Dynamics of Vegetation

(The Wilson Company, New York: 1949)
Frederic E. Clements

PLANT SUCCESSION AND HUMAN FOLLIES N.B.– The photographs that accompany this chapter are in a separate file 1. THE NATURE AND ROLE OF PLANT SUCCESSION Dynamic ecology concerns itself first and foremost with causes and processes and in consequence its dominant theme is one of change. The recognition of the basic fact of succession in vegetation has become more and more general in recent years, although at first it had been applied largely to local terrains such as dunes, bogs and swamps, with little recognition of their relation to the climax about them. As time has passed, however, an appreciation of the values to be obtained from a wider knowledge of the extent and character of the successive changes in the vegetative cover, has developed. As a consequence, federal agencies and large institutions have turned to intensive and extensive research into the subject, with results of the greatest practical importance to forestry, agriculture range management and soil conservation. Darwin once said that every traveler should be a botanist, since plants furnish the chief embellishment of all landscapes. Today it may be asserted with equal warrant that the traveler should be an ecologist if he is to understand the changes wrought by nature and by man upon the countenance of Mother Earth. Even the everlasting hills are not ageless, for they are worn down by wind and water; lakes are filled, rivers, grow old, and swamps become dry land subject to the plow. Intimately connected with these changes, hastening or retarding them, and in turn being modified by them, are the populations of living things, interacting in a maze of causes and effects of endless variety. Most responsive of these in the plant cover, forming the pattern of a complex community in which animals and primitive man in particular find shelter and homes and from which they draw food and materials. Every such community is essentially an organism, of a higher order than an individual geranium, robin or chimpanzee, but possessing structure and development, and a coordination of functions and parts similar in many respects. Like them, it is a unified mechanism in which the whole is greater than the sum of its parts and hence it constitutes a new kind of organic being with novel properties. Plant communities arise, grow, mature, attain old age and die from natural causes or by accident. They regularly reproduce themselves after partial destruction by fire, lumbering, clearing, or other disturbance, regenerating new parts, not altogether unlike the process by which a lobster grows a new claw or a lizard a tail. The final or adult community is termed a

logging. and of Pikes Peak. as well as of deciphering past event. restoration and rehabilitation under way or projected in the present national program (Plate 1B) . on rocky ridge and cliff. or indirectly by grazing. Each climax is the product of its particular climate and hence the indicator of it. and passing though a series of communities to end finally in the climax proper to each climate. on the exfoliating domes of Yosemite and in the sinter and diatom basins of YelIowstone geysers. and other high summits of the Rocky Mountains. and the prairies of many kinds of grasses. Thus.—Examples of the growth of climaxes. the use of indicators furnishes a method of primary importance (Chapter III). Related to these and hence of signal interest as seeming far out of place. and for all projects of utilization. Sierra Nevada. in sand-dunes and badlands. erosion. forestry. Plants Indicate Conditions. progressing slowly or rapidly in accordance as the site is water. and likewise its further story. and the (cascades.—Everyone is familiar in a general way with the great climaxes of our country and especially with the two most extensive. all survivals of a distant time of glacial advance when the arctic tundra moved far to the south. Kinds of Succession. Mount Rainier.—The significant outcome of these relations is that both species and communities serve as measures or indexes of conditions and hence are known to the ecologist as indicators. Mount Washington. draining. flooding and water supplies. cultivation and so forth. climax and Succession have not only great practical applications. and its process of growth is known as succession. a ruling class drawn usually from trees or grasses and best fitted to the climate concerned. flooding. and its major effects can be read with almost as much certainty as though recorded on the spot by an eye-witness (Plate lA). erosion. In addition to these are the great transcontinental forest of spruce and fir to the northward and the Barren Grounds of sedge and lichen stretching along the Arctic Circle from ocean to ocean. but others with a much longer life-span occur in pond and lakelet. maple chestnut and oaks. settlement. The primary indications have to do with climate and soil and the outstanding changes of the past. The Great Plant Climaxes. from the series of transient populations that pass across the scene. Most frequently seen are those due to disturbances caused by man. but woven into this pattern is the infinite variety wrought by man. and of various subordinate groups. agriculture. but also provide the open sesame by which traveler or nature-lover may unlock the pages of nature's book and read the past and present of every landscape. Each of these great communities consists of certain dominants. directly through fire. but they also possess the clairvoyance of forecasting future changes and the possibility of controlling them. rock or actual soil. .climax by reason of the fact that it is thy highest type of social organism capable of growing in a particular climate. grazing. Each of these processes has its own indicator communities. of their childhood and adolescence are to be found everywhere within the corresponding climate. the eastern forest of beech. among which the flowering herbs of woodland and prairie are the most conspicuous and familiar. Wherever an area is bare or is denuded by natural agencies or by man and his animals. and thus serves as the point of departure for all the disturbances brought about by man. In connection with land classification. are the alpine tundras of Mount Katahdin. development begins. They indicate not merely the present features of climate and soil. Mount Whitney.

