Darwinism (1889) : An exposition of the theory of natural selection, with some of its applications E-Book
Alfred Russel Wallace
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- Herausgeber: Krill Press
- Kategorie: Geisteswissenschaft
- Sprache: Englisch
- Veröffentlichungsjahr: 2016
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DARWINISM (1889) : AN EXPOSITION OF THE THEORY OF NATURAL SELECTION, WITH SOME OF ITS APPLICATIONS
Alfred Russel Wallace
DOSSIER PRESS
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TABLE OF CONTENTS
PREFACE TO SECOND EDITION
PREFACE TO FIRST EDITION
CHAPTER I
FOOTNOTES:
CHAPTER II
FOOTNOTES:
CHAPTER III
FOOTNOTES:
CHAPTER IV
FOOTNOTES:
CHAPTER V
FOOTNOTES:
CHAPTER VI
FOOTNOTES:
CHAPTER VII
FOOTNOTES:
CHAPTER VIII
FOOTNOTES:
CHAPTER IX
FOOTNOTES:
CHAPTER X
FOOTNOTES:
CHAPTER XI
FOOTNOTES:
CHAPTER XII
FOOTNOTES:
CHAPTER XIII
FOOTNOTES:
CHAPTER XIV
FOOTNOTES:
CHAPTER XV
FOOTNOTES:
Darwinism (1889) : An exposition of the theory of natural selection, with some of its applications
By
Alfred Russel Wallace
Darwinism (1889) : An exposition of the theory of natural selection, with some of its applications
Published by Dossier Press
New York City, NY
First published circa 1913
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PREFACE TO SECOND EDITION
THE PRESENT EDITION IS A reprint of the first, with a few verbal corrections and the alteration of some erroneous or doubtful statements. Of these latter the following are the most important:—
P. . The statement as to the fulmar petrel, which Professor A. Newton assures me is erroneous, has been modified.
P. . A note is added as to Darwin’s statement about the missel and song-thrushes in Scotland.
P. . An error as to the differently-coloured herds of cattle in the Falkland Islands, is corrected.
PARKSTONE, DORSET
August, 1889.
PREFACE TO FIRST EDITION
THE PRESENT WORK TREATS THE problem of the Origin of Species on the same general lines as were adopted by Darwin; but from the standpoint reached after nearly thirty years of discussion, with an abundance of new facts and the advocacy of many new or old theories.
While not attempting to deal, even in outline, with the vast subject of evolution in general, an endeavour has been made to give such an account of the theory of Natural Selection as may enable any intelligent reader to obtain a clear conception of Darwin’s work, and to understand something of the power and range of his great principle.
Darwin wrote for a generation which had not accepted evolution, and which poured contempt on those who upheld the derivation of species from species by any natural law of descent. He did his work so well that “descent with modification” is now universally accepted as the order of nature in the organic world; and the rising generation of naturalists can hardly realise the novelty of this idea, or that their fathers considered it a scientific heresy to be condemned rather than seriously discussed.
The objections now made to Darwin’s theory apply, solely, to the particular means by which the change of species has been brought about, not to the fact of that change. The objectors seek to minimise the agency of natural selection and to subordinate it to laws of variation, of use and disuse, of intelligence, and of heredity. These views and objections are urged with much force and more confidence, and for the most part by the modern school of laboratory naturalists, to whom the peculiarities and distinctions of species, as such, their distribution and their affinities, have little interest as compared with the problems of histology and embryology, of physiology and morphology. Their work in these departments is of the greatest interest and of the highest importance, but it is not the kind of work which, by itself, enables one to form a sound judgment on the questions involved in the action of the law of natural selection. These rest mainly on the external and vital relations of species to species in a state of nature—on what has been well termed by Semper the “physiology of organisms,” rather than on the anatomy or physiology of organs.
It has always been considered a weakness in Darwin’s work that he based his theory, primarily, on the evidence of variation in domesticated animals and cultivated plants. I have endeavoured to secure a firm foundation for the theory in the variations of organisms in a state of nature; and as the exact amount and precise character of these variations is of paramount importance in the numerous problems that arise when we apply the theory to explain the facts of nature, I have endeavoured, by means of a series of diagrams, to exhibit to the eye the actual variations as they are found to exist in a sufficient number of species. By doing this, not only does the reader obtain a better and more precise idea of variation than can be given by any number of tabular statements or cases of extreme individual variation, but we obtain a basis of fact by which to test the statements and objections usually put forth on the subject of specific variability; and it will be found that, throughout the work, I have frequently to appeal to these diagrams and the facts they illustrate, just as Darwin was accustomed to appeal to the facts of variation among dogs and pigeons.
I have also made what appears to me an important change in the arrangement of the subject. Instead of treating first the comparatively difficult and unfamiliar details of variation, I commence with the Struggle for Existence, which is really the fundamental phenomenon on which natural selection depends, while the particular facts which illustrate it are comparatively familiar and very interesting. It has the further advantage that, after discussing variation and the effects of artificial selection, we proceed at once to explain how natural selection acts.
Among the subjects of novelty or interest discussed in this volume, and which have important bearings on the theory of natural selection, are: (1) A proof that all specific characters are (or once have been) either useful in themselves or correlated with useful characters (Chap. VI); (2) a proof that natural selection can, in certain cases, increase the sterility of crosses (Chap. VII); (3) a fuller discussion of the colour relations of animals, with additional facts and arguments on the origin of sexual differences of colour (Chaps. VIII-X); (4) an attempted solution of the difficulty presented by the occurrence of both very simple and very complex modes of securing the cross-fertilisation of plants (Chap. XI); (5) some fresh facts and arguments on the wind-carriage of seeds, and its bearing on the wide dispersal of many arctic and alpine plants (Chap. XII); (6) some new illustrations of the non-heredity of acquired characters, and a proof that the effects of use and disuse, even if inherited, must be overpowered by natural selection (Chap. XIV); and (7) a new argument as to the nature and origin of the moral and intellectual faculties of man (Chap. XV).
