Assembled here for the first time in one volume are forty classic papers that have laid the foundations of modern ecology. Whether by posing new problems, demonstrating important effects, or stimulating new research, these papers have made substantial contributions to an understanding of ecological processes, and they continue to influence the field today.
The papers span nearly nine decades of ecological research, from 1887 on, and are organized in six sections: foundational papers, theoretical advances, synthetic statements, methodological developments, field studies, and ecological experiments. Selections range from Connell's elegant account of experiments with barnacles to Watt's encyclopedic natural history, from a visionary exposition by Grinnell of the concept of niche to a seminal essay by Hutchinson on diversity.
Six original essays by contemporary ecologists and a historian of ecology place the selections in context and discuss their continued relevance to current research. This combination of classic papers and fresh commentaries makes Foundations of Ecology both a convenient reference to papers often cited today and an essential guide to the intellectual and conceptual roots of the field.
Published with the Ecological Society of America.
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About the Author
James H. Brown is a Distinguished Professor of Biology at the University of New Mexico, and past president of the International Biogeography Society.
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Foundations of EcologyClassic Papers with Commentaries
THE UNIVERSITY OF CHICAGO PRESSCopyright © 1991 The University of Chicago
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Chapter OneArticle IX.—The Lake as a Microcosm. By Stephen A. Forbes.
A lake is to the naturalist a chapter out of the history of a primeval time, for the conditions of life there are primitive, the forms of life are, as a whole, relatively low and, ancient, and the system of organic interactions by which they influence and control each other has remained substantially unchanged from a remote geological period.
The animals of such a body of water are, as a whole, remarkably isolated—closely related among themselves in all their interests, but so far independent of the land about them that if every terrestrial animal were suddenly annihilated it would doubtless be long before the general multitude of the inhabitants of the lake would feel the effects of this event in any important way. It is an islet of older, lower life in the midst of the higher, more recent life of the surounding region. It forms a little world within itself—a microcosm within which all the elemental forces are at work and the play of life goes on in full, but on so small a scale as to bring it easily within the mental grasp.
Nowhere can one see more clearly illustrated what may be called the sensibility of such an organic complex, expressed by the fact that whatever affects any species belonging to it, must have its influence of some sort upon the whole assemblage. He will thus be made to see the impossibility of studying completely any form out of relation to the other forms; the necessity for taking a comprehensive survey of the whole as a condition to a satisfactory understanding of any part. If one wishes to become acquainted with the black bass, for example, he will learn but little if he limits himself to that species. He must evidently study also the species upon which it depends for its existence, and the various conditions upon which these depend. He must likewise study the species with which it comes in competition, and the entire system of conditions affecting their prosperity; and by the time he has studied all these sufficiently he will find that he has run through the whole complicated mechanism of the aquatic life of the locality, both animal and vegetable, of which his species forms but a single element.
It is under the influence of these general ideas that I propose to examine briefly to-night the lacustrine life of Illinois, drawing my data from collections and observations made during recent years by myself and my assistants of the State Laboratory of Natural History,
The lakes of Illinois are of two kinds, fluviatile and water-shed. The fluviatile lakes, which are much the more numerous and important, are appendages of the river systems of the state, being situated in the river bottoms and connected with the adjacent streams by periodical overflows. Their fauna is therefore substantially that of the rivers themselves, and the two should, of course, be studied together.
They are probably in all cases either parts of former river channels, which have been cut off and abandoned by the current as the river changed its course, or else are tracts of the high-water beds of streams over which, for one reason or another, the periodical deposit of sediment has gone on less rapidly than over the surrounding area, and which have thus come to form depressions in the surface which retain the waters of overflow longer than the higher lands adjacent. Most of the numerous "horseshoe lakes" belong to the first of these varieties, and the "bluff-lakes," situated along the borders of the bottoms, are many of them examples of the second.
These fluviatile lakes are most important breeding grounds and reservoirs of life, especially as they are protected from the filth and poison of towns and manufactories by which the running waters of the state are yearly more deeply defiled.
