Research methods in ecology

by Frederic E. Clements

CARTOGRAPHY

230. Value of cartographic methods. Chart, map, and photograph are records indispensable to the systematic study of vegetation. They serve not merely to preserve the facts ascertained, and to permit their ready comparison, but they also put a premium upon accurate methods, and consequently bring to light many points otherwise overlooked. For ecology, they have the value which drawings possess in taxonomy, in that they make clear at a glance what pages of description fail to indicate. They are the fundamental material of comparative phytogeography, and in all careful vegetational study their use is no longer optional but obligatory. Hence it is obvious that cartographic methods should be clear and simple, and that they should be uniform, so that charts and maps of widely separated formations may be directly compared without difficulty. It is not to be expected that uniform methods will come into general use immediately, but a proper appreciation of the obligation that rests upon every ecologist to make his results both easily comprehensible and usable will serve to produce this very necessary result. In the treatment that follows, as elsewhere, no attempt is made to describe the general cartographic methods used by other ecologists, notably Flahault. The methods employed by the author form a complete system, which has proved valuable, and for various reasons it alone is discussed here.

231. Standard scale. The question of the scale to which charts and maps are to be made is of primary importance. The general principle is that the ratio between area and drawing should be as small as possible. Moreover, charts and maps of the same character should always be drawn to the same scale, unless a good reason to the contrary exists. The ideal scale is 1:1, which is manifestly an impossibility. This is approached most nearly in the quadrat chart where the scale is 10:1. Charts of definite areas are made on a scale as large as possible, while maps of formations, regions, etc., are necessarily drawn upon a very small scale. General maps designed to show the distribution of species and formations, or the vegetation of continents, are usually not drawn with reference to a scale at all. While it is manifestly impossible to use the same scale for charts and maps, it is feasible and desirable that they be constructed upon scales readily convertible into each other. This is most satisfactorily accomplished by means of the decimal system, and the various type scales are 10:1, 100:1, 1000:1, etc. The first two or three scales are used for charts of quadrats, transects, and circles; the remaining ones are employed in making maps of large areas. No attempt has been made to draw an absolute line between charts and maps, but an endeavor is made to restrict the term chart to the record of the number and position of plants, while maps deal with the arrangement and location of formational areas. It is hardly necessary to point out the reasons why all charts and maps should be based upon the decimal system of scales. Experience will furnish the very best of arguments.

232. Color scheme. The first requisite for the graphic representation of formations, regions, etc., is that each class of formations be invariably indicated by the same color. It is also necessary that the colors and shades be easily distinguishable, and it is at least desirable that they be referred to the different classes in some consistent sequence. Uniformity in all these points is greatly to be desired at the hands of all ecologists. Here, as in the case of the standard scale, uniformity will be found the more desirable the more impossible it is made by ignoring it. In the use of color to represent regions and provinces, on maps too small to indicate formations, the color of each division is represented by the color of its dominant formation; thus the prairie province is colored ochroleucus on account of the color used to represent prairie formations, the boreal-subalpine zone atrovirens on account of the typical coniferous forests, etc. No endeavor has been made to take account of the various types of formations, e. g., the different coniferous forests, as this is a problem to be worked out for more local maps in various shades of dark green, etc. The following color scheme which has been based upon the points made above is proposed as a satisfactory solution of the problem. The color standard used is that of Saccardo’s Chromotaxia.

I. Hydrophytic Formations: blue

1. Marine: cyaneus 2. Brackish: ardesiacus 3. Freshwater: caeruleus 4. Swamps and marshes: caesius

A. Forest formations: green

1. Coniferous forests: atrovirens 2. Broadleaved evergreen forests: viridis 3. Deciduous forests: flavovirens

1. Meadows: melleus 2. Prairies: ochroleucus

1. Fields: ruber 2. Groves and orchards: atropurpureus 3. Wastes: purpureus

1. Deserts: isabellinus 2. Plains and steppes: avellaneus 3. Saline formations: umbrinus 4. Arctic-alpine formations: testaceus

