oN THE ORIGIN OF SPECIES.“But with regard to the material world, we can at least go so far as this—we can perceive that events are brought about not by insulated in- terpositions of Divine power, exerted in each pasticular case, but by the establishment of general laws.” W. Warwnit: Bridgewater Treatise. “To conclude, therefore, let no man out of a weak conceit of sobriety, or an ill-applied moderation, think or maintain, tbat a man can search too far or be too well studied in the book of God's word, or in the book of God’s works; divinity or philosophy ; but rather let men endeavour an endless progress or proficience in both.” Bacon: Advancement of Learning. “The only distinct meaning of the word ‘natural’ is stated, fized, or ‘esttled ; since what is natural as much requires and presupposes an intelli- gent agent to render it #0, é.¢. to effect it continually or at stated times, an ‘what is supernatural or miraculous does to effect it for once. Borur: Analogy of Revealed Religion, Bromley, Kent, October Ist, 1859.ON THE ORIGIN OF SPECIES MEANS OF NATURAL SELECTION, PRESERVATION OF FAVOURED RAOES IN THE STRUGGLE FOR LIFE. BY CHARLES DARWIN, M.A., YELLOW OF THE XOTAL, GEOLOGICAL, LINZZAX, ETO. SOOIRTIES ; avrdon oF “J0URNAL OF EESEARONES DUNIXO x. x, 8, BEAGLE’ YorAGR ROUND i ‘TER WORLD.” 4. EDITION, RETIMED 49D AVOMENTED BY THE AUTHOR, NEW YORK: D. APPLETON AND COMPANY, 43 & 445 BROADWAY. M.DC00.1x1,KE 73%2PREFACE. CONTRIBUTED BY THE AUTHOR TO THIS AMERICAN EDITION, I wu here attempt to give a brief, but I fear imper- fect, sketch of the progress of opinion on the Origin of Species. The great majority of naturalists have believed that species were immutable productions and have been separately created: this view has been ably maintained by many authors. A few naturalists, and several who have not particularly studied natural history, believe, on the other hand, that species undergo modification, and that the existing forms of life have descended by true gen- eration from pre-existing forms. Passing over authors of the classical period, and likewise Demaillet and Buffon, with whose writings I am not familiar, Lamarck was the first man, whose view that species undergo change ex- cited much attention, This justly celebrated naturalist published his Philosophie Zoologique in 1809, and his In- - troduction to his Hist. Nat. des animaux sans Vertdbres in 1815, in which works he upholds the doctrine that spe- cies are descended from each other. He seems to have been chiefly led to this conclusion by the difficulty of dis- tinguishing species and varieties,—by the almost perfect gradation of the forms in certain groups, and by the anal- ogy of domestic productions. With respect to the means of modification, he attributed something to the action ofvi PREFACE. external conditions, something to the crossing of already existing forms, and much to use and disuse or the effects of habit. To this latter agency he seems to attribute all the beautiful adaptations in nature,—such as the long neck of the giraffe for browsing on the branches of trees.* But he likewise believed in a law of progressive develop- ment; and as all the forms of life thus tended to progress, in order to account for the presence of very simple pro- ductions at the present day, he maintained that such forms ‘were now spontaneously generated. . Geoffroy Saint Hilaire, as is stated in his Life by his Son, as early as 1795, suspected that what we call species are various degenerations of the same type. It was not until 1828 that he published his conviction that the same forms have not been perpetuated since the origin of all things. Geoffroy seems to have relied chiefly on the con- ditions of life, or the “monde ambiant,” as the cause of change ; but he was cautious, and, as his son says, “ C’est donc un problame a reserver entidrement a avenir, sup- posé méme que Vavenir doive avoir prise sur lui.” In England, the Hon, and Rev. W. Herbert, after- wards Dean of Manchester, in his work on the Amarylli- dacem (1887, p. 19, 389), declares that “horticultural experiments have established, beyond the possibility of refutation, that botanical species are only a higher and more permanent class of varieties.” He extends the same view to animals. The Dean believes that single species of each genus were created in an originally highly plastic condition, and that these by intercrossing and by varia- tion have produced all our existing species. It