraisal of land-use capability. As noted by Marsh and Gibbons ( 1940), millions of acres of former cropland were so extensively damaged by erosion as to be submarginal for agriculture. "On much of this the forest is creeping back. Some day these lands may again be needed for cultivation; meanwhile, there may be no better or cheaper means for rebuilding them than restoration to forests." The long-term hazards of grain farming on what had been the central grasslands were evident in the dry thirties, and Thornthwaite (1941) summarized concepts strongly conditioned by the immediate past: In a semiarid climate like that of the Great Plains, wide climatic fluctuations are to be expected. Although it is not yet possible to forecast a specific drought year, it is possible to determine drought frequency and the probability of its occurrence. A stable economy can be achieved only if agriculture is adapted to the entire range of climatic conditions. This would necessitate returning to a grazing economy, in which pasturing of cattle on the natural and restored range is supplemented by the production of forage and feed crops in areas where flood irrigation is possible, and elsewhere in the rainy years on soils that are resistant to deterioration by wind and water. Such a change in land use requires an increase in the size of farms to a point where cultivation and grazing can both be controlled. It is well known that the most extensive and thorough application of land-use capability principles has been by the Soil Conservation Service. At the inception of this program in 1933, it was evident that there must be a systematic and objective basis for appraising sites of different potential on the same cropping unit. This realization led to the development of eight capability classes (Hockensmith and Steele, 1943) by means of which a land-use plan of the farm could be drawn. The significance of this approach in the management of wildlife is dis- cussed in Chapter 4. It is inherent in such a system that the most intensive management a site can sustain (for a yield of plant and animal products) should be determined scientifically on a basis of soil, topography, and climate. The individual operator cannot do this for himself, so it becomes a function of public programs where, logically, ultimate public welfare is

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24 Land Use and Wildlife Resources an important frame of reference. In the final analysis, considerations of demand and marketability are involved, and the question of what an area can produce becomes one of what it should produce. In this con- nection the influences of a growing population and a changing technol- ogy pose problems with which we shall be particularly concerned in chapters to follow. BIOLOGICAL INTERPRETATION OF HISTORY The history of our land and wildlife, interpreted in the context of modern ecological knowledge, is a study of "management experiments" on a vast scale. Unfortunately, there were no controls, and often the records are poor. Conclusions must be regarded in some degree as provisional. There can be no doubt, however, concerning the key role played by vegetation in determining the nature of animal communities and the density of populations. We may accept it as elementary in wildlife management that a desired species must be produced by first under- standing its place in the succession of communities and then finding the means of renewing, through effective and practicable kinds of "disturbance," the conditions in which it lives. Every environmental change must be viewed as a balance of values, for the processes that wipe out one organism may well benefit another. As Marsh (1864) said, ". . . in the husbandry of Nature there are no fallows." Wildlife management of the past was extensive and largely fortui- tous. Now, and in time to come, for the service of vastly more people, it must be intensive and purposeful. Opportunities for the kinds of recreation that depend on privacy are being diminished at the source. Somewhere a balance of values must be struck as we fashion our way of life for the future. In this endeavor it is a major asset to have a correct appraisal of what has happened in the past. REFERENCES Aldous, A. E. 1934. Effects of burning on Kansas bluestem pastures. Kan. Agr. Exp. Sta. Tech. Bull. 38. Ammann, G. A. 1957. The prairie grouse of Michigan. Mich. Dep. Conserv. Game Div. 200 p. Ashburn, P. M. 1947. The ranks of death: a medical history of the conquest of America. Coward-McCann, New York. 298 p.