233. Formation and vegetation maps are detailed maps of a single formation or a series of them, showing the formational limits, and when the scale is not too small, the ecotones of zones and consocies. In the cases where the topography is level, as sometimes happens in mapping single formations, the chain and pedometer must be used to ascertain the size of the different areas. Indeed in all mapping of vegetation, the methods of surveying are directly applicable. Over large areas, however, it is not necessary that limits be drawn with mathematical accuracy, and for the purposes of the ecologist, the plane table and camera are satisfactory substitutes for the surveyor’s transit, at least in the present aspect of the subject. When the formation or group of formations is commanded by an elevation of some height, the latter is used as a base. A plane table is established upon it and the topographical and vegetational features are recorded in the usual way. This map is usually supplemented by a series of views from the same base. Indeed it has come to be recognized that a complete series of photographs of this kind give a more valuable record than the plane table, and that the construction of an accurate map from them is an easy matter. Since the camera saves much time and energy also, it is used almost exclusively to furnish the data for map making. In hilly, and especially in mountainous regions, the photographic method is indispensable. Its application is extremely simple. A central hill or mountain is selected, and from it a series of views is taken so that the edge of one exactly meets the edge of the other. This is an extremely important matter, and demands much nicety of judgment. The camera is kept in the same spot, and after each exposure it is turned as the operator looks through it until a landmark at one edge just passes from view at the other. As soon as the new position is determined, the tripod screw is turned to hold the box firmly in position. In case of a slight jar, the exact position should again be obtained. If the series is accurately made, the resulting prints will give a complete panoramic view of the region, without overlap or omission. For this purpose, a 6½ × 8½ camera is desirable, since the topographic and vegetational features are larger and stand out more distinctly. A large camera requires fewer changes of position, and hence saves time and reduces the chance of error. A 4 × 5 camera serves the purpose sufficiently well, though it requires a little more care in operation on account of the greater number of exposures necessary. This may be avoided in some degree by the use of a wide-angle lens if the depth of the area is not too great. Whatever camera may be used, a telephoto lens is a very desirable adjunct, since it enables one to choose between three different sizes of the view without changing the position of the camera. To avoid possible confusion, the exposures are always made from right to left, and the plates are used in the numerical order of their holders. For the same reason the landmarks are described and numbered in their proper order. The prints obtained are mounted on a card in sequence. The view map may be preserved in this form, or it may be reduced or enlarged by making a copy to the size desired. Outline maps of topography may be traced from the resulting negative, and the formations filled in by means of the proper colors. The most satisfactory method, however, is to have the original views or the copy printed “light” and to color the formations just as they appear there, with all the wealth of topographic and vegetational detail. If a detailed topographic map alone is desired, this is traced directly from the large copy.

234. Continental maps. A method of determining the general outlines of regions, provinces, and vegetational zones as a preliminary to their detailed study has been used successfully for several years.[^[22]] This is based upon provincial and continental maps on which are traced the geographical areas of the species of genera typical of the various formations. Detail topographic maps of the prairie province and the North American continent have been used for this purpose. A number of the facies of extensive and representative formations of the different portions of the continent are selected and grouped according to genera. One map is devoted to each genus, unless the number of species is large. In this case a number of maps are used, since the limits are apt to become confused. The range of each species is determined from all the reliable sources, and a corresponding line is drawn upon the map to delimit its geographical area. The limits of the area of each species are drawn in a different color, and the name of the species printed in the same color in the legend. Although this work has as yet been done only for the trees of North America, and for the grasses and principal species of the prairie province, it promises to constitute a final method for the limitation of vegetational divisions. It is clear that if the original data concerning ranges are accurate, the increasing study of formations will do little more than rectify the detailed course of the limiting line, since in most cases facies and formations coincide in distribution. The limiting line or ecotone of a zone or province is a composite obtained from the limits of certain representative facies and principal species, and checked by the limits of species typical of the contiguous vegetations. Thus, the boreal-subalpine zone is clearly outlined by combining the limits of Populus tremuloides, Larix americana, Pinus banksiana, Abies balsamea, Picea mariana, Picea canadensis, and Betula papyracea, and checking the results by the areal limits of the hardwoods and grasses to the southward.