THE STRUCTURE OF THE FORMATION

Fig. 70. Alternating gravel slides on Mounts Cameron and Palsgrove, from the comparison of which the initial development of the talus succession has been reconstructed.

329. Method of alternating stages.The period of time through which a primary succession operates is usually too great to make a complete study possible within a single lifetime. Secondary successions run their course much more quickly, and a decade will sometimes suffice for stabilization,though even here the period is normally longer. The longest and most complex succession, however, may be accurately studied in a region, where several examples of the same succession occur in different stages of development. In the same region, the physical factors of one example of a particular succession are essentially identical with those of another example in the same stage. If one is in an initial stage, and the other in an intermediate condition, the development of the former makes it possible to reestablish more or less completely the life history of the latter. The same connection may be made between intermediate and ultimate stages, and it is thus possible to determine with considerable accuracy and within a few years the sequence of stages in a succession that requires a century or more for its complete development. In the Rocky mountains, gravel slides (talus slopes) are remarkably frequent. They occur in all stages of development, and the alternating slides of different ages furnish an almost perfect record of this succession. This method lacks the absolute finality which can be obtained by following a succession in one spot from its inception to final stabilization, but it is alone feasible for long successions, i. e., those extending over a score or more of years. When it comes to be universally recognized as a plain duty for each investigator to leave an exact and complete record in quadrat maps and quadrat photographs of the stages studied by him, it will be a simple task for the botanists of one generation to finish the investigations of succession begun by their predecessors.

330. The relict methodof studying succession is next in importance to the method of alternating areas. The two in fact are supplementary, and should be used together whenever relicts are present. This method is based upon the law of successive maxima, viz., the number of species and of individuals in each stage constantly increases up to a certain maximum, after which it gradually decreases before the forms of the next stage. In accordance with this, secondary species usually disappear first, principal species next, and facies last of all. There are notable exceptions to this, however, and the safest plan is to use the relict method only when principal species or facies are left as evidence. An additional reason for this is that secondary species are more likely to be common to two or more formations. In the majority of cases, the relict is not modified, and is readily recognized as belonging properly to a previous stage. This is true of herbs in all the stages of grassland, and in the initial ones of forest succession. The herbs and shrubs of earlier stages, which persist in the final forest stages, are necessarily modified, often in such a degree as to become distinct ecads, or species. The facies of the stages which precede the ultimate forest are rarely modified. The application of the relict method, together with the modificationjust described, is nicely illustrated by the balsam-spruce formation at Minnehaha. Of the initial gravel slide stage, the relicts areVagnera stellataandGalium boreale, the one modified intoVagnera leptopetala, and the other intoG. boreale hylocolum. The thicket stage is represented byHolodiscus dumosa, greatly changed in form and branching, and in the shape and structure of the leaf. The most striking relict of the aspen formation is the facies itself,Populus tremuloides. The tall slender trunks of dead aspens are found in practically every balsam-spruce forest. In many places, living trees are still found, with small, straggling crowns, which are vainly trying to outgrow the surrounding conifers. Of the aspen undergrowth,Rosa sayii,Helianthella parryi,Frasera speciosa,Zygadenus elegans,Castilleia confusa,Gentiana acuta, andSolidago orophilaremain more or less modified by the diffuse light. It is still a question whether the aspen stage passes directly into the balsam-spruce forest, or whether a pine forest intervenes. The presence of bothPinus ponderosaandP. flexilis, which are scattered more or less uniformly through the formation, furnishes strong evidence for the latter view.

Fig. 71. Relict spruces and aspens, showing the character of the succession immediately preceding the burn succession now developing.

The lifetime of forest and thicket stages of successions is ascertained by counting the annual rings of the stumps of facies. This is a perfectly feasiblemethod for many woodland formations where stumps already abound or where a fire has occurred, and it is but rarely necessary to cut down trees for this purpose. When trees or shrubs are present as relicts, the same method is used to determine the length of time taken by the development of the corresponding stages.

331.Since all the structures exhibited by formations, such as zones, layers, consocies, etc., are to be referred to zonation or alternation, these principles are first considered in detail. This, then, constitutes the basis for a consideration of the structure of a normal formation, with special reference to the different parts that compose it. The investigation of formational structure, since the latter is the result of aggregation, invasion, and succession, is accomplished by instruments, quadrats, etc., in the manner already indicated under development, and no further discussion of it is necessary here.

332. Concept.The recognition of vegetation zones dates from Tournefort[40], who found that, while the plants of Armenia occupied the foot of Mount Ararat, the vegetation of the slopes above contained many species of southern Europe. Still higher appeared a flora similar to that of Sweden, and on the summit grew arctic plants, such as those of Lapland.