Although I maintain, and even enforce, my differences from some of Darwin’s views, my whole work tends forcibly to illustrate the overwhelming importance of Natural Selection over all other agencies in the production of new species. I thus take up Darwin’s earlier position, from which he somewhat receded in the later editions of his works, on account of criticisms and objections which I have endeavoured to show are unsound. Even in rejecting that phase of sexual selection depending on female choice, I insist on the greater efficacy of natural selection. This is pre-eminently the Darwinian doctrine, and I therefore claim for my book the position of being the advocate of pure Darwinism.
I wish to express my obligation to Mr. Francis Darwin for lending me some of his father’s unused notes, and to many other friends for facts or information, which have, I believe, been acknowledged either in the text or footnotes. Mr. James Sime has kindly read over the proofs and given me many useful suggestions; and I have to thank Professor Meldola, Mr. Hemsley, and Mr. E.B. Poulton for valuable notes or corrections in the later chapters in which their special subjects are touched upon.
GODALMING, March 1889.
WHAT ARE “SPECIES” AND WHAT IS MEANT BY THEIR “ORIGIN”
Definition of species—Special creation—The early transmutationists—Scientific opinion before Darwin—The problem before Darwin—The change of opinion effected by Darwin—The Darwinian theory—Proposed mode of treatment of the subject
THE STRUGGLE FOR EXISTENCE
Its importance—The struggle among plants—Among animals—Illustrative cases—Succession of trees in forests of Denmark—The struggle for existence on the Pampas—Increase of organisms in a geometrical ratio—Examples of rapid increase of animals—Rapid increase and wide spread of plants—Great fertility not essential to rapid increase—Struggle between closely allied species most severe—The ethical aspect of the struggle for existence
THE VARIABILITY OF SPECIES IN A STATE OF NATURE
Importance of variability—Popular ideas regarding it—Variability of the lower animals—The variability of insects—Variation among lizardsVariation among birds—Diagrams of bird-variation—Number of varying individuals—Variation in the mammalia—Variation in internal organs—Variations in the skull—Variations in the habits of animals—The variability of plants—Species which vary little—Concluding remarks
VARIATION OF DOMESTICATED ANIMALS AND CULTIVATED PLANTS
The facts of variation and artificial selection—Proofs of the generality of variation—Variations of apples and melons—Variations of flowers—Variations of domestic animals—Domestic pigeons—Acclimatisation—Circumstances favourable to selection by man—Conditions favourable to variation—Concluding remarks
NATURAL SELECTION BY VARIATION AND SURVIVAL OF THE FITTEST
Effect of struggle for existence under unchanged conditions—The effect under change of conditions—Divergence of character—In insects—In birds—In mammalia—Divergence leads to a maximum of life in each area—Closely allied species inhabit distinct areas—Adaptation to conditions at various periods of life—The continued existence of low forms of life—Extinction of low types among the higher animals—Circumstances favourable to the origin of new species—Probable origin of the dippers—The importance of isolation—On the advance of organisation by natural selection—Summary of the first five chapters
DIFFICULTIES AND OBJECTIONS
Difficulty as to smallness of variations—As to the right variations occurring when required—The beginnings of important organs—The mammary glands—The eyes of flatfish—Origin of the eye—Useless or non-adaptive characters—Recent extension of the region of utility in plants—The same in animals—Uses of tails—Of the horns of deer—Of the scale-ornamentation of reptiles—Instability of non-adaptive characters—Delboeuf’s law—No “specific” character proved to be useless—The swamping effects of intercrossing—Isolation as preventing intercrossing—Gulick on the effects of isolation—Cases in which isolation is ineffective
ON THE INFERTILITY OF CROSSES BETWEEN DISTINCT SPECIES AND THE USUAL STERILITY OF THEIR HYBRID OFFSPRING
Statement of the problem—Extreme susceptibility of the reproductive functions—Reciprocal crosses—Individual differences in respect to cross-fertilisation—Dimorphism and trimorphism among plants—Cases of the fertility of hybrids and of the infertility of mongrels—The effects of close interbreeding—Mr. Huth’s objections—Fertile hybrids among animals—Fertility of hybrids among plants—Cases of sterility of mongrels—Parallelism between crossing and change of conditions—Remarks on the facts of hybridity—Sterility due to changed conditions and usually correlated with other characters—Correlation of colour with constitutional peculiarities—The isolation of varieties by selective association—The influence of natural selection upon sterility and fertility—Physiological selection—Summary and concluding remarks
THE ORIGIN AND USES OF COLOUR IN ANIMALS
The Darwinian theory threw new light on organic colour—The problem to be solved—The constancy of animal colour indicates utility—Colour and environment—Arctic animals white—Exceptions prove the rule—Desert, forest, nocturnal, and oceanic animals—General theories of animal colour—Variable protective colouring—Mr. Poulton’s experiments—Special or local colour adaptations—Imitation of particular objects—How they have been produced—Special protective colouring of butterflies—Protective resemblance among marine animals—Protection by terrifying enemies—Alluring coloration—The coloration of birds’ eggs—Colour as a means of recognition—Summary of the preceding exposition—Influence of locality or of climate on colour—Concluding remarks
WARNING COLORATION AND MIMICRY
The skunk as an example of warning coloration—Warning colours among insects—Butterflies—Caterpillars—Mimicry—How mimicry has been produced—Heliconidae—Perfection of the imitation—Other cases of mimicry among Lepidoptera—Mimicry among protected groups—Its explanation—Extension of the principle—Mimicry in other orders of insects—Mimicry among the vertebrata—Snakes—The rattlesnake and the cobra—Mimicry among birds—Objections to the theory of mimicry—Concluding remarks on warning colours and mimicry
COLOURS AND ORNAMENTS CHARACTERISTIC OF SEX