is curious how completely my grandfather, Dr, Erasmus Darwin, anticipated these erroneous views in his Zoonomia (vol. i. pp. 500-510), pub- lished in 1794,PREFACE. vil In 1848-44, Prof. Haldeman (in the Boston (U. 8.) Journal of Nat. Hist., vol. iv., p. 468) has ably given the arguments for and against the hypothesis of the develop- ment and modification of species: he seems to me to lean towards the side of change. The Vestiges of Creation appeared in 1844. In tho Jast or tenth and much improved edition (1853, p. 155), the anonymous author says: “The proposition determined on after much consideration is, that the several series of animated beings, from the simplest and oldest up to the highest and most recent, are, under the providence of God, the results, first, of an impulse which has been imparted to the forms of life, advancing them, in definite times, by generation, through grades of organisation terminating in the highest dicotyledons and vertebrata, these grades be- ing few in number, and generally marked by intervals of organic character which we find to be a practical difficulty in ascertaining affinities; second, of another impulse con- nected with the vital forces, tending in the course of gen- erations to modify organic structures in accordance with external circumstances, as food, the nature of the habitat and the meteoric agencies, these being the ‘adaptations’ of the natural theologian.” The author apparently be- Hieves that organisation progresses by sudden leaps; but that the effects produced by the conditions of life are gradual. The author argues with much force on general grounds that species are not immutable productions. But, T cannot see how the two supposed “impulses” account in a scientific sense for the numerous and beautiful co- adaptations, which we gee throughout nature ;—I cannot see that we thus gain any insight how, for instance, a woodpecker has become adapted to its peculiar habits of life. The work, from its powerful and brilliant style,though displaying in the earlier editions little accurate knowledge and a great want of scientific caution, immedi- ately had a very wide circulation. In my opinion, it has done excellent service in calling in this country attention to the subject, and in removing prejudices. In 1846, the veteran geologist, M. J. d’Omalius @’Hal- loz, published in an excellent, though short, paper (Bulle- tins de Acad. Roy. Bruxelles, tom. xiii., p. 581), his opin- ion that it is more probable that new species have been produced by descent with modification, than that they have been separately created: the author first promul- gated this opinion in 1831. M. Isidore Geoffroy Saint Hilaire, in his Lectures de- livered in 1850 (of which a Résumé appeared in the Re- vue et Mag. de Zoolog., Jan. 1851), briefly gives his reasons for believing that specific characters “ sont fixes, pour chaque espéce, tant qu’elle se perpétue au milieu des mémes circonstances, ils se modifient, si les circonstances ambiantes viennent 4 changer.” “En résumé, V’observa- tion des animaux sauvages démontre déja la variabilité Uimitée des espdces. Les expériences sur les animaux sau- vages devenus domestiques, et sur les animaux domes- tiques redevenus sauvages, la démontrent plus clairement encore. Ces mémes expériences prouvent, de plus, que Jes differences produites peuvent étre de valeur géné- igue.?? Mr. Herbert Spencer, in an Essay (originally pub- lished in the Leader, March, 1852, and republished in his Essays, 1858), has contrasted the theories of the creation and development of organic beings with remarkable skill and force. He argues from the analogy of domestic pro- ductions, from the changes which the embryos of many species undergo, from the difficulty of distinguishing spe-PREFACE. ix cics and varieties, and from the principle of general gra- dation, that species have been modified ; and he attributes the modification to the change of circumstances. The author (1855) has treated Psychology on the principle of the necessary acquirement of each mental power and ca- pacity by gradation. In 1852 (Revue Horticole, p. 102), M. Naudin, a dis- tinguished botanist,* has expressly stated his belief that species aro formed in an analogous manner as varieties are under cultivation; and the latter process he attributes to man’s power of selection. But he does not show how se- lection can act under nature. He believes, like Dean Herbert, that species when nascent