As the historical summary shows, the concept of zonation is the oldest in phytogeography. Notwithstanding this, it has never been clearly defined, nor has there been any detailed investigation of the phenomenon itself, or of the causes which produce it. Zones are so common, and often so clearly marked, that they invite study, but no serious attempt has heretofore been made to analyze zonation, or to formulate a definite method of investigating it. Zonation is the practically universal response of plants to the quantitative distribution of physical factors in nature. In almost all habitats, one or more of the physical factors present decreases gradually in passing away from the point of greatest intensity. The result is that the plants of the habitat arrange themselves in belts about this point, their position being determined by their relation to the factor concerned. Close investigation will show that there is hardly a formation that is entirely without zonation, though in many cases the zones are incomplete or obscure for various reasons. Zonation is as characteristic of vegetation as a whole as it is of its unit, the formation, a fact long ago recognized in temperature zones. A continentalclimate, however, often results in the interruption of these, with the consequence that these belts of vegetation are not always continuous.

333. Growth.The causes that produce zones are either biological or physical: the first have to do with some characteristic of the plant, the second with the physical features of the habitat. Biological causes arise from the method of growth, from the manner of dissemination, or from the reaction of the species upon the habitat. The formation of circles as a result of radial growth is a well-known occurrence with certain plants, but it is much more common than is supposed. In the case of agarics, this phenomenon has long been known under the name of “fairy-rings.” It is found in a large number of moulds, and is characteristic of early stages of the mycelium of the powdery mildews. It occurs in nearly all maculicole fungi, and is exhibited by certain xylogenous fungi, such asHysterographium. Among the foliose lichens, it is a common occurrence with the rock forms ofParmelia,Placodium,Physcia, andLecanora, and with the earth forms ofParmeliaandPeltigera. The thalloid liverworts show a similar radial growth. The flowering plants, and many mosses also, furnish good examples of this sort of growth in those species which simulate the form of the mycelium or thallus. These are the species that form mats, turfs, or carpets. Alpine mat formers, such asSilene acaulis,Paronychia pulvinata,Arenaria sajanesis, etc., are typical examples. Xerophytic, turf-forming species ofMuhlenbergia,Sporobolus,Bouteloua,Festuca,Poa, and other grasses form striking ring-like mats, while creeping species ofEuphorbia,Portulaca,Amarantus, etc., produce circular areas. Rosettes, bunch-grasses, and many ordinary rootstalk plants spread rapidly by runners and rhizomes. The direction of growth is often indeterminate in these also, and is in consequence more or less bilateral or unilateral. Growth results in zonation only when the older central portions of the individual or mass die away, leaving an ever-widening belt of younger plants or parts. This phenomenon is doubtless due in part to the greater age of the central portion, but seems to arise chiefly from the demands made by the young and actively growing parts upon the water of the soil. There may possibly be an exhaustion of nutritive content, as in the case of the fungi, but this seems improbable for the reason that young plants of the same and other species thrive in these areas. It must not be inferred that these miniature growth zones increase in size until they pass into zones of formations. Growth contributes its share to the production of these, but there is no genetic connection between a tiny plant zone and a zone of vegetation.

Radial and bilateral growth play an important part in formational zones in so far as they are related to migration. The growth of the runner or rhizome itself is a very effective means of dissemination, while the seeding of the plants thus carried away from the central mass is most effective at the edge of the newly occupied area. This holds with equal force for plants with a mycelium or a thallus. The circular area becomes larger year by year. Sooner or later, the younger, more vigorous, and more completely occupied circumference passes into a more or less complete zone. This will result from the reaction of the central individuals upon the habitat, so that they are readily displaced by invaders, or from their increasing senility and dying out, or from the invasion of forms which seed more abundantly and successfully. This result will only be the more marked if the radiating migrants reach a belt of ground especially favorable to their ecesis. In this connection it must be carefully noted that vegetation pressure, before which weaker plants are generally supposed to flee, or by which they are thought to be forced out into less desirable situations, is little more than a fanciful term for radial growth and migration. It has been shown under invasion that disseminules move into vegetation masses, as well as away from them, the outward movement alone being conspicuous, because it is only at the margin and beyond that they find the necessary water and light for growth.

334. Reactions.Certain reactions of plants upon habitats produce zonation. The zones of fungi are doubtless caused by the exhaustion of the organic matter present, while in lichens and mosses the decrease in nutritive content has something to do with the disappearance of the central mass. In the mats of flowering plants, the connection is much less certain. The reaction of a forest or thicket, or even of a tall herbaceous layer, is an extremely important factor in the production of zonation. The factor chiefly concerned here is light. Its intensity is greatest at the edge of the formation and just below the primary layer; the light becomes increasingly diffuse toward the center of the forest, and toward the ground. In response to this, both lateral and vertical zones appear. The former are more or less incomplete, and are only in part due to differences in illumination. The vertical zones or layers are characteristic of forest and thickets, and are caused directly by differences in light intensity.