Sex colours in the mollusca and crustacea—In insects—In butterflies and moths—Probable causes of these colours—Sexual selection as a supposed cause—Sexual coloration of birds—Cause of dull colours of female birds—Relation of sex colour to nesting habits—Sexual colours of other vertebrates—Sexual selection by the struggles of males—Sexual characters due to natural selection—Decorative plumage of males and its effect on the females—Display of decorative plumage by the males—A theory of animal coloration—The origin of accessory plumes—Development of accessory plumes and their display—The effect of female preference will be neutralised by natural selection—General laws of animal coloration—Concluding remarks
THE SPECIAL COLOURS OF PLANTS: THEIR ORIGIN AND PURPOSE
The general colour relations of plants—Colours of fruits—The meaning of nuts—Edible or attractive fruits—The colours of flowers—Modes of securing cross-fertilisation—The interpretation of the facts—Summary of additional facts bearing on insect fertilisation—Fertilisation of flowers by birds—Self-fertilisation of flowers—Difficulties and contradictions—Intercrossing not necessarily advantageous—Supposed evil results of close interbreeding—How the struggle for existence acts among flowers—Flowers the product of insect agency—Concluding remarks on colour in nature
THE GEOGRAPHICAL DISTRIBUTION OF ORGANISMS
The facts to be explained—The conditions which have determined distribution—The permanence of oceans—Oceanic and continental areas—Madagascar and New Zealand—The teachings of the thousand-fathom line—The distribution of marsupials—The distribution of tapirs—Powers of dispersal as illustrated by insular organisms—Birds and insects at sea—Insects at great altitudes—The dispersal of plants—Dispersal of seeds by the wind—Mineral matter carried by the wind—Objections to the theory of wind-dispersal answered—Explanation of north temperate plants in the southern hemisphere—No proof of glaciation in the tropics—Lower temperature not needed to explain the facts—Concluding remarks
THE GEOLOGICAL EVIDENCES OF EVOLUTION
What we may expect—The number of known species of extinct animals—Causes of the imperfection of the geological record—Geological evidences of evolution—Shells—Crocodiles—The rhinoceros tribe—The pedigree of the horse tribe—Development of deer’s horns—Brain development—Local relations of fossil and living animals—Cause of extinction of large animals—Indications of general progress in plants and animals—The progressive development of plants—Possible cause of sudden late appearance of exogens—Geological distribution of insects—Geological succession of vertebrata—Concluding remarks
FUNDAMENTAL PROBLEMS IN RELATION TO VARIATION AND HEREDITY
Fundamental difficulties and objections—Mr. Herbert Spencer’s factors of organic evolution—Disuse and effects of withdrawal of natural selection—Supposed effects of disuse among wild animals—Difficulty as to co-adaptation of parts by variation and selection—Direct action of the environment—The American school of evolutionists—Origin of the feet of the ungulates—Supposed action of animal intelligence—Semper on the direct influence of the environment—Professor Geddes’s theory of variation in plants—Objections to the theory—On the origin of spines—Variation and selection overpower the effects of use and disuse—Supposed action of the environment in imitating variations—Weismann’s theory of heredity—The cause of variation—The non-heredity of acquired characters—The theory of instinct—Concluding remarks
DARWINISM APPLIED TO MAN
General identity of human and animal structure—Rudiments and variations showing relation of man to other mammals—The embryonic development of man and other mammalia—Diseases common to man and the lower animals—The animals most nearly allied to man—The brains of man and apes—External differences of man and apes—Summary of the animal characteristics of man—The geological antiquity of man—The probable birthplace of man—The origin of the moral and intellectual nature of man—The argument from continuity—The origin of the mathematical faculty—The origin of the musical and artistic faculties—Independent proof that these faculties have not been developed by natural selection—The interpretation of the facts—Concluding remarks
CHAPTER I
WHAT ARE “SPECIES,” AND WHAT IS MEANT BY THEIR “ORIGIN”
Definition of species—Special creation—The early Transmutationists—Scientific opinion before Darwin—The problem before Darwin—The change of opinion effected by Darwin—The Darwinian theory—Proposed mode of treatment of the subject.
The title of Mr. Darwin’s great work is—On the Origin of Species by means of Natural Selection and the Preservation of Favoured Races in the Struggle for Life. In order to appreciate fully the aim and object of this work, and the change which it has effected not only in natural history but in many other sciences, it is necessary to form a clear conception of the meaning of the term “species,” to know what was the general belief regarding them at the time when Mr. Darwin’s book first appeared, and to understand what he meant, and what was generally meant, by discovering their “origin.” It is for want of this preliminary knowledge that the majority of educated persons who are not naturalists are so ready to accept the innumerable objections, criticisms, and difficulties of its opponents as proofs that the Darwinian theory is unsound, while it also renders them unable to appreciate, or even to comprehend, the vast change which that theory has effected in the whole mass of thought and opinion on the great question of evolution.
The term “species” was thus defined by the celebrated botanist De Candolle: “A species is a collection of all the individuals which resemble each other more than they resemble anything else, which can by mutual fecundation produce fertile individuals, and which reproduce themselves by generation, in such a manner that we may from analogy suppose them all to have sprung from one single individual.” And the zoologist Swainson gives a somewhat similar definition: “A species, in the usual acceptation of the term, is an animal which, in a state of nature, is distinguished by certain peculiarities of form, size, colour, or other circumstances, from another animal. It propagates, ‘after its kind,’ individuals perfectly resembling the parent; its peculiarities, therefore, are permanent.”
