Chapter II: Preface: v (1)

CHAPTER

I. AIM, CONTENT, AND POINT OF VIEW 1

II. THE VALUE AND METHOD OF ECOLOGICAL SURVEYS 23

III. FIELD STUDY 36

IV. THE COLLECTION, PRESERVATION, AND DETERMINATION OF SPECIMENS 49

V. REFERENCES TO SCIENTIFIC TECHNIQUE 55

1. The Scientific Method 55

2. Directions for Collecting and Preserving Specimens,
Photographing, Surveying, and Other Phases
of Technique 57

3. The Preparation of Papers for Publication and
on Proof Reading 65

VI. REFERENCES TO IMPORTANT SOURCES OF INFORMATION ON THE LIFE HISTORIES AND HABITS OF INSECTS AND ALLIED INVERTEBRATES 70

VII. THE LAWS OF ENVIRONMENTAL CHANGE, OR THE “ORDERLY SEQUENCE OF EXTERNAL NATURE.”(THE DYNAMIC OR PROCESS RELATIONS OF THE ENVIRONMENT) 79

VIII. THE LAWS OF ORDERLY SEQUENCE OF METABOLISM, GROWTH, DEVELOPMENT, PHYSIOLOGICAL CONDITIONS, AND BEHAVIOR, OR “THE LIVING ORGANISM AND THE CHANGES WHICH TAKE PLACE IN IT.” (THE DYNAMIC OR PROCESS RELATIONS OF THE ANIMAL) 92

1. General Physiology and Development 95

2. A Selection of Physiological and Ecological Papers 100

3. Animal Behavior as a Process 102

4. A List of Selected Reviews and Bibliographies 107

5. A Selection of References on Life Histories and
Behavior 108

IX. THE CONTINUOUS PROCESS OF ADJUSTMENT BETWEEN THE ENVIRONMENT AND THE ANIMAL, WITH SPECIAL REFERENCE TO OTHER ORGANISMS. (THE DYNAMIC OR PROCESS RELATIONS OF ANIMAL ASSOCIATIONS AND AGGREGATIONS) 122

1. The Struggle for Existence 123

2. The Dynamic Relations of Associations and Aggregations,
with Special Reference to Animal Associations 130

_a._ The Relation of Animals to Pollination and to
Plant Galls 141

_b._ Subterranean and Cave Associations 143

_c._ Selected References on Aggregations and Associations 145

INDEX 151

INDEX TO NAMES 179

LIST OF FIGURES

FIG.
1. An Oyster Habitat on the South Carolina Coast _Frontispiece_

2. Struggle for Existence on a Clam Flat. Overcrowded
Condition _facing page_ 7

3. Struggle for Existence on a Clam Flat. Destruction
of Clams by Predaceous Animals ” ” 8

4. A Small Mountain Stream as an Animal Environment ” ” 86

5. A Small River as an Animal Environment ” ” 87

6. An Illinois Prairie Remnant as an Animal Environment ” ” 139

7. An Illinois Deciduous Forest as an Animal Environment ” ” 146

RELATION OF ECOLOGY TO BIOLOGY

“I shall try to show that life is response to the order of nature.... But if it be admitted, it follows that biology is the study of response, and that the study of that order of nature to which response is made is as well within its province as the study of the living organism which responds, for all the knowledge we can get of both these aspects of nature is needed as a preparation for the study of that relation between them which constitutes life.”

“To study life we must consider three things:

_first_, the orderly sequence of external nature;

_second_, the living organism and the changes which take place in it; and,

_third_, that continuous adjustment between the two sets of phenomena which constitutes life.”

“The physical sciences deal with the external world, and in the laboratory we study the structure and activities of organisms by very similar methods; but if we stop there, neglecting the relation of the living being to its environment, our study is not biology or the science of life.”

W. K. BROOKS.

ANIMAL ECOLOGY

I. AIM, CONTENT, AND POINT OF VIEW

Ecology has no aim, but ecologists have. The problems of the ecologist are not fundamentally different from those of any other kind of naturalist. The superficial differences in aim are due to the different points of view, or methods of approach, rather than to any essential difference in the character of the problems.

The essentially biological core of ecology may be best shown by considering the relation which this science bears to other branches of biology, a relation which has been admirably expressed by the eminent physiologist, Burdon-Sanderson (’94, pp. 438-439), as follows:

“Now the first thing that strikes us in beginning to think about the activities of an organism is that they are naturally distinguishable into two kinds, according as we consider the action of the whole organism in its relation to the external world or to other organisms, or the action of the parts or organs in their relation to each other. The distinction to which we are thus led between the _internal_ and _external_ relation of plants and animals has of course always existed, but has only lately come into such prominence that it divides biologists more or less completely into two camps--on the one hand those who make it their aim to investigate the actions of the organism and its parts by the accepted methods of physics and chemistry, carrying this investigation as far as the conditions under which each process manifests itself will permit; on the other, those who interest themselves rather in considering the place which each organism occupies, and the part which it plays in the economy of nature. It is apparent that the two lines of inquiry, although they equally relate to what the organism _does_, rather than to what it is, and therefore both have equal right to be included in the one great science of life, or biology, yet lead in directions which are scarcely even parallel. So marked, indeed, is the distinction, that Professor Haeckel some twenty years ago proposed to separate the study of organisms with reference to their place in nature under the designation of ‘œcology,’ defining it as comprising ‘the relations of the animal to its organic as well as to its inorganic environment, particularly its friendly or hostile relations to those animals or plants with which it comes into direct contact.’[1] Whether this term expresses it or not, the distinction is a fundamental one. Whether with the œcologist we regard the organism in relation to the world, or with the physiologist as a wonderful complex of vital energies, the two branches have this in common, that both studies fix their attention, not on stuffed animals, butterflies in cases, or even microscopical sections of the animal or plant body--all of which relate to the framework of life--but on life itself.”