Fig. 72. Zones ofCyperus erythrorrhizusproduced by the recession of the shore-line.

335. Physical factors.The physical causes of zonation are by far the most important. They arise from differences in temperature, water, and light. In the large, temperature differences are the most important, producing the great zones of vegetation. In a particular region or habitat, variations of water-content and humidity are controlling, while light, as shown above, is important in the reactions of forest and thicket. Physical factors produce zonation in a habitat or a series of habitats, when there is either a gradual and cumulative, or an abrupt change in their intensity. Gradual, slight changes are typical of single habitats; abrupt, marked changes of a series of habitats. This modification of a decisive factor tends to operate in all directions from the place of greatest intensity, producing a characteristic symmetry of the habitat with reference to the factor concerned. If the area of greatest amount is linear, the shading-out will take place in two directions, and the symmetry will be bilateral, a condition well illustrated by rivers. On the other hand, a central intense area will shade out in all directions, giving rise to radial symmetry, as in ponds, lakes, etc. The essential connection between these is evident where a stream broadens into a lake, or the latter is the source of a stream, where a mountain ridge breaks up into isolated peaks, or where a peninsula or landspit is cut into islands. The line that connects the points of accumulated or abrupt change in the symmetry is a stress line orecotone. Ecotones are well-marked between formations, particularly where the medium changes; they are less distinct within formations. It is obvious that an ecotone separates twodifferent series of zones in the one case, and merely two distinct zones in the other.

Fig. 73. Regional zones on a spur of Pike’s Peak (3,800 m.); the forest consists ofPicea engelmanniiandPinus aristata, the forewold isSalix pseudolapponum, and the grassland, alpine meadow (Carex-Campanula-coryphium).

336. Physiographic symmetry.The physical symmetry of a habitat depends upon the distribution of water in it, and this is profoundly affected by the soil and the physiography. The influence of precipitation is slight or lacking, as it is nearly uniform throughout the habitat; the effects of wind and humidity are more localized. Differences of soil rarely obtain within a single habitat, though often occurring in a zoned series. The strikingly zonal structure or arrangement of habitats is nearly always due to differences in water-content produced by physiographic factors, slope, exposure, surface, and altitude. The effect of these upon water-content and humidity is obvious. Wherever appreciable physiographic differences occur, there will be central areas of excess and deficiency in water-content, between which there is a symmetrical modification of this factor. Peaks are typical examples of areas of deficiency, lakes and oceans of areas of excess. When these areas are extreme and close to each other, the resulting zonation will be marked; when they are moderate, particularly if they are widely separated,the zones produced are obscure. Asymmetry of a habitat or a region practically does not exist. Central areas of excess and deficiency may be very large and in consequence fail to seem symmetrical, or the space between them so great that the symmetry is not conspicuous, but they are everywhere present, acting as foci for the intervening areas.

The response of vegetation to habitat is so intimate that physiographic symmetry everywhere produces vegetational symmetry, which finds its ready expression in plant zones. The reaction of vegetation upon habitat causes biological symmetry, typical of growth zones and light zones. From these facts it is clear that zonation will be regularly characteristic of the vegetative covering. The zonal arrangement of formations is usually very evident; the zones of a formation are often obscured, or, where the latter occupies a uniform central area of excess or deficiency, they are rudimentary or lacking, as in shallow ponds. Zones are frequently imperfect, though rarely entirely absent in new soils, such as talus. They are rendered obscure in several ways. In the initial stages of a succession, as well as in the transitions between the various stages, the plant population is so scattered, so transient, or so dense as to respond not at all to a degree of symmetry which produces marked zonation in later formations. The alternation of conspicuous species not only causes great interruption of zones, but often also completely conceals the zonation of other species, such as the grasses, which, though of more importance in the formation, have a lower habit of growth. Furthermore, the ecotones of one factor may run at right angles to those of another, and the resulting series of zones mutually obscure each other. Finally, such a physiographic feature as a hill may have its symmetry interrupted by ridges or ravines, which deflect the zones downward or upward, or cause them to disappear altogether, while the shallows or depths of a pond or lake may have the same effect. An entire absence of zones, i. e., azonation, is exceptional in vegetation. Almost all cases that seem to exhibit it may be shown by careful examination to arise in one of the several ways indicated above.