secondary successions following fire or cultivation may take no more than a half-century for the complete cycle. cat-tails. grasses of progressively higher demands replace each other in forming a permanent cover. Succession on Rock. . As these grow and decay. yielding to mosses as a thin soil appears in crack and crevice. The early invaders are lowly annuals of small requirements. the general course is the same as that in succession from water. To he a sand-binder. the chief task of the pioneers is to convert rock into soil and to increase the water rather than diminish it. wild rice and reed-grass can invade usually in this order. mountain peaks. in water up to about twenty feet. When the ruling caste of woody plants is once established in a forest climate. In the prairie region. since the rainfall is nt sufficient for the development to continue to forest.—Probably the most familiar kind of succession is that found in standing water. In the prairies. but likewise bring about their own downfall by sha1lowing the water so that bulrushes. In the miniature deserts of rock-surfaces only the humblest plants can thrive. the succession terminates with a community of drouth-resisting grasses. hornworts and the like. As the soil increases in depth. to keep its head above the sand as the latter heaps up about it (Plate 3A). Thy soil usually is neither removed nor impaired. the Course of succession is quite different. cat-tails. Succession in Water. inasmuch as grasses are followed by shrubs and these by trees in the case of a forest climate ( Plate 2A). either forbs or grasses. After many years. which etch the surface and slowly produce a thin layer of dust. By contract with water-plants. tiny saxifrages and other "rock-breaking" herbs enter. lava fields and boulders everywhere. and hundreds of :ears to fill a lakelet to the point where meadow or woodland can flourish on the humus soil.—Succession on sand-dunes takes place more rapidly and dramatically since soil of a sort is already present and the major problem is to fix the shifting sand and enrich it with plant remains. bulrushes and sedges. a plant must not only be well-anchored and hold sand. and an abandoned field in the prairie may be reclaimed by the grasses in a decade or two. or in some cases to the latter directly (Plate 2B). and in the case of fire is often enriched by the minerals liberated.Primary successions on granite may require a thousand years or more between the pioneering crustlike lichens and the climax forest of oak or pine. the pond is gradually filled to the level at which floating plants can push in and take possession. which in their turn yield to grasses and afterwards to shrubs. trees of small demands and rapid growth overshadow the shrub stage. By contrast. while in forest regions the grasses yield ultimately to shrubs and trees ( Plate 3B). and these are followed after an interval by grasses.—On rocks ridges. The remains of these accumulate even more rapidly and in a few decades the pool may become a wet meadow covered with sedges. Succession in Soils. hut it must also be able to catch the load borne by the wind and even more important. pondweeds. These then rule as conquerors for a while. such as lichens and mosses which are capable of enduring dessication for months. The reconstruction of the adult community is a simpler and still more rapid process where fire or clearing has destroyed the climax. From this stage. The first settlers are crust-like lichens. and later yield to thy invading phalanx of climax trees of slower growth but greater permanence. leafy species gradually invade and carry the task forward. which gradually stabilize small areas for the entrance of an ascending series of perennials. with its communities of pond-lilies. The pioneer colony of this series is founded by submerged stoneworts.