[1] These he identifies with “those complicated mutual relations which Darwin designates as conditions of the struggle for existence.” Along with chorology--the distribution of animals--œcology constitutes what he calls _Relations-physiologie_. Haeckel, “Entwickelungsgang u. Aufgaben der Zoologie,” Jenaische Zeitschr., 1869, Vol. V, p. 353.

The quotations from Brooks, on a preceding page, show even more explicitly the intimate relation which exists between biology and ecology. At first glance they may seem to prove almost too much--that biology and ecology are synonymous. They show at least that ecology is concerned with fundamental biological problems--the responses of organisms to their complete environments.

The relations which different branches of ecology bear to one another may be discussed under three headings, individual, aggregate, and associational ecology. These phases are superficially so distinct that students of one branch may be almost unaware of the existence of the coördinate branches and may not realize that each is a part of the larger unit.

_Individual Ecology._--The study of individual ecology is the investigation of the development (_process_ of formation) of the structure, function, and behavior of a given individual or kind of animal from the standpoint of its relations and responses to the complete environment. All ecologically significant facts should be considered. Such a study may be devoted to an animal, as, for example, a bumblebee, a crawfish, or a garter snake, and may be limited to a single habitat or locality, or extended throughout the entire geographic range of an animal. From this standpoint the individual studied becomes the hub of the microcosm, from which all relations and responses radiate. Most of the physiological studies of ecological bearing and many investigations of animal behavior have been made from this viewpoint. The organism is thus considered as an agent which, transforming and utilizing substance and energy, produces a varied number of physiological conditions and forms of activity, which in turn furnish the basis for the constant process of response between the organism and its environment.

_Aggregate Ecology._--The study of aggregate ecology is the investigation of the ecological development, relations, and responses of animals based upon hereditary or taxonomic units, as in a family community, or in genera, families, orders, etc. These groups or aggregates are made the basis for the ecologic study, as a hive of bees, birds, dragon flies (Odonata), the genus _Bombus_. From this approach the activities and responses of the group are traced throughout all environments and associations within the area studied, or throughout the world, and its responses and adjustments to the whole environment receive primary attention. The hereditary or taxonomic unit is here the hub of the microcosm. Perhaps most of the contributions to ecology by the taxonomists are made from this standpoint. Here also the aggregate is considered as an agent or entity which produces many kinds of activities and adjustments to the environment.

_Associational Ecology._--Associational ecology is devoted to the investigation of the development, interrelations, and responses of animals which are grouped or associated in the same habitats and environments. In this case the associates in a given association and habitat are considered as a unit, whose activities and interrelations and responses are investigated in the same manner as if it were a single animal. The interactions among members of an association are to be compared to the similar relations existing between the different cells, organs, or activities of a single individual. Such groupings have a composition which has developed into an arrangement, or “spacing,” of individuals within it, and which produces a particular plan or pattern, as a result of the innumerable responsive activities on the part of the individuals which live together. For example, when the animals living in a small brook, the littoral zone of a lake, in a colony of breeding gulls, or on the floor of a forest, are treated as a unit, the entire history of the animals in the habitat is considered as a response to the conditions of life.

In this form of study the association becomes the center of all radiating relations and responses. Such an association is an agent which transforms substance and energy, producing varied physiological conditions and responses in the continuous process of adjustment “which constitutes life.” The physiological needs and states of an association have as real existence in individual animals as have similar needs in the cell or cells which compose the animal body. The mere statement of the facts of such relations is enough to make valid such a comparison.

For the associational aspect of ecology the German naturalist, Möbius, proposed in 1877 the term “biocœnosis.” The meaning of this he expressed very clearly and concisely, and on account of its relatively obscure publication, in a paper devoted to oyster culture, it has not gained the circulation among zoölogists which its importance merits. His statement (Möbius, ’83, p. 723) is as follows:

“Every oyster-bed is thus, to a certain degree, a community of living beings, a collection of species, and a massing of individuals, which find here everything necessary for their growth and continuance, such as suitable soil, sufficient food, the requisite percentage of salt, and a temperature favorable to their development. Each species which lives here is represented by the greatest number of individuals which can grow to maturity subject to the conditions which surround them, for among all species the number of individuals which arrive at maturity at each breeding period is much smaller than the number of germs produced at that time. The total number of mature individuals of all the species living together in any region is the sum of the survivors of all the germs which have been produced at all past breeding or brood periods; and this sum of matured germs represents a certain quantum of life which enters into a certain number of individuals, and which, as does all life, gains permanence by means of transmission. Science possesses, as yet, no word by which such a community of living beings may be designated; no word for a community where the sum of species and individuals, beings mutually limited and selected under the average external conditions of life, have, by means of transmission, continued in possession of a certain definite territory. I propose the word _Biocænosis_[2] for such a community. Any change in any of the relative factors of a biocönose produces changes in other factors of the same. If, at any time, one of the external conditions of life should deviate for a long time from its ordinary mean, the entire biocönose, or community, would be transformed. It would also be transformed, if the number of individuals of a particular species increased or diminished through the instrumentality of man, or if one species entirely disappeared from, or a new species entered into, the community.” (See Figure 1).