were more plastic. He lays weight on what he calls the principle of finality, “puissance mystérieuse, indéterminée ; fatalité pour les uns; pour Ids autres, volonté providentielle, dont action incessante sur les étres vivants détermine, & toutes les 6poques de l’existence du monde, la forme, le volume ot la durée de chacun d’eux, en raison de sa destinée dans Vordre de choses dont il fait partie. C'est cette puissance qui harmonise chaque membre a l’ensemble en l’appropri- ant a la fonction qu’il doit remplir dans Porganisme géné- ral de la nature, foriction qui est pour Ini ea raison d’étre.” In 1853, a celebrated geologist, Count Keyserling (Bul- letin de'la Soc. Geolog., 2d ser., tom. x., p. 357) suggested that as new diseases, supposed to have been caused by some miasma, have arisen and spread over the world, so at-certain periods the germs of existing species may have been chemically affected by circumambient molecules of 8 particular nature, and thus have given rise to new forms. ® M, Lecoq, another French botanist, entertains, I believe, analo views on the modifcation and descent of species, -x PREVACE, The “ Philosophy of Creation” has been treated in an admirable manner by the Rev. Baden Powell, in his Es- says on the Unity of Worlds, 1855. Nothing can be more striking than the manner in which he shows that the in- troduction of new species is “a regular, not a casual, phenomenon,” or, as Sir John Herschel expresses it, “a natural in contradistinction to a miraculous process.” I think this work can hardly have failed to have produced a great effect in every philosophical mind. The third volume of the Journal of the Linnean So- ciety (August, 1858) contains papers by Mr. Wallace and myself, in which, as stated in the introductory remarks to this volume, the theory of Natural Selection is promul- gated. In June, 1859, Professor Huxley gave a lecture be- fore the Royal Institution on the Persistent Types of Ani- mal Life. ‘Referring to such cases, he remarks: “It is difficult to comprehend the meaning of such facts as these, if we suppose that each species of animal and plant, or each great type of organisation, was formed and placed upon the surface of the globe, at long intervals, by a dis- tinct act of creative power ; and it is well to recollect that such an assumption is as unsupported by tradition or reve- lation as it is opposed to the general analogy of nature. If, on the other hand, we view ‘ Persistent Types’ in rela- tion to that hypothesis which supposes the species living at any time to be the result of the gradual modification of pre-existing species —a hypothesis which, though un- proven, and sadly damaged by some of its supporters, is yet the only one to which physiology lends any counte- nance—their existence would seem to show that the amount of modification which living beings have under- gone during geological time is but very small in relationPREFACE. a to the whole series of changes which they have suf- fered.” In November, 1859, the first edition of this work was published. In December, 1859, Dr. Hooker published his Introduction to the Tasmanian Flora: in the first part of this admirable essay he admits the truth of the descent and modification of species; and supports this doctrine by many original and valuable observations. Cuantes Dazwor Down, Buouzrr, Kxxr, Feb. 1860.CONTENTS CHAPTER I. ‘VARIATION UXDEE DOMESTICATION. (Causes of Variability—Effects of Habit—Correlation of Growth—Inberitance—Char- ‘acter of Domestic Varletiee—Dificulty of distinguishing between Varieties ‘and Species—Origin of Domestio Varieties from one or more Speoiee—Domestic Pigeons, their Differences and Origin—Principle of Belection anciently followed, ite Effecte—Mothodical and Unconscious Selection—Unknown Origin of our Domes ‘tio Produtions—Circamstances favourable to Man's power of Selection, . 