To illustrate these definitions we will take two common English birds, the rook (Corvus frugilegus) and the crow (Corvus corone). These are distinct species, because, in the first place, they always differ from each other in certain slight peculiarities of structure, form, and habits, and, in the second place, because rooks always produce rooks, and crows produce crows, and they do not interbreed. It was therefore concluded that all the rooks in the world had descended from a single pair of rooks, and the crows in like manner from a single pair of crows, while it was considered impossible that crows could have descended from rooks or vice versâ. The “origin” of the first pair of each kind was a mystery. Similar remarks may be applied to our two common plants, the sweet violet (Viola odorata) and the dog violet (Viola canina). These also produce their like and never produce each other or intermingle, and they were therefore each supposed to have sprung from a single individual whose “origin” was unknown. But besides the crow and the rook there are about thirty other kinds of birds in various parts of the world, all so much like our species that they receive the common name of crows; and some of them differ less from each other than does our crow from our rook. These are all species of the genus Corvus, and were therefore believed to have been always as distinct as they are now, neither more nor less, and to have each descended from one pair of ancestral crows of the same identical species, which themselves had an unknown “origin.” Of violets there are more than a hundred different kinds in various parts of the world, all differing very slightly from each other and forming distinct species of the genus Viola. But, as these also each produce their like and do not intermingle, it was believed that every one of them had always been as distinct from all the others as it is now, that all the individuals of each kind had descended from one ancestor, but that the “origin” of these hundred slightly differing ancestors was unknown. In the words of Sir John Herschel, quoted by Mr. Darwin, the origin of such species was “the mystery of mysteries.”
The Early Transmutationists.
A few great naturalists, struck by the very slight difference between many of these species, and the numerous links that exist between the most different forms of animals and plants, and also observing that a great many species do vary considerably in their forms, colours, and habits, conceived the idea that they might be all produced one from the other. The most eminent of these writers was a great French naturalist, Lamarck, who published an elaborate work, the Philosophie Zoologique, in which he endeavoured to prove that all animals whatever are descended from other species of animals. He attributed the change of species chiefly to the effect of changes in the conditions of life—such as climate, food, etc.—and especially to the desires and efforts of the animals themselves to improve their condition, leading to a modification of form or size in certain parts, owing to the well-known physiological law that all organs are strengthened by constant use, while they are weakened or even completely lost by disuse. The arguments of Lamarck did not, however, satisfy naturalists, and though a few adopted the view that closely allied species had descended from each other, the general belief of the educated public was, that each species was a “special creation” quite independent of all others; while the great body of naturalists equally held, that the change from one species to another by any known law or cause was impossible, and that the “origin of species” was an unsolved and probably insoluble problem. The only other important work dealing with the question was the celebrated Vestiges of Creation, published anonymously, but now acknowledged to have been written by the late Robert Chambers. In this work the action of general laws was traced throughout the universe as a system of growth and development, and it was argued that the various species of animals and plants had been produced in orderly succession from each other by the action of unknown laws of development aided by the action of external conditions. Although this work had a considerable effect in influencing public opinion as to the extreme improbability of the doctrine of the independent “special creation” of each species, it had little effect upon naturalists, because it made no attempt to grapple with the problem in detail, or to show in any single case how the allied species of a genus could have arisen, and have preserved their numerous slight and apparently purposeless differences from each other. No clue whatever was afforded to a law which should produce from any one species one or more slightly differing but yet permanently distinct species, nor was any reason given why such slight yet constant differences should exist at all.
Scientific Opinion before Darwin.
In order to show how little effect these writers had upon the public mind, I will quote a few passages from the writings of Sir Charles Lyell, as representing the opinions of the most advanced thinkers in the period immediately preceding that of Darwin’s work. When recapitulating the facts and arguments in favour of the invariability and permanence of species, he says: “The entire variation from the original type which any given kind of change can produce may usually be effected in a brief period of time, after which no further deviation can be obtained by continuing to alter the circumstances, though ever so gradually, indefinite divergence either in the way of improvement or deterioration being prevented, and the least possible excess beyond the defined limits being fatal to the existence of the individual.” In another place he maintains that “varieties of some species may differ more than other species do from each other without shaking our confidence in the reality of species.” He further adduces certain facts in geology as being, in his opinion, “fatal to the theory of progressive development,” and he explains the fact that there are so often distinct species in countries of similar climate and vegetation by “special creations” in each country; and these conclusions were arrived at after a careful study of Lamarck’s work, a full abstract of which is given in the earlier editions of the Principles of Geology.
Professor Agassiz, one of the greatest naturalists of the last generation, went even further, and maintained not only that each species was specially created, but that it was created in the proportions and in the localities in which we now find it to exist. The following extract from his very instructive book on Lake Superior explains this view: “There are in animals peculiar adaptations which are characteristic of their species, and which cannot be supposed to have arisen from subordinate influences. Those which live in shoals cannot be supposed to have been created in single pairs. Those which are made to be the food of others cannot have been created in the same proportions as those which live upon them. Those which are everywhere found in innumerable specimens must have been introduced in numbers capable of maintaining their normal proportions to those which live isolated and are comparatively and constantly fewer. For we know that this harmony in the numerical proportions between animals is one of the great laws of nature. The circumstance that species occur within definite limits where no obstacles prevent their wider distribution leads to the further inference that these limits were assigned to them from the beginning, and so we should come to the final conclusion that the order which prevails throughout nature is intentional, that it is regulated by the limits marked out on the first day of creation, and that it has been maintained unchanged through ages with no other modifications than those which the higher intellectual powers of man enable him to impose on some few animals more closely connected with him.”