Forces Concerned in Succession. The plants and animals of the community also exhibit many essential interactions.—Succession depends for its opportunity upon the production of bare or denuded areas. and purely agricultural stages of human society. Better known to us is the series of invasions that have swept over England. hunter. by the various Pueblan cultures of the Southwest. and Norman. The herbs are conquered by bushes and low shrubs. and minerals especially. these are succeeded by taller shrubs. first within the family and then spreading to larger and larger units under a slow but inevitable compulsion. Roman. The actual growth of the community is regulated by certain processes or functions by means of which soil and climate produce their effects. and shrubs gain successively a short period of mastery. though everywhere prevalent in prehistoric and ancient times. such relations become much more varied and important. homesteader. Many perennials and shrubs survive the fire and their root-sprouts soon appear in large number. the return of forest or prairie often requiring but two or three decades. The first recorded succession is that of Chellean. . and trees then begin to straggle into the copses.Mosses and liverworts appear almost at once. Brython. light. Mongol. A somewhat similar course is followed in cultivated fields that art allowed to "go back". and during the first full season a complete cover of annual forbs and grasses may be formed. After a few decades a young climax forest is again in possession. as in the pollination of flowers by insects and humming-birds. Persian. They exhibit aggregation and migration: reacting upon the environment and increasing control of it. Macedonian. and by the trapper. Chaldean. or augments the moisture of the air. reducing the light and taking the lion's share of the water in the soil (Plate 65). Aztec and the Spaniard in Mexico. by which individuals are brought together to form communities. Succession of Races and Cultures. A similar succession on our own continent is illustrated by the Maya. Babylonian. each succeeding community becoming less con-trolled by soil or terrain and more by climatic factors until the adult stage or climax is attained. increases the organic matter in the soil. Annual weeds dominate for a few years. especially in the hunting. The initial processes are aggregation and migration. grasses. pioneer. Mousterian. Toltec. and out of this has gradually emerged a new function. Solutrean and Magdalenian peoples in Europe. the actual conquest being brought about by the outcome of the competition for water. cooperation. from Sumerian to Akkadian. Assyrian. or take more or less complete control by means of sprouts. Succession has been less clearly perceived in human communities. When man enters the situation. Within each community there is likewise a certain amount of cooperation. Dane. and urbanite in the Middle West (Plate 20A). but the driving force back of it is climate. Goidel. Competition has been rife between and within them. while the most complex has been the sequence of races in Mesopotamia. the term itself indicating some popular appreciation of the process of succession. These react upon the soil and then upon the local climate to render conditions at first more favourable to themselves and later to the invaders that are to replace them.—lt is obvious that human communities are subject to the control of climate and soil—to what have often been called geographic influences. Tartar and Turk. involving Pict. Amorite. Angle and Saxon. as seen in the reaction that produces shade. pastoral. minimizes the effect of wind. while modern rivalries disclose certain aspects of it. Achulean. in some of which the mutual benefit is striking. gradually overtopping the herbs. and the usual communities of perennials.