1d CHAPTER II. VARIATION UNDER NATURE. ‘Varlabillty—Indlvidual differencee—Doubtful specles—Wide ranging, mush diffused, ‘and common species vary most—Species of the larger genera in any country vary ‘more than the species of the smaller geners—Many of the species of the larger sroeraresmmle variates tn being very clos, bus wnagually related to eteh other, and in having restricted ranges, =. rr) OHAPTER III. STRUGGLE FoR EXIsTAXCE. ‘Bears on natural selection—The term used in a wide sense—Geometrical powers of increase—Repid increase of naturalised animals and plante—Natare of the checks to inerease—Compotition untversal—Eifecta of climate—Protection from the num ‘bor of individuale—Complex relations of all animals and plants thronghout nature ‘Struggle for life most severe between individuals and varieties of the eame spe- ‘cles;; often severe between species of the same genus—The relation of organism toorganism the most important of allrelations, . . - ss CHAPTER Iv. NATURAL SELEOTION. ‘oatural Selection—its power compared with man’s selection—its power on charscters Of trifling importance—ita power at all ages and on both sexes—Sexual Belection—6 conTENTs. On the generality of intercrouses betwoen individuals of the same spectes—Cir cumstances favourable and unfavourable to Natural Selection, namely, intereroms- ng, isolation, number of individuals—-Slow action—Extinotion caused by Natural Belection—Divergetce of Character, related to the diversity of inhabitants of any small ares, and fo naturallnation— Action of Natural Belection, through Divergence ‘of Character and Extinction, on the descendants from a common parent—Eaplains the Grouping of allorganto beings, . - ee we CHAPTER Y. Laws oF VARIATION. Effects of external conditlons—Use and disuse, combined with natural selection j ‘organs of flight and of vislon—Acelimatisation—Correlation of growth—Compem- sation and economy of growth—False correlations—Maltiple, rudimentary, and lowly organised structures varlable—Parta developed in an unusual manner are Iighly variable : spooiflc characters more variable than generis: secondary sexual characters variable—Speoles of the same genus vary in an analogous manner—Re- versions tolong lost characters—Summary, = ss sss 10 CHAPTER VI. DIFFICULTIES OX THEORY. Difficulties on the theory of descent with modlffcation—Transitlons—Absence oF ity of transitional varieties—Traneitions in habits of llfe—Diversified habits in 0 same spectes—Spectes with habite widely different from those of thelr allloe— Organs of extreme perfection—Means of transition—Cases of difficulty—Naturs 100 faott saltum—Organs of small tmportance—Osgans not {n all cases absolutely perfect—The law of Unity of Type and of the Conditions of Exlatence embraced ‘by the theory of Natural Selection, » =. eee 136 CHAPTER VII. rxstrxor. Instincts comparable with habits, but different in thelr origin—Instincta graduated— Apbides and ante—Instincts varlable—Domestio instincts, thelr origin—Nataral {nstinete of the cuckoo, ostrich, and parasitic boes—Slave-making ants—Hivo-beo, ta cell-making instinet—Dificulties on the theory of the Natural Selection of instinote—Neuter or sterile insects—Summary, . . 5 ss 188 CHAPTER VIII. mYrBRrprs: ‘Distinction between the sterility of first crosses and of hybrids—Sterility various in ogree, not universal, affected by close interbreeding, removed by domestication— Laws governing the sterility of bybrids—Sterility not a special endowment, but Incidental on other differences—Csusos of the sterility of first crosses and of bybride—Parallelism botwoen the effects of changed conditions of life and crossComPanns: 2 ing—-Fertility of varieties when erossed and of their mongrel offspring not enh ‘Yorsal—Hybrids and mongrels compared independently of their fertility—Sam en ee ns, CHAPTER Ix, (ON THE IMPERFECTION OF THE GEOLOGICAL RECORD. On the absence of intermediate varieties at the present day—On the nature of extinet intermediate varieties ; on their number—On the vast lapse of time, as tnferred from the rate of deposition and of denudstion—On the poorness of our palmovto- logical collections—On the intermittence of geological formations—On the absence of intermediate varieties in apy one formation—On the sudden appearance of groups of species—On thelr sudden appearance in the lowest known fosaliferous CHAPTER X. (ON THE GEOLOGICAL SUCCESSION OF ORGANIC BEIXGS. ‘On the slow and successive appearance of new species—On thelr different rates of ‘change—Species once lost do not reappear—Groups of species follow the same gen- coral rules in thelr appearance and disappearance as do single speclee—On Extine- ‘tioo—On simultaneous changes in the forms of life throughout the world—On the ‘affinities of extinct spectos to each other and to living species—On the state of de- ‘velopment of ancient forme—On the succession of the same types within the same ‘areas—Summary of preceding and present chapters, 9. =. ss 1B CHAPTER XI. G@roGRAPSICAL DisTRIBUTION. ‘Present distribution cannot be accounted for by