These opinions of some of the most eminent and influential writers of the pre-Darwinian age seem to us, now, either altogether obsolete or positively absurd; but they nevertheless exhibit the mental condition of even the most advanced section of scientific men on the problem of the nature and origin of species. They render it clear that, notwithstanding the vast knowledge and ingenious reasoning of Lamarck, and the more general exposition of the subject by the author of the Vestiges of Creation, the first step had not been taken towards a satisfactory explanation of the derivation of any one species from any other. Such eminent naturalists as Geoffroy Saint Hilaire, Dean Herbert, Professor Grant, Von Buch, and some others, had expressed their belief that species arose as simple varieties, and that the species of each genus were all descended from a common ancestor; but none of them gave a clue as to the law or the method by which the change had been effected. This was still “the great mystery.” As to the further question—how far this common descent could be carried; whether distinct families, such as crows and thrushes, could possibly have descended from each other; or, whether all birds, including such widely distinct types as wrens, eagles, ostriches, and ducks, could all be the modified descendants of a common ancestor; or, still further, whether mammalia, birds, reptiles, and fishes, could all have had a common origin;—these questions had hardly come up for discussion at all, for it was felt that, while the very first step along the road of “transmutation of species” (as it was then called) had not been made, it was quite useless to speculate as to how far it might be possible to travel in the same direction, or where the road would ultimately lead to.
The Problem before Darwin.
It is clear, then, that what was understood by the “origin” or the “transmutation” of species before Darwin’s work appeared, was the comparatively simple question whether the allied species of each genus had or had not been derived from one another and, remotely, from some common ancestor, by the ordinary method of reproduction and by means of laws and conditions still in action and capable of being thoroughly investigated. If any naturalist had been asked at that day whether, supposing it to be clearly shown that all the different species of each genus had been derived from some one ancestral species, and that a full and complete explanation were to be given of how each minute difference in form, colour, or structure might have originated, and how the several peculiarities of habit and of geographical distribution might have been brought about—whether, if this were done, the “origin of species” would be discovered, the great mystery solved, he would undoubtedly have replied in the affirmative. He would probably have added that he never expected any such marvellous discovery to be made in his lifetime. But so much as this assuredly Mr. Darwin has done, not only in the opinion of his disciples and admirers, but by the admissions of those who doubt the completeness of his explanations. For almost all their objections and difficulties apply to those larger differences which separate genera, families, and orders from each other, not to those which separate one species from the species to which it is most nearly allied, and from the remaining species of the same genus. They adduce such difficulties as the first development of the eye, or of the milk-producing glands of the mammalia; the wonderful instincts of bees and of ants; the complex arrangements for the fertilisation of orchids, and numerous other points of structure or habit, as not being satisfactorily explained. But it is evident that these peculiarities had their origin at a very remote period of the earth’s history, and no theory, however complete, can do more than afford a probable conjecture as to how they were produced. Our ignorance of the state of the earth’s surface and of the conditions of life at those remote periods is very great; thousands of animals and plants must have existed of which we have no record; while we are usually without any information as to the habits and general life-history even of those of which we possess some fragmentary remains; so that the truest and most complete theory would not enable us to solve all the difficult problems which the whole course of the development of life upon our globe presents to us.
What we may expect a true theory to do is to enable us to comprehend and follow out in some detail those changes in the form, structure, and relations of animals and plants which are effected in short periods of time, geologically speaking, and which are now going on around us. We may expect it to explain satisfactorily most of the lesser and superficial differences which distinguish one species from another. We may expect it to throw light on the mutual relations of the animals and plants which live together in any one country, and to give some rational account of the phenomena presented by their distribution in different parts of the world. And, lastly, we may expect it to explain many difficulties and to harmonise many incongruities in the excessively complex affinities and relations of living things. All this the Darwinian theory undoubtedly does. It shows us how, by means of some of the most universal and ever-acting laws in nature, new species are necessarily produced, while the old species become extinct; and it enables us to understand how the continuous action of these laws during the long periods with which geology makes us acquainted is calculated to bring about those greater differences presented by the distinct genera, families, and orders into which all living things are classified by naturalists. The differences which these present are all of the same nature as those presented by the species of many large genera, but much greater in amount; and they can all be explained by the action of the same general laws and by the extinction of a larger or smaller number of intermediate species. Whether the distinctions between the higher groups termed Classes and Sub-kingdoms may be accounted for in the same way is a much more difficult question. The differences which separate the mammals, birds, reptiles, and fishes from each other, though vast, yet seem of the same nature as those which distinguish a mouse from an elephant or a swallow from a goose. But the vertebrate animals, the mollusca, and the insects, are so radically distinct in their whole organisation and in the very plan of their structure, that objectors may not unreasonably doubt whether they can all have been derived from a common ancestor by means of the very same laws as have sufficed for the differentiation of the various species of birds or of reptiles.
The Change of Opinion effected by Darwin.
The point I wish especially to urge is this. Before Darwin’s work appeared, the great majority of naturalists, and almost without exception the whole literary and scientific world, held firmly to the belief that species were realities, and had not been derived from other species by any process accessible to us; the different species of crow and of violet they are now, and to have originated by some totally unknown process so far removed from ordinary reproduction that it was usually spoken of as “special creation.” There was, then, no question of the origin of families, orders, and classes, because the very first step of all, the “origin of species,” was believed to be an insoluble problem. But now this is all changed. The whole scientific and literary world, even the whole educated public, accepts, as a matter of common knowledge, the origin of species from other allied species by the ordinary process of natural birth. The idea of special creation or any altogether exceptional mode of production is absolutely extinct! Yet more: this is held also to apply to many higher groups as well as to the species of a genus, and not even Mr. Darwin’s severest critics venture to suggest that the primeval bird, reptile, or fish must have been “specially created.” And this vast, this totally unprecedented change in public opinion has been the result of the work of one man, and was brought about in the short space of twenty years! This is the answer to those who continue to maintain that the “origin of species” is not yet discovered; that there are still doubts and difficulties; that there are divergencies of structure so great that we cannot understand how they had their beginning. We may admit all this, just as we may admit that there are enormous difficulties in the way of a complete comprehension of the origin and nature of all the parts of the solar system and of the stellar universe. But we claim for Darwin that he is the Newton of natural history, and that, just so surely as that the discovery and demonstration by Newton of the law of gravitation established order in place of chaos and laid a sure foundation for all future study of the starry heavens, so surely has Darwin, by his discovery of the law of natural selection and his demonstration of the great principle of the preservation of useful variations in the struggle for life, not only thrown a flood of light on the process of development of the whole organic world, but also established a firm foundation for all future study of nature.
