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Darwinism (1889).
by Alfred Russel Wallace.
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._ 30. The statement as to the fulmar petrel, which Professor A.
Newton a.s.sures me is erroneous, has been modified.
_P._ 34. A note is added as to Darwin's statement about the missel and song-thrushes in Scotland.
_P._ 172. 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 ill.u.s.trate, 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 ill.u.s.trate 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 s.e.xual 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 ill.u.s.trations 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 ill.u.s.trate 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 s.e.xual 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.
G.o.dALMING, _March 1889_.
CHAPTER I
WHAT ARE "SPECIES," AND WHAT IS MEANT BY THEIR "ORIGIN"
Definition of species--Special creation--The early Trans.m.u.tationists--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 t.i.tle 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 ma.s.s 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 a.n.a.logy 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 circ.u.mstances, from another animal. It propagates, 'after its kind,' individuals perfectly resembling the parent; its peculiarities, therefore, are permanent."[1]
To ill.u.s.trate 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 versa_. 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 Trans.m.u.tationists_.
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 pa.s.sages 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 circ.u.mstances, 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_.[2]
Professor Aga.s.siz, 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 circ.u.mstance that species occur within definite limits where no obstacles prevent their wider distribution leads to the further inference that these limits were a.s.signed 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."[3]
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 "trans.m.u.tation 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 "trans.m.u.tation" 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 a.s.suredly 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 cla.s.sified 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 Cla.s.ses 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 cla.s.ses, 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 s.p.a.ce 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 pa.s.sage 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 dispa.s.sionate 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."