[2] From βίος, life, and κοινόειν, to have something in common.

FIG. 2.--Struggle for Existence on a Clam Flat. Showing the overcrowded condition of a colony of Soft Clams (_Mya arenaria_) on Rowley Reef, Massachusetts. The pits mark the position of the living clams. Photo. by Belding, loaned by Mass. Comms. Fisheries and Game. ]

The three methods of approach to ecological study are not so distinct as they appear at first thought. With perfecting knowledge the network of interrelations increases and the paths converge. Then also the study of the individual behavior of “social” animals, as ants, white ants, bees, or birds which live and breed in colonies, shows transitional stages from the individual unit to that of the family, the colony, and on to the association. Yet the advantage of each point of view should be recognized as an aid in the analysis and synthesis of any problem.

Some students feel that the study of individual ecology should precede that of the associational. Within certain limits this is true, but if our general knowledge of biology had waited for the perfection of our knowledge of the individual cells of animals, the results would have been disastrous to all concerned. Even now our knowledge of these subjects is very incomplete. For similar reasons there should be no delay in studying animal aggregates and associations.

A combination of ecological and taxonomic study generally appeals most strongly to those students who have made a specialty of some group of animals. They are familiar with certain forms, have some confidence in taxonomic methods, and frequently have given some attention to habits, life histories, and to collecting. To those who like the descriptive aspect of taxonomy, ecological studies also offer a new field for further description and classification. At present perhaps the majority of ecological students have entered the subject through taxonomy. It is the almost universal verdict of such students that it has required much effort on their part to make the change in the point of view. Such a change cannot be made by a simple resolve, but requires a modification of the habits of the mind, which will be attended by a distinct consciousness of effort. As in other habits, reversion to the older attitude of mind is very easy. This change in point of view is a problem in habit formation, a study of the mental behavior of the ecologist, which is in reality the main topic thus far discussed. One may attempt to make such a change and find that he does not have sufficient modifiability to make it permanent, so that it is only for the moment, during actual collision with some stubborn fact, that he is able to realize ecological relations and an ecological point of view.

FIG. 3.--Struggle for Existence on a Clam Flat. Showing the destruction of a colony of Soft Clams by Horse-shoe Crabs (_Limulus_) and Cockles (_Lunatia_), Rowley Reef, Massachusetts. Photo. by Belding, loaned by Mass. Comms. Fisheries and Game. ]

To the physiologist, however, individual ecology tends to appeal most strongly, and he, perhaps on account of the preponderance of analytical methods in his work, feels that this is the safest and most important aspect. This statement is perhaps also true of most students of animal behavior. This is largely due to the great present need of analytical methods in these lines, and perhaps indicates a stage in the development of their science rather than a permanent condition. Later a synthetic development will probably become more prominent, and with it will come a change in estimating relative values. Generally physiologists allow for a greater influence of the environment than do many other students. They are impressed with the dependence of organisms upon their environment, and the study of their reactions only reinforces this conception.

The ecologist who studies the responses of animals cannot help being impressed with the processes of adjustment, and with adaptation as a _process_. It is adaptation as a process, rather than as a product, which perhaps interests him most, and emphasis needs to be placed upon this distinction. The problem of adaptation as a process may be a different and separate one from that of evolution, but individual animals must have shown adjustive adaptation, or there could have been no perpetuation to continue the struggle of adjustment. Ecological problems are likely to raise a question as to the relative importance of adaptation and evolution--if they are separate problems. The present generation has perhaps been more deeply impressed by evolution as a process, than by adaptation as a process.

The ecology of living animals is only the latest chapter in the volume on this subject; the preceding chapters will contain a history of the indefinitely long series of ecological responses which have taken place in the geologic past. Here is where the ecologist and paleontologist and geologist find common ground. The ecology of living animals must furnish us with whatever firm basis we have for the interpretation of the conditions of life in the past, upon which the paleontologist, stratigrapher, or paleogeographer must depend, at least in part, for his interpretations.

With still another training and interest, as in the case of those especially interested in human affairs, such as the sociologist, the physician, the sanitary expert, and the agriculturist, we may ultimately expect a greater appreciation for the associational aspect because of the social or associational character of human society. The associational is the phase of animal activity which may be considered as the form of animal behavior which has developed into the human social relations. It is a response to the complete organic and inorganic environment.

It is rather natural that in a relatively newly recognized subject like ecology this human aspect has not been very fully discussed. For practical reasons the ecology of man has been developed largely independent of that of animals; just as human physiology and psychology have been developed relatively independent of comparative or general physiological psychology. To the mutual advantage of these subjects they are now rapidly converging, and we may anticipate a similar relation between general animal ecology and the ecology of man. In a general treatise on animal ecology the human phase should not receive undue emphasis any more than it should in a general physiology of animals or in a comparative psychology. But, nevertheless, the relationships of man and his animal associates (slaves, domestic animals, rats, mice, parasites, etc.) form as truly an animal association as do those of the animals which live associated in some forest glade; and in all probability, before any approximately complete understanding can be had of the human associations, their roots and principles of activity must be known and understood in the less aristocratic portion of his animal relatives.