In order to show the view Darwin took of his own work, and what it was that he alone claimed to have done, the concluding passage of the introduction to the Origin of Species should be carefully considered. It is as follows: “Although much remains obscure, and will long remain obscure, I can entertain no doubt, after the most deliberate and dispassionate judgment of which I am capable, that the view which most naturalists until recently entertained and which I formerly entertained—namely, that each species has been independently created—is erroneous. I am fully convinced that species are not immutable; but that those belonging to what are called the same genera are lineal descendants of some other and generally extinct species, in the same manner as the acknowledged varieties of any one species are the descendants of that species. Furthermore, I am convinced that Natural Selection has been the most important, but not the exclusive, means of modification.”
It should be especially noted that all which is here claimed is now almost universally admitted, while the criticisms of Darwin’s works refer almost exclusively to those numerous questions which, as he himself says, “will long remain obscure.”
The Darwinian Theory.
As it will be necessary, in the following chapters, to set forth a considerable body of facts in almost every department of natural history, in order to establish the fundamental propositions on which the theory of natural selection rests, I propose to give a preliminary statement of what the theory really is, in order that the reader may better appreciate the necessity for discussing so many details, and may thus feel a more enlightened interest in them. Many of the facts to be adduced are so novel and so curious that they are sure to be appreciated by every one who takes an interest in nature, but unless the need of them is clearly seen it may be thought that time is being wasted on mere curious details and strange facts which have little bearing on the question at issue.
The theory of natural selection rests on two main classes of facts which apply to all organised beings without exception, and which thus take rank as fundamental principles or laws. The first is, the power of rapid multiplication in a geometrical progression; the second, that the offspring always vary slightly from the parents, though generally very closely resembling them. From the first fact or law there follows, necessarily, a constant struggle for existence; because, while the offspring always exceed the parents in number, generally to an enormous extent, yet the total number of living organisms in the world does not, and cannot, increase year by year. Consequently every year, on the average, as many die as are born, plants as well as animals; and the majority die premature deaths. They kill each other in a thousand different ways; they starve each other by some consuming the food that others want; they are destroyed largely by the powers of nature—by cold and heat, by rain and storm, by flood and fire. There is thus a perpetual struggle among them which shall live and which shall die; and this struggle is tremendously severe, because so few can possibly remain alive—one in five, one in ten, often only one in a hundred or even one in a thousand.
Then comes the question, Why do some live rather than others? If all the individuals of each species were exactly alike in every respect, we could only say it is a matter of chance. But they are not alike. We find that they vary in many different ways. Some are stronger, some swifter, some hardier in constitution, some more cunning. An obscure colour may render concealment more easy for some, keener sight may enable others to discover prey or escape from an enemy better than their fellows. Among plants the smallest differences may be useful or the reverse. The earliest and strongest shoots may escape the slug; their greater vigour may enable them to flower and seed earlier in a wet autumn; plants best armed with spines or hairs may escape being devoured; those whose flowers are most conspicuous may be soonest fertilised by insects. We cannot doubt that, on the whole, any beneficial variations will give the possessors of it a greater probability of living through the tremendous ordeal they have to undergo. There may be something left to chance, but on the whole the fittest will survive.
Then we have another important fact to consider, the principle of heredity or transmission of variations. If we grow plants from seed or breed any kind of animals year after year, consuming or giving away all the increase we do not wish to keep just as they come to hand, our plants or animals will continue much the same; but if every year we carefully save the best seed to sow and the finest or brightest coloured animals to breed from, we shall soon find that an improvement will take place, and that the average quality of our stock will be raised. This is the way in which all our fine garden fruits and vegetables and flowers have been produced, as well as all our splendid breeds of domestic animals; and they have thus become in many cases so different from the wild races from which they originally sprang as to be hardly recognisable as the same. It is therefore proved that if any particular kind of variation is preserved and bred from, the variation itself goes on increasing in amount to an enormous extent; and the bearing of this on the question of the origin of species is most important. For if in each generation of a given animal or plant the fittest survive to continue the breed, then whatever may be the special peculiarity that causes “fitness” in the particular case, that peculiarity will go on increasing and strengthening so long as it is useful to the species. But the moment it has reached its maximum of usefulness, and some other quality or modification would help in the struggle, then the individuals which vary in the new direction will survive; and thus a species may be gradually modified, first in one direction, then in another, till it differs from the original parent form as much as the greyhound differs from any wild dog or the cauliflower from any wild plant. But animals or plants which thus differ in a state of nature are always classed as distinct species, and thus we see how, by the continuous survival of the fittest or the preservation of favoured races in the struggle for life, new species may be originated.
This self-acting process which, by means of a few easily demonstrated groups of facts, brings about change in the organic world, and keeps each species in harmony with the conditions of its existence, will appear to some persons so clear and simple as to need no further demonstration. But to the great majority of naturalists and men of science endless difficulties and objections arise, owing to the wonderful variety of animal and vegetable forms, and the intricate relations of the different species and groups of species with each other; and it was to answer as many of these objections as possible, and to show that the more we know of nature the more we find it to harmonise with the development hypothesis, that Darwin devoted the whole of his life to collecting facts and making experiments, the record of a portion of which he has given us in a series of twelve masterly volumes.