337.Two kinds of zonation are distinguished with reference to the direction in which the controlling factor changes. When this is horizontal, as with water-content and temperature, zonation will be lateral; when it is vertical, as in the case of light, the zonation is vertical. There exists an intimate connection between the two in forests, where the secondary layer of small trees and shrubs is continuous with a belt of trees and shrubs around the central nucleus, and the lower layers of bushes and herbaceous plants with similar zones still further out. This connection doubtless arises fromthe fact that conditions are unfavorable to the facies, outside of the nucleus as well as beneath it. Floristically, each layer and its corresponding zone are distinct, as the one consists of shade, the other of sun species. Lateral zonation is radial when the habitat or physiographic feature is more or less circular in form, and it is bilateral, when the latter is elongated or linear. Vertical zonation is unilateral.

338. Radial zonationis regularly characteristic of elevations and depressions. From the form of the earth, it reaches its larger expression in the girdles of vegetation corresponding to the zones of temperature. The zones of mountain peaks are likewise due largely to temperature, though humidity is a very important factor also. Mountain zones are normally quite perfect. The zonation of islands, hills, etc., is due to water-content. In the former, the zones are usually quite regular and complete; in the latter, they are often incomplete or obscured. Prairies and steppes are not zoned as units, but are complexes of more or less zonal hills and ridges. Ponds, lakes, and seas regularly exhibit complete zones, except in those shallow ponds where the depth is so slight that what is ordinarily a marginal zone is able to extend over the entire bottom. The line between an elevation and a depression, i. e., the edge of the water level, is the most sharply defined of all ecotones. It separates two series of zones, each of which constitutes a formation. One of these is regularly hydrophytic, the other is usually mesophytic. The line between the two can rarely be drawn at the water’s edge, as this is not a constant, owing to waves, tides, or periodical rise and fall. There is in consequence a more or less variable transition zone of amphibious plants, which are, however, to be referred to the hydrophytic formation. Nearly all forest formations serve as a center about which are arranged several somewhat complete zones. As a rule, these merge into a single heterogeneous zone of thickets.

339. Bilateral zonationdiffers from radial only in as much as it deals with linear elevations and depressions instead of circular ones. With this difference, the zones of ranges and ridges correspond exactly to those of peaks and hills, while the same relation is evident between the zones of streams, and of lakes and ponds. The ecotones are identical except as to form; they are linear in the one and circular in the other. Incompleteness is more frequently found in bilateral zonation, though this is a question of distance or extent, rather than one of symmetry.

340. Vertical zonationis peculiar in that there is no primary ecotone present, on either side of which zones arrange themselves with reference tothe factor concerned. This arises from the fact that the controlling factor is light, which impinges upon the habitat in such manner as to shade out in but one direction, i. e., downward. Vertical zones appear in bodies of water, on account of the absorption of light by the water. In a general way, it is possible to distinguish bottom, plancton, and surface zones, consisting almost wholly of algae. There is little question that minor zones exist, especially in lakes and seas, but these await further investigation. The most characteristic vertical zones occur in forests, where the primary layer of trees acts as a screen. The density of this screen determines the number of zones found beneath it. In extreme cases the foliage is so dense that the light beneath is insufficient even for mosses and lichens. As a rule, however, there will be one or more zones present. In an ordinary deciduous forest, the layers below the facies are five or six in number: (1) a secondary layer of small trees and shrubs, (2) a tertiary layer of bushes, (3) an upper herbaceous layer of tall herbs, (4) a middle herbaceous layer, (5) a lower herbaceous layer, (6) a ground layer of mosses, lichens, other fungi, and algae. The upper layers are often discontinuous, the lower ones are more and more continuous. As a forest becomes denser, its layers disappear from the upper downward, the ground layer always being the last to disappear because of its ability to grow in very diffuse light. A vertically zoned formation shows a complex series of reactions. The primary layer determines the amount of heat, light, water, wind, etc., for the subordinate layers in general. Each of these layers then further determines the amount for those below it, the ground layer being subject in some degree to the control of every layer above it. This accounts probably for the definiteness and permanence of this layer. The degree to which the lower layers influence the upper by reacting upon the habitat is not known. It is evident that this influence must be considerable by virtue of their control of the water supply in the upper soil strata, by virtue of their transpiration, their decomposition, etc.

The ecotone between two formations is never a sharp line, but it is an area of varying width. The edge of this area which is contiguous to one formation marks the limit for species of the other. Both formations disappear in this transition zone, but in opposite directions. The overlapping which produces such zones arises from the fact that the physical factors tend to approach each other at the line of contact between formations, and that many species are more or less adjustable to conditions not too dissimilar.