The amount and variety of animal life contained in them as well as in the streams related to them is extremely variable, depending chiefly on the frequency, extent, and duration of the spring and summer overflows. This is, in fact, the characteristic and peculiar feature of life in these waters. There is perhaps no better illustration of the methods by which the flexible system of organic life adapts itself, without injury, to widely and rapidly fluctuating conditions. Whenever the waters of the river remain for a long time far beyond their banks, the breeding grounds of fishes and other animals are immensely extended, and their food supplies increased to a corresponding degree. The slow or stagnant backwaters of such an overflow afford the best situations possible for the development of myriads of Entomostraca, which furnish, in turn, abundant food for young fishes of all descriptions. There thus results an outpouring of life—an extraordinary multiplication of nearly every species, most prompt and rapid, generally speaking, in such as have the highest reproductive rate, that is to say, in those which produce the largest average number of eggs and young for each adult.
The first to feel this tremendous impulse are the protophytes and Protozoa, upon which most of the Entomostraca and certain minute insect larvae depend for food. This sudden development of their food resources causes, of course, a corresponding increase in the numbers of the latter classes, and, through them, of all sorts of fishes. The first rishes to feel the force of this tidal wave of life are the rapidly-breeding, non-predaceous kinds; and the last, the game fishes, which derive from the others their principal food supplies. Evidently each of these classes must act as a check upon the one preceding it. The development of animalcules is arrested and soon sent back below its highest point by the consequent development of Entomostraca; the latter, again, are met, checked, and reduced in number by the innumerable shoals of fishes with which the water speedily swarms. In this way a general adjustment of numbers to the new conditions would finally be reached spontaneously; but long before any such settled balance can be established, often of course before the full effect of this upward influence has been exhibited, a new cause of disturbance intervenes in the disappearance of the overflow. As the waters retire, the lakes are again defined; the teeming life which they contain is restricted within daily narrower bounds, and a fearful slaughter follows; the lower and more defenceless animals are penned up more and more closely with their predaceous enemies, and these thrive for a time to an extraordinary degree. To trace the further consequences of this oscillation would take me too far. Enough has been said to illustrate the general idea that the life of waters subject to periodical expansions of considerable duration, is peculiarly unstable and nuctuating; mat eacn species swings, pendulum-like but irregularly, between a highest and a lowest point, and that this fluctuation affects the different classes successively, in the order of their dependence upon each other for food.
Where a water-shed is a nearly level plateau with slight irregularities of the surface many of these will probably be imperfectly drained, and the accumulating waters will form either marshes or lakes according to the depth of the depressions. Highland marshes of this character are seen in Ford, Livingston, and adjacent counties,* between the headwaters of the Illinois and Wabash systems; and an area of water-shed lakes occurs in Lake and McHenry counties, in northern Illinois.
The latter region is everywhere broken by low, irregular ridges of glacial drift, with no rock but boulders anywhere in sight. The intervening hollows are of every variety, from mere sink-holes, either dry or occupied by ponds, to expanses of several square miles, forming marshes or lakes.
This is, in fact, the southern end of a broad lake belt which borders Lakes Michigan and Superior on the west and south, extending through eastern and northern Wisconsin and northwestern Minnesota, and occupying the plateau which separates the headwaters of the St. Lawrence from those of the Mississippi. These lakes are of glacial origin, some filling beds excavated in the solid rock, and others collecting the surface waters in hollows of the drift. The latter class, to which all the Illinois lakes belong, may lie either parallel to the line of glacial action, occupying valleys between adjacent lateral moraines, or transverse to that line and bounded by terminal moraines. Those of our own state all drain at present into the Illinois through the Des Plaines and Fox; but as the terraces around their borders indicate a former water-level considerably higher than the present one it is likely that some of them once emptied eastward into Lake Michigan. Several of these lakes are clear and beautiful sheets of water, with sandy or gravelly beaches, and shores bold and broken enough to relieve them from monotony. Sportsmen long ago discovered their advantages and club-houses and places of summer resort are numerous on the borders of the most attractive and easily accessible. They offer also an unusually rich field to the naturalist, and their zoology and botany should be better known.
The conditions of aquatic life are here in marked contrast to those afforded by the fluviatile lakes already mentioned. Connected with each other or with adjacent streams only by slender rivulets, varying but little in level with the change of the season and scarcely at all from year to year, they are characterized by an isolation, independence, and uniformity which can be found nowhere else within our limits.
Among these Illinois lakes I did considerable work during October of two successive years, using the sounding line, deep-sea thermometer, towing net, dredge, and trawl in six lakes of northern Illinois, and in Geneva Lake, Wisconsin, just across the line. Upon one of these Illinois lakes I spent a week in October, and an assistant, Prof. H. Garman, now of the University, spent two more, making as thorough a physical and zoological survey of this lake as was possible at that season of the year.