Proposed Mode of Treatment of the Subject.
It is evidently of the most vital importance to any theory that its foundations should be absolutely secure. It is therefore necessary to show, by a wide and comprehensive array of facts, that animals and plants do perpetually vary in the manner and to the amount requisite; and that this takes place in wild animals as well as in those which are domesticated. It is necessary also to prove that all organisms do tend to increase at the great rate alleged, and that this increase actually occurs, under favourable conditions. We have to prove, further, that variations of all kinds can be increased and accumulated by selection; and that the struggle for existence to the extent here indicated actually occurs in nature, and leads to the continued preservation of favourable variations.
These matters will be discussed in the four succeeding chapters, though in a somewhat different order—the struggle for existence and the power of rapid multiplication, which is its cause, occupying the first place, as comprising those facts which are the most fundamental and those which can be perfectly explained without any reference to the less generally understood facts of variation. These chapters will be followed by a discussion of certain difficulties, and of the vexed question of hybridity. Then will come a rather full account of the more important of the complex relations of organisms to each other and to the earth itself, which are either fully explained or greatly elucidated by the theory. The concluding chapter will treat of the origin of man and his relations to the lower animals.
FOOTNOTES:
Geography and Classification of Animals, p. 350.
These expressions occur in Chapter IX. of the earlier editions (to the ninth) of the Principles of Geology.
L. Agassiz, Lake Superior, p. 377.
CHAPTER II
THE STRUGGLE FOR EXISTENCE
Its importance—The struggle among plants—Among animals—Illustrative cases—Succession of trees in forests of Denmark—The struggle for existence on the Pampas—Increase of organisms in a geometrical ratio—Examples of great powers of increase of animals—Rapid increase and wide spread of plants—Great fertility not essential to rapid increase—Struggle between closely allied species most severe—The ethical aspect of the struggle for existence.
There is perhaps no phenomenon of nature that is at once so important, so universal; and so little understood, as the struggle for existence continually going on among all organised beings. To most persons nature appears calm, orderly, and peaceful. They see the birds singing in the trees, the insects hovering over the flowers, the squirrel climbing among the tree-tops, and all living things in the possession of health and vigour, and in the enjoyment of a sunny existence. But they do not see, and hardly ever think of, the means by which this beauty and harmony and enjoyment is brought about. They do not see the constant and daily search after food, the failure to obtain which means weakness or death; the constant effort to escape enemies; the ever-recurring struggle against the forces of nature. This daily and hourly struggle, this incessant warfare, is nevertheless the very means by which much of the beauty and harmony and enjoyment in nature is produced, and also affords one of the most important elements in bringing about the origin of species. We must, therefore, devote some time to the consideration of its various aspects and of the many curious phenomena to which it gives rise.
It is a matter of common observation that if weeds are allowed to grow unchecked in a garden they will soon destroy a number of the flowers. It is not so commonly known that if a garden is left to become altogether wild, the weeds that first take possession of it, often covering the whole surface of the ground with two or three different kinds, will themselves be supplanted by others, so that in a few years many of the original flowers and of the earliest weeds may alike have disappeared. This is one of the very simplest cases of the struggle for existence, resulting in the successive displacement of one set of species by another; but the exact causes of this displacement are by no means of such a simple nature. All the plants concerned may be perfectly hardy, all may grow freely from seed, yet when left alone for a number of years, each set is in turn driven out by a succeeding set, till at the end of a considerable period—a century or a few centuries perhaps—hardly one of the plants which first monopolised the ground would be found there.
Another phenomenon of an analogous kind is presented by the different behaviour of introduced wild plants or animals into countries apparently quite as well suited to them as those which they naturally inhabit. Agassiz, in his work on Lake Superior, states that the roadside weeds of the northeastern United States, to the number of 130 species, are all European, the native weeds having disappeared westwards; and in New Zealand there are no less than 250 species of naturalised European plants, more than 100 species of which have spread widely over the country, often displacing the native vegetation. On the other hand, of the many hundreds of hardy plants which produce seed freely in our gardens, very few ever run wild, and hardly any have become common. Even attempts to naturalise suitable plants usually fail; for A. de Candolle states that several botanists of Paris, Geneva, and especially of Montpellier, have sown the seeds of many hundreds of species of hardy exotic plants in what appeared to be the most favourable situations, but that, in hardly a single case, has any one of them become naturalised. Even a plant like the potato—so widely cultivated, so hardy, and so well adapted to spread by means of its many-eyed tubers—has not established itself in a wild state in any part of Europe. It would be thought that Australian plants would easily run wild in New Zealand. But Sir Joseph Hooker informs us that the late Mr. Bidwell habitually scattered Australian seeds during his extensive travels in New Zealand, yet only two or three Australian plants appear to have established themselves in that country, and these only in cultivated or newly moved soil.
These few illustrations sufficiently show that all the plants of a country are, as De Candolle says, at war with each other, each one struggling to occupy ground at the expense of its neighbour. But, besides this direct competition, there is one not less powerful arising from the exposure of almost all plants to destruction by animals. The buds are destroyed by birds, the leaves by caterpillars, the seeds by weevils; some insects bore into the trunk, others burrow in the twigs and leaves; slugs devour the young seedlings and the tender shoots, wire-worms gnaw the roots. Herbivorous mammals devour many species bodily, while some uproot and devour the buried tubers.