341. Vegetation zones.As a fundamental expression of progressive change in the amount of heat and water, zonation is the most important feature of vegetation. It constitutes the sole basis for the division of continental as well as insular vegetation. The continent of North Americafurnishes striking proof of the truth of this. Conforming to the gradual decrease of temperature and water-content northward, three primary belts of vegetation stretch across the continent from east to west. These are forest, grassland, and polar desert. The first is further divided into the secondary zones of broad-leaved evergreen, deciduous, and needle-leaved forests. At right angles to this temperature-water symmetry lies a symmetry due to water alone, in accordance with which forest belts touch the oceans, but give way in the interior to grasslands, and these to deserts. It is at once evident that the mutual interruption of these two series of zones has produced the primary features of North America vegetation, i. e., tropical forests where heat and water are excessive, deserts where either is unusually deficient, grassland when one is low, the other moderate, and deciduous and coniferous forests, where the water-content is as least moderate and the temperature not too low. Such a simple yet fundamental division has been modified, however, by the disturbing effect which three continental mountain systems have had upon humidity and upon temperature symmetry. The two are intimately interwoven. The lowering of temperature due to altitude produces the precipitation of the wind-borne moisture upon those slopes which look toward the quarter from which the prevailing winds blow. A mountain range thus makes an abrupt change in the symmetry, and renders impossible the gradual change from forest to grassland and desert. The Appalachian system is not sufficiently high to produce a pronounced effect, and forests extend far beyond it into the interior before passing into prairies and plains. On the other hand, the influence of the Rocky mountains and the Sierra Nevada is very marked. The latter rise to a great height relatively near the coast, and condense upon their western slopes nearly all of the moisture brought from the Pacific. The Rocky mountains have the same effect upon the much drier winds that blow from the east, and the two systems in consequence enclose a parched desert. This series of major zones thus becomes, starting at the east, forest, grassland, desert, and forest, instead of the more symmetrical series, forest, grassland, desert, grassland, forest, which would prevail were it not for these barriers. This actual series of major zones undergoes further interruption by the action of these mountain systems in deflecting northern isotherms far to the south. This action is greatest in the high ranges, the Rocky mountains and the Sierras, and least in the lower Appalachians. Its result is to carry the polar deserts of the north far southward along the crests of the mountains, and to extend the boreal coniferous forests much further south along their slopes. In the Appalachians, this means no more than the extension of a long tongue of conifers into the mass of deciduous forests, and the occasional appearance of an isolated peak. In the western ranges, it produces two symmetricalseries of minor mountain zones, forest, alpine grassland or desert, and forest, to say nothing of the foot-hill and timber-line zones of thicket.

There seems to be no good reason for distinguishing the zones of mountains as regions. The term itself is inapplicable, as it has no reference to zonation, and is used much more frequently as a term of general application. Its use tends to obscure also the essential identity of the so-called vertical zones of mountains with the major continental zones, an identity which can not be insisted upon too strongly. For the sake of clearness, it is important to distinguish all belts of vegetation as zones, though it is evident that these are not all of the same rank. The following division of the vegetation of North America is based upon the fundamental principles of continental symmetry and the community of continental and mountain zones.

342. Concept.The term alternation is used to designate that phenomenon of vegetation, in which a formation recurs at different places in aregion, or a species at separate points in a formation. Although it is a fundamental feature of vegetation, it has been recognized but recently.[41]

Alternation is the response of vegetation to the heterogeneity of the surface of the earth. It is in sharp contrast to zonation, inasmuch as it is directly caused by asymmetry in the topography. In consequence, it deals with the subdivisions of zones, arising from physical differences within the symmetrical area. It deals with vegetation areas of every rank below that of major zone, with the habitat and geographical areas of species, and, in a certain way, with the correspondence of vicarious genera. The breaking up of vegetation into formations is a striking example of alternation. The same phenomenon occurs in every formation, producing consocies and minor plant groups, and everywhere giving variation to its surface and structure. The essential idea involved in this principle is the recurrence of like formations, consocies, or groups, which are more or less separated by formations, consocies, or groups differing from them. It is an exact expression of the primary law of association that heterogeneity of structure varies directly as the extent and complexity of the habitat, or the series of habitats. Vegetation is made up of what are superficially homogeneous formations, but upon analysis these are seen to contain consocies. The latter, though more uniform than formations, break up into groups, each of which still shows a characteristic heterogeneity arising from the varying number and arrangement of its constituent species.