The recognition of the associational aspect of ecology, as well as that human ecology is a part of general animal ecology, is of recent origin. This is very well shown in the following quotation from Huxley (1854. On the Educational Value of the Natural History Sciences):

“Biology deals only with living beings as isolated things--treats only of the life of the individual: but there is a higher division of science still, which considers living beings as aggregates--which deals with the relation of living beings one to another--the science which _observes_ men--whose _experiments_ are made by nations one upon another, in battlefields--whose _general propositions_ are embodied in history, morality, and religion--whose _deductions_ lead to our happiness or our misery--and whose _verifications_ so often come too late, and serve only

‘To point a moral, or adorn a tale’--

I mean the science of Society or _Sociology_.”

At a later date (1876. On the Study of Biology) Huxley says: “For whatever view we may entertain about the nature of man, one thing is perfectly certain, that he is a living creature. Hence, if our definition is to be interpreted strictly, we must include man and all his ways and works under the head of Biology; in which case, we should find that psychology, politics, and political economy would be absorbed into the province of Biology. In strict logic no one can object to this course.... The real fact is that we biologists are a self-sacrificing people ... [so that] we feel that we have more than sufficient territory.... But I should like you to recollect that that is a sacrifice, and that you should not be surprised if it occasionally happens that you see a biologist apparently trespassing in the region of philosophy or politics; or meddling with human education; because, after all, that is a part of his kingdom which he has only voluntarily forsaken.”

Whether sociology is regarded as a response of man to his fellows or to the whole of his environment is inconsequential in its bearing upon whether or not it is ecological. The response of man, as an animal, to a part or the whole of his environment is strictly ecological. Huxley recognized one relation very clearly, and that is that the ecological relations of _individuals_ do not currently include the higher synthesis which deals with them as associations, or “aggregates” as he terms them. So far as known to the writer, human activities in general have never been fully and comprehensively oriented from the ecological standpoint, even by the humanitarians themselves, although some important preliminary steps have been taken. It looks as if such a viewpoint might give a new unity to all studies of human relations.

There is still another class of persons, particularly teachers and isolated students, who desire first of all to understand and interpret their own vicinity, and who will inquire which of the three plans their work best fits. If such a one begins with the detailed study of each species, the general survey will not be completed during his lifetime. If he uses the larger taxonomic units, he may survey the field by going over the same ground again and again, with each of the different groups successively in mind, until the entire field has been surveyed. Or, lastly, he may divide the area into associations and study the animals which are found living together, and by studying one association after another he may cover the entire field. A teacher will find certain important advantages in this plan, and certain disadvantages. One of the most important considerations in its favor is that such a study results in a familiarity with the kinds of animals one actually finds in natural groups, as when his class is on an excursion. The natural history which a farmer, a fisherman, a summer vacationist, or a sportsman acquires is grouped in this same manner. Thus to a large number of people this is the natural method of approach, and is generally of most permanent value, except possibly to some professional teachers or zoölogists. One of its greatest disadvantages is that in most of the literature which one must use, the animals are not grouped in this way, but taxinomically.

The individual, aggregate, and associational methods of study are in themselves subject to diverse angles of approach, and each has its particular advantages and disadvantages. Of the methods of approach mention will be made of three only, the descriptive, the comparative, and the genetic or method of processes. The descriptive method must develop to some degree before the genetic problems can be adequately stated, and the mature development of the genetic may, and generally does, lag far behind that of the descriptive. The reason for this is simple, for it is evident that it is much easier to describe what we see than it is to explain how it originated or its process of development. At present biology as a science is mainly in the descriptive stage, though it is slowly but surely becoming explanatory and genetic. The developmental or explanatory method is so difficult that every possible expedient--observation, comparison, reflection, experiment, etc.--must be used to secure the proper development of the main phases of ecology. There is a marked tendency in the naturalist to master one system of work, as observation or experiment, and to use it as a tool almost exclusively, turning from one phase of the subject to another, and continuing the use of the same method. This way of working is favorable to a good technique, but its weakness is that it often tends to give its user a feeling of the great superiority and reliability of the result reached by his method, and a correspondingly less appreciative recognition of results secured by other methods. To observe, to experiment, to reflect, to dissect, to stain, and to collect are only partial methods of investigation, and this fact should be realized and be kept in mind when estimating values and planning work.

The aim of the ecologist is professedly genetic or explanatory because it is the study of _responses_ to all conditions of the complete environment. But these responses must be described, and the conditions influencing them as well, so that a descriptive aspect is an essential part in all phases of ecology. In the study of the responses of an individual, an order, or an association, pure description of the responses is necessary; but a description which will at once describe and show the working of the processes by which the results were produced, is of quite a different order. This phase of explanation has been most concisely expressed and applied by the students of the physical sciences, and biologists may profit much from a study of their methods.

When, however, we turn to the viewpoint of the development of the science of ecology as a whole, a symmetrical development of the subject is most desirable. The preponderating influence of any special point of view tends, like dominance in general, to smother or suppress other germinating and competing ideas. The different special interests each have their advantages and disadvantages, as does a general interest. Diversity in students leads to diversity in the development of the subject, and a variety of emotional appeals to the student has its advantages. And just as the special student should devote some attention to the general bearing of his work, so also should the student of the general aspects cultivate some special field of interest.