and dry years were regarded merely as fleeting if unpleasant interludes in the westward march of empire. irrigating the valleys. until the barrier of an inhospitable frontier was thought to have vanished forever. railway and speculator with lands to sell. wave after wave of settlers has flung down the gauntlet to nature. the judgment of many individuals.—The solution of all problems that involve natural vegetation directly or indirectly. they would have met the hazards of successive frontiers more effectively or perhaps have refused the struggle against them. the task might have lagged endlessly had it not been for the conjunction of drouth. The ebbing of one wave was lost to view in the advancing crest of the next. It has sung of the con-quest of nature. as in the maintenance of all surface natural resources. APPLICATIO:N OF PLANT SUCCESSION TO HUMAN NEEDS The applications of the principles of plant succession to human problems and natural industries are manifold. However. and the tragic consequences of his short-sightedness were minimized or ignored. perhaps none was so potent as the illusion. and particularly of afforestation and reforestation. and it lies at the root of systems of forest management. In consequence. that increased rainfall followed cultivation and that each disastrous drouth must perforce be the last. the study of the chief plant communities of a region affords the best measure of the climate and its possibilities. and hence to regulated grazing and the utilization of the public domain. of heroism and success. in which millions of dollars in the Burkburnett oil-fields were involved. It is indispensable to land classification. fostered alike by state. turning the prairie sod with the plow.2. and burning with no heed of the morrow. and the conservation of water-supplies for irrigation and urban use. In addition. while the minor ones will reveal the significant variations of soil and topography. Had the newcomers been skilled in reading nature's books. An outstanding instance of their practical importance js to be found in the litigation between Texas and Oklahoma over the location of the boundary formed by the Red River. succession is invoked for its benefits in the rotation of crops. Clearing with the ax. and of the building of new commonwealths. When disturbance takes a hand. but in direct proportion to man's destructiveness. dust-storm. of un-witting destruction and unfulfilled hopes. The decision of the United States Supreme Court in favour of Texas was based upon the evidence obtained by application to the problem. Even less observant and intelligent was man in the face of his own destruction of capital in terms of land and cover. as is generally recognized in the case of fire and clearing. erosion and floods. It is the chief tool in the control of run-off. until the salvation of the semi-arid regions could only be brought about from outside. and the pioneer treading on his heels in the unending quest for a home. Among the many reasons for their failure to do either. is more dependable than that of one and the verdict of many different kinds of plants grouped in a community is much better. Even at this. The processes of nature were to be reversed by the wishing-wand of man. The widespread use of these is exemplified in all the disturbances wrought by man in the vegetation of the globe. these primary indications may be greatly modified and the . and flood to a degree and on a scale never before known. rests upon the fact that every plant or community is an indicator of the conditions about it and hence of the causes that lead to these. including game. of the principles of plant succession as laid down in 1916. caring little and knowing less of her resourceful and inexorable ways. Basis for Accurate Prediction. The epic of the West has long celebrated the frontiersman ever pressing onward in the search for open spaces. but it has muted the themes of failure and tragedy.

from which is derived their greatest value in human situations.pattern becomes much more complex. soil. but they also forecast what will happen in the future and hence serve as the basis for control of all kinds. Fortunately. but the outcome is still tinged with human optimism and befogged by the occurrence of wet and dry years. and the respective parts ascribed to climate. The vague terms "marginal" and "sub-marginal" should be dropped and all the lands of the West reorganized on the basis of regulated utilization grounded solely upon fact and not upon illusion. but they can be turned to account only as the indications of climate and terrain are understood and heeded. such mosaics can be disentangled by careful scrutiny. Further evidence is furnished by the heavy toll taken by erosion. intentional or otherwise. the belts of assured practice must be shifted eastward to the extent of several inches of rainfall. such as water-supplies. This is still true in spite of the fact that settlement throughout the West has demonstrated by trial and error what cannot be expected of the land in the various climates. In addition. but also the sequence in which changes occur. Changes in Practice Demanded. can be predicted with much definiteness (Chapter III). the facts are adequate to permit a classification upon the basis of proper use. The wind-breaks of the shelter-belt project are likewise to be regarded as crops. Between indicators on one hand and actual experience on the other. largely for the direct values in terms of lumber and forage. In a program of rehabilitation. The problems of forestry and grazing center about the maintenance of the natural climax. Tragedy of Erosion and Flood. but increasingly also for the indirect ones. Nevertheless. control of erosion and floods. as evidenced by the farmer's fight against weeds and by the fate of abandoned fields. quite out of harmony with the climate and climax of the prairie and hence to be maintained against the grass dominants only by exceptional means and in sites where soil or terrain is especially favorable. In short. but they are perennial woody ones. and recreation in the widest sense. plant communities not only reflect the controlling factors.