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Historical Perspective Barnes, C. P., and F. I. Marschner. 1958. Our wealth of land resources. p. 10-18, In Land. The yearbook of agriculture 1958. U.S. Department of Agriculture. U.S. Government Printing Office, Washington, D.C. Biswell, H. H. 1963. Research in wildland fire ecology in California. Tall Timbers Fire Ecol. Conf. Proc., 2d Annul Conf. Bond, R. M. 1945. Range rodents and plant succession. 10th N. Amer. Wildl. Conf. Trans. 25 Box, T. W. 1967. Brush, fire, and west Texas rangeland. Tall Timbers Fire Ecol. Conf. Proc., 6th Annul Conf. Burroughs, R. D. 1961. The natural history of the Lewis and Clark Expedition. Michigan State University Press, East Lansing. Carrier, L. 1923. The beginnings of agriculture in America. McGraw-Hill Book Co., New York. 323 p. Chepil, W. S. 1957. Erosion of soil by wind, p. 308-314. In Soil. The yearbook of agriculture 1957. U.S. Department of Agriculture. U.S. Government Printing Office, Washington, D.C. Chittenden, H. M. 1935. The American fur trade of the far west. Barnes & Noble, New York. 2 vol. Clements, F. E. 1936. Nature and structure of the climax. J. Ecol. 24:252-284. Clements, F. E., and V. E. Shelford. 1939. Bio-ecology. John Wiley & Sons, New York. 425 p. Curtis, J. T. 1959. The vegetation of Wisconsin. University of Wisconsin Press, Madison. 657 p. Dansereau, P. 1957. Biogeography: an ecological perspective. Ronald Press, New York. 394 p. Darling, F. F. 1956. Man's ecological dominance through domesticated animals on wild lands, p. 778-787. In W. L. Thomas (ed.) Man's role in changing the face of the earth. University of Chicago Press, Chicago. Day, G. M. 1953. The Indian as an ecological factor in the northeastern forest. Ecology 34(2):329-346. Deevey, E. S. 1949. Biogeography of the Pleistocene (Part 1: Europe and North America). Geol. Soc. Amer. Bull. 60: 1315-1416. Deevey, E. S., and R. F. Flint. 1957. Postglacial hypsithermal interval. Science 125: 182-184. Dice, L. R. 1943. The biotic provinces of North America. University of Michigan Press, Ann Arbor. 78 p. Edminster, F-. C. 1954. American game birds. C. Scribuer's Sons, New York. 590 p. Edwards, E. E. 1940. American agriculture-the first 300 years, p. 171-27~. In Farmers in a changing world. The yearbook of agriculture 1940. U.S. Depart- ment of Agriculture. U.S. Government Printing Office, Washington, D.C. Flint, R. F. 1957. Glacial and Pleistocene geology. John Wiley & Sons, New York. 553 p. Garren, K. H. 1943. Effects of fire on the vegetation of the southeastern U.S. Bot. Rev.9:617-654. Gleason, H. A. 1913. Relation of forest distribution and prairie fires in the middle west. Torreya 13: 173-181 . Gleason, H. A. 1922. The vegetational history of the Middle West. Ass. Amer. Geogr. Annu.12:3 9-85.