I now propose to give you in this paper a brief general account of the physical characters and the fauna of these lakes, and of the relations of the one to the other; to compare, in a general way, the animal assemblages which they contain with those of Lake Michigan –where also I did some weeks of active aquatic work in 1881—and with those of the fluviatile lakes of central Illinois; to make some similar comparisons with the lakes of Europe; and, finally, to reach the subject which has given the title to this paper—to study the system of natural interactions by which this mere collocation of plants and animals has been organized as a stable and prosperous community.
First let us endeavor to form the mental picture. To make this more graphic and true to the facts, I will describe to you some typical lakes among those in which we worked; and will then do what I can to furnish you the materials for a picture of the life that swims and creeps and crawls and burrows and climbs through the water, in and on the bottom, and among the feathery water-plants with which large areas of these lakes are filled.
Fox Lake, in the western border of Lake county, lies in the form of a broad irregular crescent, truncate at the ends, and with the concavity of the crescent to the northwest. The northern end is broadest and communicates with Petite Lake. Two points projecting inward from the southern shore form three broad bays. The western end opens into Nippisink Lake, Crab Island separating the two. Fox River enters the lake from the north, just eastward of this island, and flows directly through the Nippisink. The length of a curved line extending through the central part of this lake, from end to end, is very nearly three miles, and the width of the widest part is about a mile and a quarter. The shores are bold, broken, and wooded, except to the north, where they are marshy and flat. All the northern and eastern part of the lake was visibly shallow—covered with weeds and feeding waterfowl, and I made no soundings there. The water there was probably nowhere more than two fathoms in depth, and over most of that area was doubtless under one and a half. In the western part, five lines of soundings were run, four of them radiating from Lippincott's Point, and the fifth crossing three of these nearly at right angles. The deepest water was found in the middle of the mouth of the western bay, where a small area of five fathoms occurs. On the line running northeast from the Point, not more than one and three fourths fathoms is found. The bottom at a short distance from the shores was everywhere a soft, deep mud. Four hauls of the dredge were made in the western bay, and the surface net was dragged about a mile.
Long Lake differs from this especially in its isolation, and in its smaller size. It is about a mile and a half in length by a mile in breadth. Its banks are all bold except at the western end, where a marshy valley traversed by a small creek connects it with Fox Lake, at a distance of about two miles. The deepest sounding made was six and a half fathoms, while the average depth of the deepest part of the bed was about five fathoms.
Cedar Lake, upon which we spent a fortnight, is a pretty sheet of water, the head of a chain of six lakes which open finally into the Fox. It is about a mile in greatest diameter in each direction, with a small but charming island bank near the center, covered with bushes and vines—a favorite home of birds and wild flowers. The shores vary from rolling to bluffy except for a narrow strip of marsh through which the outlet passes, and the bottoms and margins are gravel, sand, and mud in different parts of its area. Much of the lake is shallow and full of water plants; but the southern part reaches a depth of fifty feet a short distance from the eastern bluff.
Deep Lake, the second of this chain, is of similar character, with a greatest depth of fifty-seven feet—the deepest sounding we made in these smaller lakes of Illinois. In these two lakes several temperatures were taken with a differential thermometer. In Deep Lake, for example, at fifty-seven feet I found the bottom temperature 53½— about that of ordinary well-water—when the air was 63° ; and in Cedar Lake, at forty-eight feet, the bottom was 58° when the air was 61°.
Geneva Lake, Wisconsin, is a clear and beautiful body of water about eight miles long by one and a quarter in greatest width. The banks are all high, rolling, and wooded, except at the eastern end, where its outlet rises. Its deepest water is found in its western third, where it reaches a depth of twenty-three fathoms. I made here, early in November, twelve hauls of the dredge and three of the trawl, aggregating about three miles in length, so distributed in distance and depth as to give a good idea of the invertebrate life of the lake at that season.
And now if you will kindly let this suffice for the background or setting of the picture of lacustrine life which I have undertaken to give you, I will next endeavor—not to paint in the picture; for that I have not the artistic skill. I will confine myself to the humble and safer task of supplying you the pigments, leaving it to your own constructive imaginations to put them on the canvas.