—From the foregoing. the plant cover on the ground is an epitome of past events and future possibilities. it is evident that plant communities as indicators furnish the most satisfactory method of determining the best use of the land. though it must be realized that much farm experience runs to opinion rather than knowledge. at variance with the normal climatic and successional processes.—While the drama of erosion reached its climax in the duststorms of 1934 to 1936. and the cure of the latter js to be found only in a field system that substitutes some other control for the natural cover or makes an approach to this in stripcropping. Conservation to secure similar objectives is a major concern of the national parks and of those state parks that are something more than picnic resorts. On such a basis. agriculture and large-scale construction of various sorts rests upon the destruction of the native vegetation and are faced with the necessity for maintaining an artificial situation. it must be fully recognized that millions of acres in the Great Basin and the Southwest should be withdrawn from their extra-hazardous use and reserved for their scenic or recreation values which are or may be relatively high. On the other hand. or human interference. They indicate not merely previous conditions and communities. the curtain had risen upon the scene with the advent of white men . so that the grazing-forage industry occupies the western portion of the present farming region and the latter is restricted to the rainfall zone in which severe drouth and crop-failure come but once in a decade. and the outcome of its manipulation by man. the effects of the latter are of the most immediate importance.

marked by an intensity and extent never before seen in this country. or the cover destroyed by trails or roads. so must a realization of the inevitable relation between cause and effect bc invoked to cure them. As it was. the widespread conversion of range lands into dry farms giving drouth and high winds their chance to turn the exposed soil into clouds of suffocating dust and to pile it over houses and barns in great dunes (Plates 6 and 60A). the prairje broken. This Occurred in a climate and climax quite unlike those of southern California and arose from a wholly different kind of disturbance. before the natural processes of recovery could act. the range overgrazed. In the case of watersheds that have been seriously overgrazed. a rainfall of 12 inches in 36 hours found nothing to impede its descent as a torrential flood. He too failed signally to realize that dam and levee constituted but partial answers and that the adequate solution of the joint problems of erosion and flood lay in the control of watersheds. but the success of . Wherever the forest was burned or cleared. in large part. the higher and steeper areas being sown from the air (Plate 7). together with such terracing and furrowing as the terrain demands. Since the early stages of succession have less effect in restraining runoff and erosion. it is essential to hasten the return of adequate control by means of artificial seeding. This has been done on a large scale over the burns at Montrose and Santa Barbara and with excellent results. often imperceptibly but none the less steadily. these problems were nobody's business but that of the engineer and he could cope with the damage only after jt was done. Here again the primary cause was man's interference with natural processes. proper control of erosion and flooding can be secured only by removing all stock for a few years or by reducing the number to permit the recovery needed.—Just as a proper understanding of the role of disturbance would have prevented these calamities.and the action had increased in tempo with each generation. Fire above Mont-rose and La Crescenta denuded the steep mountain slopes of their protective cover of chaparral. a method that will at the same time salvage the fertile top-soil of farms and preserve the forage-value of grazing ranges (Plate 4). In May of 1935. The factor of safety must receive every possible emphasis. The towns at the foot of the mountains were overwhelmed and many lives lost. Here also restoration may be speeded up by sowing or planting. In short. It destroyed every bridge for 300 miles along the Republican River and piled up flood waters to threaten metropolitan cities 600 miles away. Until fires can be wholly eliminated. yet the flood effects in terms of damage to property and loss of life were identical. as well as to render them impossible jn the future. every precaution must be taken to avoid the consequences that too often ensue. and such adjuncts as debris basins and check dams must be so constructed as to play an assured part (Plate 66). The Problem of Recovery. the protective role of plants was lost and the surface soil began to slip away. as well as homes and property destroyed (Plate 5). a rain of high intensity falling on the badly over-grazed slopes of the PlatteArkansas divide in Colorado wrought similar havoc in the valley levels of Colorado Springs. A few months later. all feasible steps must be taken to hasten the natural succession. he might have been stirred to action. Had the farmer of the Middle West been able to visualize the floods of the lower Mississippi or the contribution of his own land to the rapidly growing delta at its mouth. The ecological principles involved in the connection between climate and climax and the effect of disturbances upon the latter are strikingly illustrated by tragic happenings in California in 1934. Better known because more novel. laden with destroying boulders. these destructive processes are to be stopped before they begin on the head-waters of all the minor tributaries. were the dust-storms of the summer of 1934 and the spring of 1935.