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26 Land Use and Wildlife Resources Griffin, J. B. 1967. Eastern North American archaeology: a summary. Science 156: 175-191. Grinnell, J. 1923. The burrowing rodents of California as agents in soil formation. J. Mamm. 4: 137-149. Gross, A. O. 1928. The heath hen. Boston Soc. Natur. Hist. Memoirs 6(4~:491-588. Hanson, H. C. 1939. Fire in land use and management. Amer. Midl. Nat. 21 :415- 434. Hockensmith, R. D., and J. G. Steele. 1943. Classifying land for conservation farming. U.S. Dep. Agr. Farmers' Bull. 1853. Holdridge, L. R. 1947. Determination of world plant formations from simple climatic data. Science 105: 367-368. Hubbard, B. 1887. Memorials of a half-century. G. P. Putnam, New York. 581 p. Jackson, A. S. 1965. Wildfires in the Great Plains grassland. Tall Timbers Fire Ecol. Conf. Proc., 4th Annul Conf. Koford, C. B. 1958. Prairie dogs, whitefaces, and blue grama. J. Wildl. Manage. 3:78. Komarek, E. V. 1964. The natural history of lightning. Tall Timbers Fire Ecol. Conf. Proc., 3d Annul Conf. Kuchler, A. W. 1964. Potential natural vegetation of the coterminous U.S. Amer. Geogr. Soc. Spec. Publ. 36. 116p. Landstrom, K. L. 1958. How we acquired our landed estate, p. 19-27. In Land. The yearbook of agriculture 1958. U.S. Department of Agriculture. U.S. Gov- ernment Printing Office, Washington, D.C. Larson, F. 1940. The role of bison in maintaining the short grass plains. Ecology 21(2): 113-121. Lay, D. W. 1957. Browse quality and the effects of prescribed burning in southern pine forests. J. Forestry 55~5~:342-347. Lee, A. T. M., and H. H. Wooten. 1958. The ~nanagement of state lands, p. 72-86. In Land. The yearbook of agriculture 1958. U.S. Department of Agriculture. U.S. Government Printing Office, Washington, D.C. Lehmann, V. W. 1965. Fire in the range of Attwater's prairie chicken. Tall Timbers Fire Ecol. Conf. Proc., 4th Annul Conf. Leopold, A. S., and F. F. Darling. 1953. Wildlife in Alaska. Ronald Press, New York, 129 p. Little, S., Jr. 1953. Prescribed burning as a tool of forest management in the Northeastern states. J. Forestry 51~71:496-500. Lorant, S. (ed.~. 1965. The new world; the first pictures of America. Duell, Sloan & Pearce, New York. 292 p. McComb, A. L., and W. E. Loomis. 1944. Subclimax prairie. Torrey Bot. Club Bull. 71:46-76. Marsh, G. P. 1864. Man and nature: physical geography as modified by human action. C. Scribuer's, New York. 560 p. Marsh, R. E., and W. H. Gibbons. 1940. Forest-resource conservation, p. 458-488. In Farmers in a chang~ng world. The yearbook of agriculture 1940. U.S. De- partment of Agriculture. U.S. Government Printing Office, Washington, D.C. Martin, P. S. 1958. Pleistocene ecology and biogeography of North America, p.375420. In C. lIubbs (ed.~. Zoogeography. Amer. Ass. Advan. Sci. Publ.51. Washington, D.C.

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Historical Perspective Martin, P. S. 1963. The last 10,000 years. University of Arizona Press, Tucson. 87 p. Martin, P. S. 1967. Pleistocene overkill. Natur. Hist. 76~10~:32-38. Merriam, C. H. 1898. Life zones and crop zones of the U.S. Burl Biol. Surv. Bull. 10. 79 P. Mershon, W. B. 1907. The passenger pigeon. Outing Publ. Co., New York. 225 p. Mooney, J. 1928. The aboriginal population of America north of Mexico. Smith- sonian Misc. Coll. 80(7). 40 p. Muller-Beclc, H. 1966. Paleohunters in America: origins and diffusion. Science 1 52(3726): 1 19 1-1 209. Parkman, F. 1894. La Salle and the discovery of the Great West. Little, Brown & Co., Boston. 483 p. Parkman, F. 1895. Pioneers of France in the new world. Little, Brown & Co. Boston. 473 p. 27 Riebold, R. J. 1964. Large-scale prescribed burning. Tall Timbers Fire Ecol. Conf. Proc., 3d Annul Conf. Roe, F. G. 1951. The North American buffalo. University of Toronto Press, Ontario, Canada. 