Research methods in ecology

by Frederic E. Clements

358. Modification of habitat. As the final factors in ecesis and competition, water, light, and temperature control the grouping of plants into vegetation. An efficient change in one of these, or in all of them, brings about a visible adjustment in the structure of the plant group concerned. Modifications of water-content and light are readily produced in the field by drainage, irrigation, shading, clearing, etc. In fact, all the changes of habitat indicated under experimental evolution serve equally well to initiate experiments in experimental vegetation; indeed, the same experiment covers both fields. It is impracticable, however, to modify the temperature of a habitat without changing its water-content or light, and consequently the influence of temperature can not be determined through experiment by modification. The extent of the area modified should be as large as convenience will permit, in order that the number of individuals may be large enough to indicate clearly the resulting adjustment in position and arrangement. The best results can be obtained where a small separate area of a formation can be modified, e. g., where a small swamp can be drained, or a depression flooded. In the case of light, however, it is usually impossible to clear or to shade a large area, and the study must be restricted to a relatively small group of plants. In regions where lumbering is actively carried on, the consequent clearing initiates invaluable experiments over large areas, and this is likewise true of forest plantations. Modification of a large area has decided advantages in bringing out the changes in the more prominent structural features, but the causes and the details of the adjustment can be worked out much more satisfactorily in a small area.

359. Denuding. The modification of the habitat by denuding is the sole method of initiating succession by experiment. It is consequently of the most fundamental importance in investigating aggregation, ecesis, and competition, as well as the reactions exerted by the invaders of the different stages. The possibilities of denuding an entire habitat or an extensive area are not great, and the investigator must content himself with denuded quadrats, transects, and migration circles, which are small enough to permit a critical study of all the factors in succession. It is of course unnecessary that the denuding be done by the ecologist himself, provided he is able to follow the succession from the very beginning. Accordingly, it becomes possible for him to make the very best use of all those changes wrought by man in which the vegetation is destroyed over considerable areas. These are essentially natural experiments, and at this point the methods of natural and artificial habitats merge.

The manner of denuding depends in a degree upon the nature of vegetation, but, when time, convenience, and safety are all taken into account, the actual removal of the vegetation as indicated under the denuded quadrat is by far the most satisfactory. Under certain conditions, flooding or burning can be used to advantage, but cases of this kind are infrequent. The purpose of the experiment determines the kind of area to be denuded. Quadrat, transact, and migration circle are equally valuable for ecesis and competition. The quadrat is best adapted to work in a homogeneous area, while the transect is suited to a heterogeneous one characterized by zones, societies, or communities. It is an advantage to replace the denuded transect by a series of denuded quadrats, one for each zone or society, when the transect would be too long for convenience. The denuded migration circle is invaluable for aggregation and ecesis, since it makes possible the study of migration as a distinct function. A series of denuded quadrats, consisting of one or more in the different stages of a succession, furnishes important evidence concerning the development of each stage. By far the best method, however, for making a comparative study of the stages of a succession is the quadrat sequence. A quadrat is denuded each year, thus yielding a complete sequence of miniature stages through the whole course of succession. This method is especially valuable when a succession is represented by a single example, and there is no opportunity of reconstructing it by the comparison of various stages. A quadrat sequence is naturally of the greatest value if begun at the time when the first invaders appear.

360. Modification of the formation by transfer. The study of partial and intermittent invasion into an established vegetation is made through the transfer of a species or group of species by means of seeding or planting. The process differs in no way from that described for experimental evolution, except in so far that an endeavor is made to establish a family or a community, and not merely a few individuals. Transfer makes possible the critical investigation of ecesis under conditions of intense competition, as well as the study of aggregation and the origin of plant groups under these conditions. Perhaps its greatest value is in the experimental study of alternation and zonation, especially the former. It is practically impossible to determine whether alternation, especially when corresponsive, is due to physical or historical causes, i. e., migration and competition, except by means of the reciprocal transfer of the species concerned.

Field cultures for the careful study of ecesis and competition are made by transferring seeds or plants to new or denuded soils. This is practically a combination of the methods of modification and transfer. It has a unique value in making it possible to initiate artificial successions of almost any character that is desired, and to carry them out with the reactions more or less under control. This opens up an extremely important field of experimental inquiry, which promises to put the study of succession upon a much more exact basis. Competition cultures in the field are not essentially different from those under control, and they will be considered under the next method.

METHOD OF CONTROL HABITATS

Fig. 83. Simple culture of floating ecads of Ranunculus sceleratus.

361. Competition cultures. Although it is quite possible to carry on experiments in invasion and succession in the planthouse, the limited space usually available makes this undesirable, except in a few problems where control is necessary. Competition cultures, on the other hand, yield better results in the planthouse than in the field, since the physical factors and the appearance of unwelcome migrants are much more easily controlled. The possibilities of the culture method in the study of competition seem inexhaustible, and the author has found it necessary to confine his own investigations to a few of the fundamental problems. In this work, he has distinguished several kinds of cultures, based chiefly upon the species concerned and the arrangement of the individuals. Simple cultures are those in which a single species is used. The resulting group is a family, and the competition is between like individuals. In such cultures, the problem of the factors in competition is reduced to its simplest terms. Mixed cultures are based upon two or more species, and the problem is correspondingly complicated. As a rule, all the seeds have been sown at the same time in both simple and mixed cultures, but it has been found desirable to make some heterochronous cultures, in which seeds are also sown after the plants have appeared. Mixed cultures are distinguished as layered cultures, when the species are of very different height. Thus, rosettes have been grown with stemmed plants, tall slender forms with low branching ones, erect plants with twining and climbing plants, etc. Further evidence as to the nature of competition has been sought by means of ecad cultures, and factor cultures. In the former, plants of different response to water and light are grown together under the same conditions, in order to evaluate the part played by the nature of the plant. In a factor culture, the area is divided into two or more parts which are given different amounts of water or of light, in order to determine the influence of slight variations upon the same competitors. In somewhat similar fashion, an attempt has been made to ascertain the bearing of biotic factors upon competition. Cultures are easily made in which Cuscuta or parasitic fungi are used to place certain species at a disadvantage. Permanent cultures are obtained by allowing the plants to ripen and drop their seeds for several generations, just as in nature. They are indispensable for determining the final outcome of the competition between different species.