The preceding discussion of the aims and methods in ecological study has been intended to indicate some of its general bearings, and to give the student some idea of the tests or criteria which may be used to aid in steering his course through the maze of observations which he may make and the opinions which he encounters. It is of equal importance for the student to be able to perceive ecological relations as recorded by others, because one person’s experience is so limited compared with the general body of recorded fact and inference. Furthermore, there are also so many degrees and kinds of work that go by the name ecological, which may or may not be, and so many also which are truly ecological but which do not pass under that name, that it is necessary that the student shall be able to see through its diverse guises and recognize its essential character. Whenever the question arises as to the ecological character of a fact, inference, or conclusion, its ecological validity may be tested in the following way:

Do the facts, inferences, or conclusions show a _response_ to the inorganic or organic environment:

1. As an individual of a species or kind of animal?

2. As a group of taxonomically related animals?

3. As an association of interacting animals?

REFERENCES ON THE ECOLOGICAL STANDPOINT

In this I have listed only those papers which have seemed to me particularly significant because of their point of view, regardless of whether or not they are primarily zoölogical or specifically mention ecology.

BROOKS, W. K.

1899. The Foundations of Zoölogy. pp. 339. New York.

Introductory, pp. 1-29; Huxley, and the Problem of the Naturalist, pp. 33-46; Nature and Nurture, pp. 49-79.

1906. Heredity and Variation; Logical and Biological. Proc. Amer. Phil. Soc., Vol. XLV, pp. 70-76.

An extremely suggestive paper which should be read by every ecological student.

GANONG, W. F.

1907. The Organization of the Ecological Investigation of the Physiological Life-Histories of Plants. Bot. Gaz., Vol. XLIII, pp. 341-344.

1904. The Cardinal Principles of Ecology. Science, N. S., Vol. XIX, pp. 493-498.

BURDON-SANDERSON, J. S.

1894. Biology in Relation to Other Natural Sciences. Smithsonian Report for 1893, pp. 435-463.

MÖBIUS, K.

1877. Die Auster und die Austernwirthschaft. pp. 126. Berlin.

On page 72 he proposes the term “biocœnose” for the group of animals which live together in the same habitat. Not seen by writer.

1883. The Oyster and Oyster-Culture. Rep. U. S. Fish Comm., 1880, Part VIII, pp. 683-751.

Translation of preceding paper. On pp. 721-729 he discusses “An Oyster-Bank as a Biocönose, or a Social Community”; on page 723 he proposes the word “biocœnosis.” An illuminating paper.

DAHL, F.

1902. Die Ziele der vergleichenden “Ethologie” (d. i. Biologie im älteren engeren Sinne). Verh. V. Inter. Zoöl.-Cong. 1901, pp. 296-300.

1908. Grundsätze und Grundbegriffe der biocönotischen Forschung. Zool. Anz., Bd. XXXIII, pp. 349-353.

1898. Experimentell-statistische Ethologie. Verh. der Deutsch. Zool. Gesell. Bd., 1898, pp. 121-131.

1901. Was ist ein Experiment, was Statistik in der Ethologie? Biol. Centralbl., Bd. XXI, pp. 675-681.

WASMANN, E.

1901. Biologie oder Ethologie? Biol. Centralbl., Bd. XXI, pp. 391-400.

WHEELER, W. M.

1902. ‘Natural History,’ ‘Œcology’ or ‘Ethology’? Science, N. S., Vol. XV, pp. 971-976.

Advocates the use of the term ethology.

ST. HILAIRE, I. GEOFFRAY.

1859. Histoire Générale des Règnes Organiques, Vol. II.

Not seen by writer. Dr. W. M. Wheeler, of Harvard University, has kindly sent me the following note from p. 285. “‘It is to ethology therefore that the fourth part of this work is devoted, to which belongs the study of the relations of organisms within the family and the society in the aggregate and in the community.’ In a volume of the same work, page xx, St. Hilaire gives his program and speaks of the general facts belonging to ethological laws. These are defined as ‘relating to the instincts, habits and more generally to the external vital manifestations of organisms.’” About the preceding Dr. Wheeler remarks: “You see this covers precisely the field which was a few years later called ‘ecology’ by Haeckel. Apparently the part of the work in which St. Hilaire wished to give a detailed account of the ethological phenomena of animals was not published. Only three volumes of the work exist. He died November 10, 1861, without having completed the work.”

Thus ethology has priority over ecology, but to my mind this fact carries no special weight, particularly since the word has become current in botany. To use a different name for the same subject or process in botany and zoölogy is as undesirable as to use a different term for heredity in plants and in animals.

LANKESTER, E. R.

1889. Article “Zoölogy.” Ency. Britannica, 9th ed. Amer. Reprint. Vol. XXIV, pp. 842, 856.

Lankester defines “Bionomics.--The lore of the farmer, gardener, sportsman, fancier, and field naturalist, including thremmatology, or the science of breeding, and the allied teleology, or science of organic adaptations: exemplified by the patriarch Jacob, the poet Vergil, Sprengel, Kirby and Spence, Wallace, and Darwin.... Buffon (1707-1788) alone among the greater writers of the three past centuries emphasized that view of living things which we call ‘bionomics.’ Buffon deliberately opposed himself to the mere exposition of the structural resemblances and differences of animals, and, disregarding classification, devoted his treatise on natural history to a consideration of the habits of animals and their adaptations to their surroundings, whilst a special volume was devoted by him to the subject of reproduction.... Buffon is the only prominent writer who can be accorded historic rank in this study.”

As I have access to but few of Buffon’s writings, I quote the above. Bionomics is seen not to be synonymous with ecology, as defined by most students, although it includes much that is ecological. The chaotic and unorganized “lore of the farmer” has no unifying or guiding principles, and although it contains many facts, from which a science may be built, to call it science seems undesirable.