957 p. Ruess, L. A., H. H. Wooten, and F. I. Marschner. 1948. Inventory of major land uses, United States. U.S. Dep. Agr. Misc. Publ. 663. U.S. Government Printing Office, Washington, D.C. 89 p. Sauer, C. O. 1950. Grassland climax, fire, and man. J. Range Manage. 3~1): 16-21. Sauer, C. O. 1956. The agency of man on the earth, p. 49-69. In W. L. Thomas (ed.), Man's role in changing the face of the earth. University of Chicago Press, Chicago. Schorger, A. W. 1944. The prairie chicken and sharp-tailed grouse in early Wis- consin. Wis. Acad. Sci. Trans. 35: 1-59. Schorger, A. W. 1955. The passenger pigeon: its natural history and extinction. University of Wisconsin Press, Madison. 424 p. Scott, W. E., and N. H. Hoveland. 1951. Report to the people of Wisconsin on cover destruction, habitat improvement and watershed problems of the state in 195Q. Wis. Conserv. Bu11. 16~2~:3-77. Sears, P. B. 1942. Xerothermic theory. Bot. Rev. 8(10~:708-736. Sears, P. B. 1948. Forest sequence and climatic change in northeastern North America since early Wisconsin time. Ecology 29:326-333. Seton, E. T. 1929. Lives of game animals. Doubleday, Garden City, N.Y. 4 vol. Shantz, H. L. 1954. The place of grasslands in the earth's cover of vegetation. Ecology 35~2~: 143-145. Shelford, V. E. 1931. Some concepts of bioecology. Ecology 12:455-467. Shelford, V. E. 1963. The ecology of North America. University of Illinois Press Urbana. 610 p. Smith, P. W. 1957. An analysis of post-Wisconsin biogeography of the prairie peninsula region based on distributional phenomena among terrestrial verte- brate populations. Ecology 38:205-218. Smith, R. A. 1926. The land-economic survey in Michigan. Roosevelt Wildl. Bull. 3~4~: 679-692. Spinden, H. J. 1928. The population of ancient America. Geogr. Rev. 18:641-660.

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28 Land Use and Wildlife Resources Stewart, O. C. 1951. Burning and natural vegetation in the United States. Geogr. Rev. 41:317-320. Swanton, I. R. 1946. The Indians of southeastern United States. Burl Amer. Ethnol. Bull. 137. Swanton, J. R. 1952. The Indian tribes of North America. Burl Amer. Ethnol. Bull. 145. Tansley, A. G. 1935. The use and abuse of vegetational concepts and terms. Ecology 16~3~:284-307. Taylor, W. P. 1949. The biotic community concept as applied in historical geology. Tex. J. Sci. 1~1~:34-40. Thornthwaite, C. W. 1941. Climate and settlement in the Great Plains, p. 177-187. In Climate and man. The yearbook of agriculture 1941. U.S. Department of Agriculture. U.S. Government Printing Office, Washington, D.C. Thwaites, R. G. 1904. The original journals of the Lewis and Clark expedition. Dodd, Mead & Co., New York. Transeau, E. N. 1935. The prairie peninsula. Ecology 16~3~:423-437. U.S. Forest Service. 1936. The western range, p. 620. In Senate Doc. 199, 74th Congr., 2d Sess. U.S. Government Printing Office, Washington, D.C. Van Doren, M. redo. 1928. The travels of William Bartram. Dover Publ., New York. 414 p. Vestal, A. G. 1914. Internal relations of terrestrial associations. Amer. Natur. 48:413445. Weaver, J. E., and F. W. Albertson. 1956. Grasslands of the Great Plains. Johnson Publ. Co., Lincoln, Nebr. 395 p. Weaver, J. E., and F. E. Clements. 1938. Plant ecology. McGraw-Hill Book Co., New York. 601 p. Whittaker, R. H. 1953. A consideration of climax theory: the climax as a popula- tion and pattern. Ecology Monogr. 23:41-78. Wormington, H. M. 1962. A survey of early American prehistory. Amer. Sci. 50~1~: 230-242. Yarnell, A. R. 1964. Aboriginal relationships between culture and plant life in the Upper Great Lakes region. Univ. Mich. Mus. Anthrop. Pap. 23. 218 p.