When the inevitable drouth enters such situations. There is no adequate remedy for such a condition until the rains return and the weed seeds everywhere scattered by the billion. and it has already become clear that it is much simpler to formulate methods for desettlement and resettlement than to carry them out in the face of the habits and practices of a whole countryside. it should be pointed out that nature's cooperation is essential to the success of the many present endeavours to undo man's destructiveness. scenic values may often be destroyed or reduced in the very regions where they are greatest. as by the construction of main highways that steadily increase in depth of cut and height of fill. massing. To insure against this. those that attend man's destructive acts in connection with road building and similar operations appear trivial. happened and can be made to happen. None the less. . and it is in this direction that progress is most needed. as well as in devastating floods and mud-flows. color and texture is indispensable to the highest type of highway composition (Plates 8B and 70). but the body of facts concerning climax and succession clearly indicate the essential features of the plan. these may also exert serious effects in terms of safety in travel. once in possession. Precautions in Road Building. The resulting succession is then controlled to produce in a few years the final pattern desired (Plates 8A.such measures depends largely upon the whims of the weather. But there is still to be achieved a much greater insight into nature's weather moods and the familiar but obscure chain of events known as climate. this should follow the principles of natural landscaping. In most cases. However. Practically all of the great projects now under way in the West would profit enormously by the ability to anticipate the more striking fluctuations in rainfall especially. it must be won by understanding and insight. Nature's Cooperation Essential.—By comparison with these staggering consequences to human welfare and survival. imitating natural succession in some degree but assuring a protective cover the first year. the latter offer opportunity for the action of natural succession. The problem of recovery becomes much more difficult and complicated if the native cover is completely removed as a consequence of tillage. which is characterized by employing native plants in harmony with the climax in general. Since this cannot be compelled. A quarter of a century may be required to reproduce the desired forage cover and it is imperative to employ an artificial succession under control that will restore the grasses in four or five years at the most.—As an epilogue. as well as upon the kind of climate and climax. the bare topsoil is whirled away in dust storms that may span the continent. and the use of these as indicators of what ha. or heaped into unsightly ridges. In addition. but against the slowness of this must be set the certainty that the first heavy rain will cause serious and often catastrophic wash-in and slumping. Such a comprehensive attempt at rehabilitation has never before been attempted. they can be displaced but slowly by the grasses. and rests squarely upon the major rule that well-ordered variety in materials. It is as far removed as possible from the straight rows of streetside exotics too often regimented along highways. leaving a sterile waste in place of a farm (Plate 64A) . 68 and 69). Like all bare areas. effective methods have been devised for holding the loose soil by terracing and planting. the chief problem is the human one. begin the task of succession to re clothe the soil. if not in all. These have already been attained in fair measure with respect to climax community the process of succession. Although they perform this job well against both wind and water. to say nothing of costs of maintenance and repair.

should be signlized by downpours and floods. and in harmony with the behavior of protracted drouths. This is indicated by the fact that the use of rainfall records in conjunction with the sunspot cycle suggested the probability that the succession of dry years on the Pacific Coast would end in supra-normal precipitation during the season of 1934-35. . The same indexes also warranted the expectation of normal rainfall or better in the drouth region. after the unprecedented series of six dry years.The first attempts to blaze the way to predictions made a year or more in advance have shown distinct promise. and further indicated that the actual break would come about May first.