It is of course advantageous in some ways to have agreement as to the limitations of ecology, or any science, but even the more exact sciences seem to fare little better, as is shown by the following statement: “It is not long since I heard a university professor begin a lecture on physics somewhat in this way: ‘Physics is the science of matter and energy. This field is so large that it is customary at present to break off the physics of the molecule and its reactions and call it chemistry. Also to put to one side the physics of the heavenly bodies and call this a part of astronomy,’ etc.” (Strong, Science, N. S., Vol. XXXIV, p. 409, 1911.)

FORBES, S. A.

1895. On Contagious Disease in the Chinch-Bug (_Blissus leucopterus_ Say). 19th Rep. State Ent. Ill. (8th Rep. of S. A. Forbes), pp. 16-176.

In this paper Forbes defines (pp. 16-18) ecology and points out, I believe for the first time, that economic entomology is simply applied ecology. He says, “The study of œcology is thus to the economic entomologist what the study of physiology is to the physician.”

1909. Aspects of Progress in Economic Entomology. Journ. Econ. Ent., Vol. II, pp. 25-35.

Especially pp. 28-32 on the relation of ecology to economic entomology.

HERDMANN, W. A.

1896. Oceanography, Bionomics, and Aquiculture. Smithsonian Report for 1895, pp. 433-454.

EMERY, C.

1905. Éthologie, Phylogénie et Classification. C. R. 6me Cong. inter. de Zool. Berne, 1904, pp. 160-174.

CLEMENTS, F. E.

1905. The Foundations of Ecology, pp. 1-17.

Research Methods in Ecology, pp. 334. Lincoln, Nebraska.

ADAMS, CHAS. C.

1906. Introductory Note. An Ecological Survey in Northern Michigan. Ann. Rep. Mich. Geol. Surv. for 1905, pp. 11-12.

1909. The Ecological Succession of Birds. Ann. Rep. Mich. Geol. Surv. for 1908, pp. 121-154.

SHELFORD, V. E.

1912. Ecological Succession. V. Aspects of Physiological Classification. Biol. Bull., Vol. XXIII, pp. 331-370.

The standpoint of this paper is very much in harmony with that advanced in this book.

CASE, E. C.

1905. Œcological Features of Evolution. Bull. Wis. Nat. Hist. Soc., Vol. III, pp. 169-180.

WHEELER, W. M.

1905. Ethology and the Mutation Theory. Science, N. S., Vol. XXI, pp. 535-540.

FLAHAULT, C., and SCHRÖTER, C.

1910. Phytogeographical Nomenclature. Reports and Propositions. IIIᵉ Cong. Inter. de Botanique. Bruxelles, 1910. pp. 28 + x. Zurich.

WHITE, C. A.

1893. The Relation of Biology to Geological Investigation. Ann. Rep. U. S. Nat. Mus. for 1892, pp. 245-368.

This paper and the two following references illustrate the intimate relation of ecology to phases of geology.

GRABAU, A. W.

1899. The Relation of Marine Bionomy to Stratigraphy. Bull. Buffalo Soc. Nat. Sci., Vol. VI, pp. 319-367.

WALTHER, J.

1893-94. Einleitung in die Geologie als historische Wissenschaft. I. Bionomie des Meeres. II. Die Lebensweise der Meeresthiere. III. Lithogenesis der Gegenwart. Jena.

Shows the close relation between ecology and geology. The process standpoint is emphasized and the past is interpreted in terms of processes now in operation.

ADAMS, CHAS. C.

1908. Some of the Advantages of an Ecological Organization of a Natural History Museum. Proc. Amer. Associa. Museums, Vol. I, pp. 170-178.

II. THE VALUE AND METHOD OF ECOLOGICAL SURVEYS

“I cannot too strongly emphasize the fact ... that a comprehensive survey of our entire natural history is absolutely essential to a good _working knowledge_ of those parts of it which chiefly attract popular attention,--that is, its edible fishes, its injurious and beneficial insects, and its parasitic plants. Such a survey, however, should not stop with a study of the dead forms of nature, ending in mere lists and descriptions. To have an _applicable_ value, it must treat the life of the region as an organic unit, must study it _in action_, and direct principal attention to the laws of its activity.”--S. A. FORBES. 1883.

Natural history surveys have come down to us from the early days of zoölogy. These surveys have been of many kinds and have ranged from the adventurous accounts of early and daring explorers to those of such naturalists as Belt, Bates, Wallace, and Darwin, onward to the voluminous accounts of the “Biologia Centrali-Americana,” and in the _Challenger_ reports. These surveys have contributed greatly to our knowledge of the fundamental facts of zoölogy and to the training of naturalists.

The most frequent form of survey is that carried on along the lines which most nearly approach individual and aggregate ecology. Most of such surveys give only slight attention to the _responsive relation_, or only to its most general aspects. Surveys of the usual character are of great importance, and with students of taxonomic training and interests only, this form of survey occurs very naturally. Most of the governmental and state surveys and museum expeditions are developed along these lines. The frequency with which such methods are used in surveys, which are expected to produce economic results, indicates that these methods are generally considered the most satisfactory. The exceptions to this rule are mainly surveys of fresh and salt waters, and are related in some way to aquatic resources. Except when detailed individual studies of certain species or some special subject has been made, the usual form of the reports of such surveys is the annotated list. It is rarely that even brief chapters discuss the groupings of the animals as they are found associated in nature. These statements show that, judging from the past, the methods currently used cannot be depended upon for a rapid and symmetrical development of ecology, or for the best development of ecological surveys. These must be developed in a more direct and deliberate manner, by carefully planned and executed ecological investigations. It is desirable also that ecological surveys should be conducted along some one of the three main avenues of approach, individual, aggregate, and associational, in order that the science may develop symmetrically. The following are some of the reasons which may be mentioned in favor of such surveys:

As a record of the associations, their interrelations and responses to their environment--before they have become too much changed or exterminated. This is a duty to future naturalists and to future science. The animal remains in themselves are only a very incomplete record; their activities and environments are an essential part of the animals and should also be preserved.