When one sees acres of the shallower water black with water-fowl, and so clogged with weeds that a boat can scarcely be pushed through the mass; when, lifting a handful of the latter, he finds them covered with shells and alive with small crustaceans; and then, dragging a towing net for a few minutes, finds it lined with myriads of diatoms and other microscopic algae, and with multitudes of Entomostraca, he is likely to infer that these waters are everywhere swarming with life, from top to bottom and from shore to shore. If, however, he will haul a dredge for an hour or so in the deepest water he can find, he will invariably discover an area singularly barren of both plant and animal life, yielding scarcely anything but a small bivalve mollusk, a few low worms, and red larvae of gnats. These inhabit a black, deep, and almost impalpable mud or ooze, too soft and unstable to afford foothold to plants even if the lake is shallow enough to admit a sufficient quantity of light to its bottom to support vegetation. It is doubtless to this character of the bottom that the barrenness of the interior parts of these lakes is due; and this again is caused by the selective influence of gravity upon the mud and detritus washed down by rains. The heaviest and coarsest of this material necessarily settles nearest the margin, and only the finest silt reaches the remotest parts of the lakes, which, filling most slowly, remain, of course, the deepest. This ooze consists very largely, also, of a fine organic debris. The superficial part of it contains scarcely any sand, but has a greasy feel and rubs away, almost to nothing, between the fingers. The largest lakes are not therefore, as a rule, by any means the most prolific of life, but this shades inward rapidly from the shore, and becomes at no great distance almost as simple and scanty as that of a desert.
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Table of Contents
Part One - Foundational Papers
Defining Ecology as a Science
Sharon E. Kingsland
1. Stephen A. Forbes (1887)
The Lake as a Microcosm
(Bulletin of the Peoria Scientific Association, pp. 77-87. Reprinted in the Bulletin of the Illinois State Natural History Survey 15 (1925): 537-50
2. Henry Chandler Cowles (1899)
The Ecological Relations of the Vegetation on the Sand Dunes of Lake Michigan
The Botanical Gazette 27 : 97-117, 167-202, 281-308, 361-91
3. Frederic E. Clements (1936)
Nature and Structure of the Climax
The Journal of Ecology 24 : 252-84
4. H. A. Gleason (1926)
The Individualistic Concept of the Plant Association
Bulletin of the Torrey Botanical Club 53 : 7-26
5. Joseph Grinnell (1917)
The Niche-Relationships of the California Thrasher
The Auk 34 : 427-33
6. A. J. Nicholson and V. A. Bailey (1935)
The Balance of Animal Populations, Part I
Proceeding of the Zoological Society, London, no. 3, pp. 551-98
Part Two - Theoretical Advances
The Role of Theory in the Rise of Modern Ecology
Leslie A. Real and Simon A. Levin
8. Frank W. Preston (1962)
The Canonical Distribution of Commonness and Rarity, Part I
Ecology 43 : 185-215, 431-32
9. G. Evelyn Hutchinson (1957)
Population Studies: Animal Ecology and Demography. Cold Spring Harbor Symposia on Quantitative Biology 22 : 415-27
10. Lamont C. Cole (1954)
The Population Consequences of Life History Phenomena
The Quarterly Review of Biology 29 : 103-37
11. Robert M. May (1974)
Biological Populations with Non-Overlapping Generations: Stable Points, Stable Cycles, and Chaos
Science 186 : 645-47
12. Robert H. MacArthur and Eric R. Pianka (1966)
On Optimal Use of a Patchy Environment
The American Naturalist 100 : 603-9
13. Vito Volterra (1926)
Fluctuations in the Abundance of a Species Considered Mathematically
Nature 118 : 558-60
14. J. G. Skellam (1951)
Random Dispersal in Theoretical Populations
Biometrika 38 : 196-218
Part Three - Theses, Antitheses, and Syntheses
Conversational Biology and Ecological Debate
Joel G. Kingsolver and Robert T. Paine
15. A. G. Tansley (1935)
The Use and Abuse of Vegetational Concepts and Terms
Ecology 16 : 284-307
16. G. E. Hutchinson (1959)
Homage to Santa Rosalia; or, Why Are There So Many Kinds of Animals?