In animals, it is the eggs or the very young that suffer most from their various enemies; in plants, the tender seedlings when they first appear above the ground. To illustrate this latter point Mr. Darwin cleared and dug a piece of ground three feet long and two feet wide, and then marked all the seedlings of weeds and other plants which came up, noting what became of them. The total number was 357, and out of these no less than 295 were destroyed by slugs and insects. The direct strife of plant with plant is almost equally fatal when the stronger are allowed to smother the weaker. When turf is mown or closely browsed by animals, a number of strong and weak plants live together, because none are allowed to grow much beyond the rest; but Mr. Darwin found that when the plants which compose such turf are allowed to grow up freely, the stronger kill the weaker. In a plot of turf three feet by four, twenty distinct species of plants were found to be growing, and no less than nine of these perished altogether when the other species were allowed to grow up to their full size.
But besides having to protect themselves against competing plants and against destructive animals, there is a yet deadlier enemy in the forces of inorganic nature. Each species can sustain a certain amount of heat and cold, each requires a certain amount of moisture at the right season, each wants a proper amount of light or of direct sunshine, each needs certain elements in the soil; the failure of a due proportion in these inorganic conditions causes weakness, and thus leads to speedy death. The struggle for existence in plants is, therefore, threefold in character and infinite in complexity, and the result is seen in their curiously irregular distribution over the face of the earth. Not only has each country its distinct plants, but every valley, every hillside, almost every hedgerow, has a different set of plants from its adjacent valley, hillside, or hedgerow—if not always different in the actual species yet very different in comparative abundance, some which are rare in the one being common in the other. Hence it happens that slight changes of conditions often produce great changes in the flora of a country. Thus in 1740 and the two following years the larva of a moth (Phalaena graminis) committed such destruction in many of the meadows of Sweden that the grass was greatly diminished in quantity, and many plants which were before choked by the grass sprang up, and the ground became variegated with a multitude of different species of flowers. The introduction of goats into the island of St. Helena led to the entire destruction of the native forests, consisting of about a hundred distinct species of trees and shrubs, the young plants being devoured by the goats as fast as they grew up. The camel is a still greater enemy to woody vegetation than the goat, and Mr. Marsh believes that forests would soon cover considerable tracts of the Arabian and African deserts if the goat and the camel were removed from them. Even in many parts of our own country the existence of trees is dependent on the absence of cattle. Mr. Darwin observed, on some extensive heaths near Farnham, in Surrey, a few clumps of old Scotch firs, but no young trees over hundreds of acres. Some portions of the heath had, however, been enclosed a few years before, and these enclosures were crowded with young fir-trees growing too close together for all to live; and these were not sown or planted, nothing having been done to the ground beyond enclosing it so as to keep out cattle. On ascertaining this, Mr. Darwin was so much surprised that he searched among the heather in the unenclosed parts, and there he found multitudes of little trees and seedlings which had been perpetually browsed down by the cattle. In one square yard, at a point about a hundred yards from one of the old clumps of firs, he counted thirty-two little trees, and one of them had twenty-six rings of growth, showing that it had for many years tried to raise its head above the stems of the heather and had failed. Yet this heath was very extensive and very barren, and, as Mr. Darwin remarks, no one would ever have imagined that cattle would have so closely and so effectually searched it for food.
In the case of animals, the competition and struggle are more obvious. The vegetation of a given district can only support a certain number of animals, and the different kinds of plant-eaters will compete together for it. They will also have insects for their competitors, and these insects will be kept down by birds, which will thus assist the mammalia. But there will also be carnivora destroying the herbivora; while small rodents, like the lemming and some of the field-mice, often destroy so much vegetation as materially to affect the food of all the other groups of animals. Droughts, floods, severe winters, storms and hurricanes will injure these in various degrees, but no one species can be diminished in numbers without the effect being felt in various complex ways by all the rest. A few illustrations of this reciprocal action must be given.
Illustrative Cases of the Struggle for Life.
Sir Charles Lyell observes that if, by the attacks of seals or other marine foes, salmon are reduced in numbers, the consequence will be that otters, living far inland, will be deprived of food and will then destroy many young birds or quadrupeds, so that the increase of a marine animal may cause the destruction of many land animals hundreds of miles away. Mr. Darwin carefully observed the effects produced by planting a few hundred acres of Scotch fir, in Staffordshire, on part of a very extensive heath which had never been cultivated. After the planted portion was about twenty-five years old he observed that the change in the native vegetation was greater than is often seen in passing from one quite different soil to another. Besides a great change in the proportional numbers of the native heath-plants, twelve species which could not be found on the heath flourished in the plantations. The effect on the insect life must have been still greater, for six insectivorous birds which were very common in the plantations were not to be seen on the heath, which was, however, frequented by two or three different species of insectivorous birds. It would have required continued study for several years to determine all the differences in the organic life of the two areas, but the facts stated by Mr. Darwin are sufficient to show how great a change may be effected by the introduction of a single kind of tree and the keeping out of cattle.
The next case I will give in Mr. Darwin’s own words: “In several parts of the world insects determine the existence of cattle. Perhaps Paraguay offers the most curious instance of this; for here neither cattle nor horses nor dogs have ever run wild, though they swarm southward and northward in a feral state; and Azara and Rengger have shown that this is caused by the greater numbers, in Paraguay, of a certain fly which lays its eggs in the navels of these animals when first born. The increase of these flies, numerous as they are, must be habitually checked by some means, probably by other parasitic insects. Hence, if certain insectivorous birds were to decrease in Paraguay, the parasitic insects would probably increase; and this would lessen the number of the navel-frequenting flies—then cattle and horses would become feral, and this would greatly alter (as indeed I have observed in parts of South America) the vegetation: this again would largely affect the insects, and this, as we have just seen in Staffordshire, the insectivorous birds, and so onward in ever-increasing circles of complexity. Not that under nature the relations will ever be as simple as this. Battle within battle must be continually recurring with varying success; and yet in the long run the forces are so nicely balanced, that the face of nature remains for a long time uniform, though assuredly the merest trifle would give the victory to one organic being over another.”