343. Causes.The primary cause of alternation is physical asymmetry, which is everywhere present within the symmetrical areas which produce zones. This is influenced so strongly, however, by migration and plant competition (phyteris) that the consideration of this subject will gain in clearness if these are treated as separate causes. The essential relation between them must not be lost sight of, however. Migration carries disseminules into all, or only some of the different areas of a formation, or into different formations, with little respect to the physical nature of these. The physical character of these asymmetrical areas determines that some of these plants shall be established in one series of places, and some in another, while the competition between the individuals in the various areas determines the numerical value of each species as well as its persistence. These three causes are invariably present in the production of alternating areas, and originally, i. e., in new or denuded soils, the sequence is constant, viz., migration, ecesis in asymmetrical areas, and competition.

With respect to the different portions of an asymmetrical area, migration will have one of three effects: (1) it will carry disseminules into both favorable and unfavorable areas, (2) into favorable ones only, or (3) into unfavorable ones alone. From the radial nature of migration, the first case is far the most frequent; it is typical of sporostrotes, and the highly specialized spermatostrotes and carpostrotes. The effect of migration is uniform here, and alternation arises in consequence of the selective power of ecesis. It is evident that migration does not have an even indirect effect, when the disseminules are carried into none but unfavorable situations. Where the movement is into favorable places alone, alternation is the immediate result. The intermittent operation of migration and the presence of barriers are responsible for the absence of plants in situations favorable to them, and in consequence bring about a certain alternation between corresponding species.

The selective operation of physical factors upon the disseminules carried into the different parts of an asymmetrical area is the usual cause of alternation. Asymmetry alone is universal within the more conspicuous structures termed zones, down to the smallest areas which a group of plants can occupy. The difference between contiguous areas, particularly within the same habitat, is often small. It sometimes seems inefficient in the initial stages of a succession when a single species is present, but even in extreme cases its effect will be recognizable in the size and density of the individuals. Asymmetry is clearly evident in vegetation where two symmetrical series cross each other, or when a symmetry is interrupted by barrier-like elevations or depressions. Within formations, it arises from differences, often very slight, in slope, exposure, elevation, from irregularities of surface, differences in soil structure, or composition, in the amount of cover, and in the reactions of the living plants. At the last point, it is in direct connection with plant competition.

344. Competition.Much uncertainty, as well as diversity of opinion, seems still to exist in regard to the precise nature of the competition between plants that occupy the same area. It has long been admitted that the phrase, “struggle for existence,” is true of this relation only in the most figurative sense, but the feeling still prevails that, since plants live in associations, there must be something mysterious and vitalistic in their relation. No one has been able to discover anything of this nature, but nevertheless the impression remains. Such a direct relation exists only between parasites, epiphytes, and lianes, and the plants which serve to nourish or support them. In the case of plants growing on the same stratum, actual competition between plant and plant does not occur. One individual can affect another only in as much as it changes the physical factors that influence the latter. Competition is a question of the reaction of a plant upon the physical factors which encompass it, and of the effect of these modifiedfactors upon the adjacent plants. In the exact sense, two plants do not compete with each other as long as the water-content and nutrition, the heat and light are in excess of the needs of both. The moment, however, that the roots of one enter the area from which the other draws its water supply, or the foliage of one begins to overshade the leaves of the other, the reaction of the former modifies unfavorably the factors controlling the latter, and competition is at once initiated. The same relation exists throughout the process; the stronger, taller, the more branched, or the better rooted plant reacts upon the habitat, and the latter immediately exerts an unfavorable effect upon the weaker, shorter, less branched, or more poorly rooted plant. This action of plant upon habitat and of habitat upon plant is cumulative, however. An increase in the leaf surface of a plant not merely reduces the amount of light and heat available for the plant near it or beneath it, but it also renders necessary the absorption of more water and other nutritive material, and correspondingly decreases the amount available. The inevitable result is that the successful individual prospers more and more, while the less successful one loses ground in the same degree. As a consequence, the latter disappears entirely, or it is handicapped to such an extent that it fails to produce seeds, or these are reduced in number or vitality.

Competition in vegetation furnishes few instances as simple as the above, but this will serve to make clear the simplest case of ordinary competition, i. e., that in which the individuals belong to a single species. The various individuals of one species which grow together in a patch show relatively slight differences, in height, width, leaf expanse, or root surface. Still, some will have the largest surfaces for the impact of water, heat, and light, while others will have the smallest; the majority, perhaps, will occupy different places between the extremes. The former will receive more than their share of one or more factors. The reaction thus produced will operate upon the plants subject to it inversely as the amount of surface impinged upon. The usual expression of such competition is seen in the great variation in height, branching, etc., of the different individuals, and in the inability of many to produce flowers. This is particularly true of annuals, and of perennials of the same generation. In the competition between parents and offspring of the same perennial species, the former usually have so much the advantage that the younger plants are often unable to thrive or even germinate, and disappear, leaving a free space beneath and about the stronger parents. This illustrates the primary law of competition, viz., that this is closest when the individuals are most similar. Similar individuals make nearly the same demands upon the habitat, and adjust themselves least readily to their mutual reactions. The more unlike plants are, the greater the difference in their needs, and some are able to adjust themselves to the reactions of others with little or no disadvantage.