The study of original conditions is a simpler problem than after interference by man, but excessive modifications result in the simplicity due to annihilation and a corresponding imperfection of knowledge. The value of a knowledge of original conditions tends to increase with time, and will aid much in future interpretations when there is still more disturbance. Thus an important perspective may be developed which will aid in estimating relative values. At the present time the loss of records of original conditions is only beginning to be felt. The possibility of making certain records will vanish with each generation. It is not even desirable to _preserve all_, but it is evident that many ecological records should be preserved.

As the importance of ecological studies, in natural environments, comes to be more generally recognized the serious encroachments of civilization upon habitats and associations is enforced upon us. Not only are the descriptions of these associations very few in number, but the interrelations of the animals in them are even less known, and the chances of preserving adequate records before their complete extinction are becoming fewer every year. Without the least disparagement of other lines of work, one can but wonder if the naturalists of the future will commend our foresight in studying with such great diligence certain aspects of biology which might be very well delayed, while ephemeral and vanishing records are allowed to be obliterated without the least concern. These changes are generally greatest where civilized man is most dominant, and in progressive attenuations, zones, or strips, the degree of change produced by him radiates. Ecology has developed only at a late stage in civilization, after much of the environment has undergone great changes, so that in order to study the original conditions, which are of such great historic and genetic significance, he must make long journeys, or invade the swamps or sterile uplands which man has not yet been able to reduce to the average conditions best suited to his needs. This state of affairs is one which, at times, makes him thankful that there are conditions which, for the present at least, man cannot cultivate and utterly change and mutilate. Some appear to think that an interest in such original conditions is of no particular scientific value, or is largely one of sentiment; still others, that such studies have no practical value. But if we come to consider that the original primeval conditions give us our best conception of the _normal processes of nature_ and are comparable to the normal health of an organism, it puts the subject in another light. A pathological condition is, of course, a state in a natural process, as is also any disturbance of the normal order of nature by man, and each should be studied scientifically. But the science of pathology has developed best as a study of the disturbances of normal processes and is interpreted primarily in terms of the normal; and the artificial should be similarly interpreted--the natural being the basis to which all standards must be referred. A comparison may also profitably be made between natural conditions and the physiological and vital optima of organisms and to the responses which are made with departures from such conditions. Similar comparisons should be made in the study of the responses of aggregations and associations in natural environments and departures from them. No matter how much we learn, the normal must remain as the ideal, and all departures from and disturbances of such conditions must be interpreted in terms of this fundamental unit.

To study disturbed, artificial, and “pathological” conditions, without an adequate knowledge of the normal and original conditions of both the organisms and the environment, is an attempt to interpret the abnormal and artificial in terms of itself, rather than in terms of the normal. If, however, the normal is no longer preserved, then its nearest approach should be studied, but with all the more care and caution. With a proper understanding of the normal, the disturbances made by man will be capable of _interpretation in an orderly sequence strictly comparable to that found in the original and natural conditions_. The cutting down and washing of the lands, the draining and filling of depressions, the flooding of the lands, the destruction (or succession) of plant and animal associations (including crop rotation), are processes brought about or practiced by other organisms or animal agencies. An ecological standpoint gives us a consistent, comprehensive orientation of all these natural and “artificial” activities and processes, and shows the unity in all organic responses to the environment. Man’s influence in the main consists of hastening or retarding “natural processes.”

Naturalists have for a long time spoken of the “balance of nature” and of the all-pervading influence of any serious disturbance of it. This balance is, of course, only a relative condition, and not absolutely fixed. It swings from one side, then back, sometimes showing considerable amplitude in its swing, then again its moves are very slight, mere tremblings, as it were. But now and then some local catastrophic event occurs which overturns everything, as when a volcano becomes active, or some dominant association takes possession of the field,--as in the case of man,--and a new order is initiated and a new balance is developed. The mongoose in Jamaica, our English sparrow, and rabbits in Australia are the classic examples of the overturning of the local order of nature by the agency of other organisms. Obviously this balance is not a condition limited to any particular locality or group of organisms. Balance is very generally conceded to be of fundamental importance in the study of any species or group of organisms, if its place in the economy of nature is understood. A vast number of the problems of the economic zoölogist are thus problems, not so much of individual or aggregate ecology, but ones in which the balance of the whole local biotic _association_ is concerned.

This was the fact pointed out by Möbius when he studied the oyster and came to see that it must be studied not in isolation but as a member of a community, association of animals, or a biocœnosis, as he called these interrelated organisms. These facts are mentioned, as examples from a vast number that are recorded, to show that our applied or economic zoölogy and entomology are _fundamentally more closely related to associational ecology_ than to any other phase of zoölogy, and to suggest that it would be to the great advantage of the students of such problems if they clearly understood this relation. This is also an argument for the ecological organization of a vast number of natural history surveys, because the associational grouping of observations and responses gives the most intimate knowledge of the life of animals in the network of their environmental relations.