The American Naturalist 93 : 145-59
17. Nelson G. Hairston, Frederick E. Smith, and Lawrence B. Slobodkin (1960)
Community Structure, Population Control, and Competition
The American Naturalist 94 : 421-25
18. Paul R. Ehrlich and Peter H. Raven (1964)
Butterflies and Plants: A Study in Coevolution
Evolution 18 : 586-608
19. J. L. Harper (1967)
A Darwinian Approach to Plant Ecology
The Journal of Ecology 55 : 247-70
20. Thomas W. Schoener (1971)
Theory of Feeding Strategies
Annual Review of Ecology and Systematics 2 : 369-404
Part Four - Methodological Advances
New Approaches and Methods in Ecology
James H. Brown
21. Lennart von Post (1967 )
Forest Tree Pollen in South Swedish Peat Bog Deposits
Pollen et Spores 9 : 378-401. A translation by Margaret Bryan Davis and Knut Faegri of Om skogstradspollen i sydsvenska torfmosselagerfolijder (foredragsreferat) (Geolgiska Foereningen i Stockholm. Foerhandlingar 38 : 384-34), with an introduction by Knut Faegri and Johs. Iversen
22. P. H. Leslie (1945)
On the Use of Matrices in Certain Population Mathematics
Biometrika 33 : 183-212
23. L. C. Birch (1948)
The Intrinsic Rate of Natural Increase of an Insect Population
The Journal of Animal Ecology 17 : 15-26
24. C. S. Holling (1959)
The Components of Predation as Revealed by a Study of Small Mammal Predation of the European Pine Sawfly
The Canadian Entomologist 91 : 293-320
25. Warren P. Porter and David M. Gates (1969)
Thermodynamic Equilibria of Animals with Environment
Ecological Monographs 39 : 227-44
26. J. Roger Bray and J. T. Curtis (1957)
An Ordination of the Upland Forest Communities of Southern Wisconsin
Ecological Monographs 27 : 325-49
27. Eugene P. Odum (1969)
The Strategy of Ecosystem Development
Science 164 : 262-70
Part Five - Case Studies in Natural Systems
Lessons from Nature: Case Studies in Natural Systems
Robert K. Peet
28. J. Davidson and H. G. Andrewartha (1948)
The Influence of Rainfall, Evaporation and Atmospheric Temperature on Fluctuations in the Size of a Natural Population of Thrips Imaginis (Thysanoptera)
The Journal of Animal Ecology 17 : 200-222
29. John M. Teal (1962)
Energy Flow in the Salt Marsh Ecosystem of Georgia
Ecology 43 : 614-24
30. Margaret B. Davis (1969)
Climatic Changes in Southern Connecticut Recorded by Pollen Desposition at Rogers Lake
Ecology 50 : 409-22
31. Alex S. Watt (1947)
Pattern and Process in the Plant Community
The Journal of Ecology 35 : 1-22
32. Robert H. MacArthur (1958)
Population Ecology of Some Warblers of Northeastern Coniferous Forests
Ecology 39 : 599-619
33. John Langdon Brooks and Stanley I. Dodson (1965)
Predation, Body Size, and Composition of Plankton
Science 150 : 28-35
Part Six - Experimental Manipulations in Lab and Field Systems
Manipulative Experiments as Tests of Ecological Theory
Jane Lubchenco and Leslie A. Real
34. H. B. D. Kettlewell (1955)
Selection Experiments on Industrial Melanism in the Lepidoptera
Heredity 9 :323-42
35. Thomas Park (1948)
Experimental Studies of Interspecies Competition. I. Competition between Populations of the Flour Beetles, Tribolium confusum Duvall and Tribolium castaneum Herbst
Ecological Monographs 18 : 267-307
36. C. B. Huffaker (1958)
Experimental Studies on Predation: Dispersion Factors and Predator-Prey Oscillations
Hilgardia 27 : 343-83
37. Joseph H. Connell (1961)
The Influence of Interspecific Competition and Other Factors on the Distribution of the Barnacle Chthamalus stellatus
Ecology 42 : 710-23
38. Robert T. Paine (1966)
Food Web Complexity and Species Diversity
The American Naturalist 100 : 65-75
39. Daniel S. Simberloff and Edward O. Wilson (1969)
Experimental Zoogeography of Islands: The Colonization of Empty Islands
Ecology 50 : 278-96
40. Gene E. Likens, F. Herbert Bormann, Noye M. Johnson, D. W. Fisher, and Robert S. Pierce (1970)
Effects of Forest Cutting and Herbicide Treatment on Nutrient Budgets in the Hubbard Brook Watershed-Ecosystem
Ecological Monographs 40 : 23-47
List of Contributors