In accordance with the above principles, the competition is closer between species of like form than between those that are dissimilar. This similarity must be one of vegetation or habitat form, not one of systematic position. The latter is in fact of no significance, except where there is a certain correspondence between the two. Leaf, stem, and root characters determine the outcome, and those species most alike in these features will be in close competition, regardless of their taxonomic similarity or dissimilarity. This is as conclusive of the competition between the species of the same genus as it is between those belonging to genera of widely separated families. From this may be deduced a second principle of competition, viz., the closeness of the competition between the individuals of different species varies directly with their similarity in vegetation or habitat form. This principle is of primary importance in the competition which arises between occupants and invaders in the different stages of succession. Those invading species that show the greatest resemblance to occupants in leaf, stem, and root form experience the greatest difficulty in establishing themselves. The species, on the contrary, which are so unlike the occupants that they come in at a clear advantage or disadvantage, establish themselves readily, in the one case as a result of the reaction, in the other by taking a subordinate position. This principle lies at the base of the changes in succession which give a peculiar stamp to each stage. A reaction sufficient to bring about the disappearance of one stage can be produced only by the entrance of invaders so different in form as to materially or entirely change the impress of the formation. Stabilization results when the entrance of invaders of such form as to exert an efficient reaction is no longer possible. In forests, while many vegetation forms can still enter, none of these produce a reaction sufficient to place the trees at a disadvantage, and the ultimate forest stage, though it may change in composition, can not be displaced by another.

It is obvious that the vegetation forms and habitat forms of associated species are of fundamental importance in determining the course and result of competition. Identity of vegetation form regularly produces close competition, and the consequent numerical reduction or disappearance of one or more species. Dissimilarity, on the other hand, tends to eliminate competition, and to preserve the advantage of the superior form. Species of trees compete sharply with each other when found together; the same is true of shrubs, or rosettes, etc. The relation of the shrubs to the trees, or of the rosettes to the shrubs of a formation is one of subordination rather than of competition. The matter of height and width often enters here also to such a degree that the tallest herbs compete with the bushes and shrubs, and rosettes with mats or grasses. The amount and disposition of the leaf surface are decisive factors in the competition between species of the samevegetation form, in so far as this is governed by light. In those plants in which the leaves are usually erect, notably the grasses and sedges, the competition between the aerial parts is relatively slight, and the result is determined by the reactions of the underground stems and roots.

The position of the competing individuals is of the greatest importance. The distance between the plants affects directly the degree of competition, while their arrangement, whether in groups according to species or singly, exerts a marked influence by determining that the contest shall be between like forms, or unlike forms. Position is controlled primarily by the relation existing between seed-production and dissemination. It is of course influenced in large measure by the initial position taken by the invaders into a nudate area, but this is itself a result of the same phenomena. The individuals of species with great seed-production and little or no mobility usually occur in dense stands. In these, the competition is fierce, for the two reasons of similarity and density, and the result is that the plants fall far below the normal in height and width. This is an extreme example of the group arrangement. When the seed-production is small, the mobility may be great or little without seriously affecting the result. The individuals of a species of this kind will be scattered among those of other species, and the closeness of competition will depend largely upon the similarity existing between the two. The arrangement in such cases is sparse. A species with great seed-production and great mobility usually shows both kinds of arrangement, the position of the individuals and the competition between them varying accordingly. This is due to the intermittent action of distributing agents, making it possible for the seeds to fall directly to the ground during the times that winds, etc., are absent. The three types of arrangement indicated above are termed gregarious, copious, and gregario-copious. They furnish the basis for the investigation of abundance which deals essentially with the number and arrangement of the individuals of competing species. The effect of distance, i. e., the interval between individuals, upon competition is fundamental. The competition increases as the interval diminishes, and the reverse.

The view here advanced, i. e., that competition is purely physical in nature, renders untenable the current conceptions of vegetation pressure, occupation, etc. Masses of vegetation are thought to force the weaker species toward the edge, thus initiating an outward or forward pressure. As has been shown above, no such phenomenon occurs in vegetation. This movement is nothing but simple migration, followed by ecesis, and has no connection with “weaker” species, or the development of a vital pressure. The direction taken by the migrating disseminules is essentially indeterminate. Migration seems to be outward, or away from the mass, merely because theecesis is greater at the edge, where the increased dissimilarity between plant forms diminishes the competition. The actual movement is outward, but it takes place through the normal operation of competition. In this connection, it should be pointed out that the common view that plants require room is inexact, if not erroneous. This is difficult of proof, as it is impossible to distinguish room as such from the factors normally present, light, heat, water, and nutrient salts, but it seems obvious that the available amounts of these will determine the space occupied by a plant, irrespective of the room adjacent plants may allow it. The explanation of competition upon physical grounds likewise invalidates the view that plants possess spheres of influence other than the areas within which they exert a demonstrable reaction upon the physical factors present.