In addition to the balance of nature which is found within the small associational units there are the larger ones of considerable geographic extent, which the students of faunal or floral problems frequently call zones or distinct regions. Some of these are distinct ecological units, whose _dynamic status_ should be determined, so that we may know and understand whether it is in a condition of stress, a process of adjustment, or one of relative equilibrium or balance. Under present conditions in what direction does it tend to move? At what rate? The non-ecological surveys have not put these questions or worked deliberately toward a goal which will answer them. For any comprehensive study of this character we need to have determined what may be considered as a _biotic base_, optimum, or balance, toward which relations under given conditions tend, and at which an equilibrium will become established (The Auk, 1908, Vol. XXV, p. 125). Such facts underlie all of the problems involved in the interpretation of climax biotic associations, and their application by man. Cook (1909, Bull. 145, Bur. Plant Industry, U. S. Dept. Agriculture, pp. 7, 8) has expressed similar relations as follows: “Unless we can form a definite idea of the original conditions we cannot expect to judge of their influence on primitive man, nor can we determine what effects man has had upon the vegetation and other natural conditions. We need what might be called a bionomic base line, an idea of the conditions which existed before man came upon the scene, the conditions which would again supervene if the human inhabitants were withdrawn.”

It is perhaps significant that the genetic or successional relations of habitats and associations, as contrasted with their descriptive classification, both in plants and animals, have in the past been developed, not by the ecological students who live and work among conditions greatly modified by man, as in parts of Europe, but in the newer, less modified America. In this respect a parallel exists to the development of our knowledge and the process and genetic interpretation of topography, which has also developed more rapidly in America than elsewhere. The process and genetic method which has developed in this physical science has now spread to the biological sciences and has found a fertile soil there for development on account of the _relatively_ undisturbed biotic conditions which still persist in certain areas.

In this connection it may be worth while to indicate some of the ecological disadvantages under which the non-ecological surveys are carried on. As a rule, such surveys feel no strong obligation to record fully the conditions of the environment, or its developmental processes. The environment is considered as static, and not as a changing medium; it has no past or future, it has merely horizontal extension. The problem as to its _dynamic status_, whether in a condition of stress, in the process of adjustment, or in relative equilibrium, is not raised, or if it should be, it could not be handled. The student eager for new and little-known species is not the one to study such relations, at least, as a rule, this has not been his practice. So long as the success of a day’s work is measured by the length of the list of novelties secured, rather than by the quality and quantity of ecological relations discovered, such students and surveys will not contribute greatly to our knowledge of the economy of nature in the regions surveyed.

At the present time it is very difficult to secure trained men to do ecological surveying. Even a superficial examination of this paper should show that familiarity with ecologic methods and results is not one to be acquired offhand, but a knowledge which requires considerable special training; not only as much as is usually required for other kinds of zoölogical work, but generally more, because of its synthetic relational tendency which requires a broad knowledge as well as some special knowledge in several lines of biology and the allied sciences. Conventionally considered, a properly equipped physiologist must have a working knowledge of certain phases of modern physics and chemistry in addition to his grounding in biology. A properly trained anatomist should have a knowledge of physiological and developmental processes, or his anatomy is purely descriptive and static. A student of general zoölogy should be grounded not only in physiological and developmental processes, but also in the relations of the organisms to their complete environment. The ecologist requires also the grounding in physiological, developmental, and ecological processes of adjustment, but as well he must understand the processes by which the vegetation and the physical environment have been and are being developed and their method of mutual interrelations and adjustment. It is difficult for some students to develop the ecological phases in the field. There are many disadvantages to be overcome. The difficulties are similar, in some respects, to those of the ethnologist who is sent on some museum expedition. The wealthy donor of the funds may wish to see a room filled with specimens on the return of the ethnologist, so that materials which have bulk and make a showing take precedence over detailed studies of the habits, traditions, languages, and descriptions of the people, because such studies require appreciation rather than inspection for evaluation. The zoölogical student may meet with just the same kind of difficulty. His institutional authorities often judge values by the cubic foot and pound, rather than by the quality of relations discovered. The student himself who has had an extensive collecting experience, in which quantity and variety have been the ideal, finds it difficult to return from a day’s work with only a few pages of notes on the responses of the animals, and with perhaps only a few specimens.

With such an understanding of the general rules of the game we may turn to the application or art of ecology, to indicate its relation to general problems. With a grounding in the general principles of organic response to the total environment, one is able to see that the disturbances due to man are a problem in the adjustment of the highest type of animal, as a member of an animal association, to its complete environment. The “control of nature” for which men strive is the process of making the environments and associations _to order_. The disturbances in the natural order may be looked upon as so many huge experiments or trial activities in this process of adjustment.

If natural preserves are not made, how will the next generation be best grounded in the general principles of the science? Are these complex modified conditions the natural place to start the student, or should such problems be reserved for the maturely trained one? These disturbed fragmentary conditions may be likened to fragmentary fossils whose interpretation is attempted. A paleontologist whose only knowledge of animals was derived from such fragments, and who had never known a perfect living animal, would certainly be at a great disadvantage in such an investigation. The natural starting point therefore seems to be in as nearly natural normal environments and associations as is possible, and with such experience one is prepared for the more complex problems resulting from man’s activity.

By way of conclusion, some of the main advantages of ecological surveys are:

1. The _record_ of natural environments and their associations for future generations.

2. The study of natural biotic conditions giving a _perspective_ not derived in any other way.

3. The clearer conception of the _dynamic relations_ of the balance of nature, biotic base, and climax associations.

4. Emphasis of the _process and interpretative_ phase of scientific investigation over that of purely descriptive study.

5. Facilitating the invention of multiple working hypotheses which bear upon animal responses in nature.