Competition plays a very important role in alternation. It produces minor examples of alternation in the physical units of an asymmetrical series. Its greatest influence, however, is exerted in modifying the effects of asymmetry. The reaction of occupants emphasizes or reduces the effect of asymmetry, and has a corresponding action upon alternation. This result of competition is typical of succession, in which the sequence of stages arises from the interaction of occupant and invader.

345. Kinds of alternation.Alternation involves two ideas, viz., the alternation of different species or formations with each other, and the alternation of the same species or formation in similar but separate situations. This is the evident result of asymmetry, in response to which contiguous areas are dissimilar and remote ones often similar. Individuals of the same species or examples of the same formation may be said to alternate between two or more similar situations, while different species or formations are said to alternatewitheach other, occurring usually in situations different in character. From the nature of alternation, the two phenomena are invariably found together.

It is possible to distinguish three kinds of alternation: (1) of a formation, consocies, layer, facies, or species in similar situations; (2) of similar or corresponding formations, species, etc., in similar situations; (3) of facies and other species with respect to number. The last two are merely variations of the first, arising out of slight differences in the physical factors of the alternating areas, the adjacent flora, or the course of competition. The alternation of different examples of the same formation is a significant feature of greatly diversified areas, such as mountains. It is naturally much less characteristic of lands physiographically more uniform. A xerophytic formation will alternate from ridge to ridge, a mesophytic formation between the intermediate valleys; aquatic vegetation will alternate from pond to pond,or stream to stream. The appearance of new or denuded soils upon which successions establish themselves is the most important cause of the alternation of formations. The weathering of rocks in different areas of the same region produces in each a sequence of similar or identical formations. The same statement is true in general of other causes of succession, such as erosion, flooding, burning, cultivation, etc., wherever they operate upon areas physically similar and surrounded by the same type of vegetation. The areas of more or less heterogeneous formations characterized by major physical differences are occupied by consocies. In an extensive formation, the same consocies alternates from one to another of these areas that are similar. When the formation is interrupted and occurs here and there in separate examples, a consocies often alternates from one to another of these. A consocies regularly derives its character from the fact that one or more of the facies of the formation is more intimately connected with certain areas of the latter than with others. This explains why the alternations of consocies and facies are usually identical. Layers sometimes alternate between different examples of the same forest or thicket formation, when they are suppressed in some by the diffuseness of the light.

The alternation of species is a typical feature of formations; it is absent only in those rare cases where the latter consist of a single species. The areas of a habitat which show minor physical or historical (i. e., competitive) differences are occupied by groups of individuals belonging to one or more species responsive to these differences. Each of these groups will recur in all areas essentially similar, the intervals being occupied of course by slightly different groups. Such groups are constituted by gregarious or copious species of restricted adjustability. Sparse plants likewise alternate, but they necessarily play a much less conspicuous part. In habitats not too heterogeneous, a large number of species are sufficiently adjustable to the slight differences so that they occur throughout the formation. Often, to be sure, they show a characteristic response, expressed in the size or number. This is illustrated by the facies and many of the principal species of the prairie formation.Festuca,Koelera,Panicum, andAndropogonoccur throughout, except in the moist ravines which are practically meadows.Astragalus,Psoralea,Erigeron, andAstergrow everywhere on slopes and crests, but they are much more abundant in certain situations. Other plants,Lomatium,Meriolix,Anemone,Pentstemon, etc., recur in similar or identical situations upon different hills.Lomatiumalternates between sandy or sandstone crests,MeriolixandPentstemonoccur together upon dry upper slopes, whileAnemonealternates between dry slopes and crests.

Owing to the accidents of migration and competition, similar areas within a habitat are not occupied by the same species, or group of species. A speciesfound in one area will be replaced in another by a different one of the same or a different genus. The controlling factors of the area render imperative an essential identity of vegetation and habitat form, though in systematic position the plants may be very diverse. Such genera and species may be termedcorresponding. The relation between such plants is essentially alternation; it should, perhaps, be distinguished from alternation proper ascorresponsive. The prairie formation furnishes a good example of this on exposed sandy crests, upon whichLomatium,Comandra, andPentstemonalternate. Formations exhibit a similar correspondence.

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