differences in physical conditions—Im- portance of berriers—Affinity of the produetions of the same continent—Centres of crestion—Means of dispersal, by changes of elimate and of the level of the land, and by occasional means—Dispersal during the Glacial period co-extensive withtheworl, . eee OR CHAPTER XII. ‘qpocRAPmicat, pisrarsuTr0K—Continued. ‘Distribution of fresh-water ‘productions—On the inhabitants of oceanto falands—Ab- sence of Batrachians and of terrestrial mammals—On the relation of the inhabit ‘ante of lelands to those of the nearest mainland—On colonisation from the nearest source with subsequent modification—Summary of the last and present chap- CHAPTER XIII. MUTUAL AFTOOTIES oF ORGANIC BEINGS: MORPHOLOGY: muBRYOLOGY: RUDI- d MENTARY ORGANS. Ciassrrioartox, groups subordinate to groupe—Nataral system—Roles and aiftcal- ‘tes in classification, explained on the theory of descent with modiffcation—Clasch{fleation of varleties—Descent always used in clasaifieation— Analogies] or adaptive characters—Affinities, general, complex and radiating—Kxtinotion separates and defines groupe—Monrnorocr, between members of the same class, between parts of the same individual—Exsarozoor, laws of, explained by variations not super- ‘Tesing at an easty ng, and being naried at correponding age—Rivoruenrant onoaxs; their origin explained—Summary, ar) CHAPTER XIV. RECAPIFULATION AND COXCLUSION. Recapitalation of the dificalties on the theory of Natural Selection—Recapttulation of the general and special elroumstances in its favour—Canses of the general ‘ollef in the tmmutability of species—How far the theory of natural selection may ‘be extended—Etfecta of its adoption on the atndy of Natural History—Conclnding Wr, ee a Inetruction to Binder.—Tbe Diagram to front page 198,ON THE ORIGIN OF SPECIES. INTRODUOTION. ‘Wen on board H. M.S. ‘Beagle’ as naturalist, I was much struck with certain facts in the distribution of the inhabitants of South America, and in the geological rela- tions of the present to the past inhabitants of that con- eae See ae eos to me to aoe some light on the origin of species—that m of mysteries, as it has been called by one of our greatend philosophers On my return home, it occurred to me, in 1887, that some- thing might perhaps be made out on this question by patiently secumalating and reflecting on all sorts of facts which could ibly have any bearing on it. After five years’ work I allowed myself to spec late on the subject, and drew np some short notes; these I enlarged in 1844 into a sketch of the conclusions, which then seemed to me probable: from that period to the present day I have stead- ily pursued the same object. I hope that I may be ox- cused for entering on these I details, as I give them to show that I have not been hasty in coming to a decision. My work is now nearly finished ; but as it will take me two or three more years to complete it, and as my health is far from strong, I have been urged to publ this Abstract. I have more especially been induced to do this, as Mr. Wallace, who is now studying the natural history of the Malay archipelago, has arrived at almost exactly the same general conclusions that I have on the of species. Last year he sent to me a memoir on this subjest, with a request that I would forward it to Sir10 INTRODUCTION Charles Lyell, who sent it to the Linnean Society, and it is published in the third volume of the Journal of that Society. Sir C. Lyell and Dr. Hooker, who both knew of my work—the latter having read my sketch of 1844 —honoured me by thinking it advisable to publish, with Mr. Wallace’s excellent memoir, some brief extracts from my manuscripts. This Abstract, which I now publish, must necessarily be imperfect. I cannot here give references and author- ities for my several statements; and I must trust to the reader reposing some confidence in my securacy, No doubt errors will have crept in, though I hope I have always been cautious in trusting to good authorities alone. Ican here give only the general conclusions at which I have arrived, with a few facts in illustration, but which, L hope, in most cases will suffice. No one can feel more sensible than I do of the necessity of hereafter publishing in detail all the facts, with references, on which my con- clusions have been grounded ; and I hope in a future work to do this, For I'am well aware that scarcely a single joint is discussed in this volume on which facts cannot e adduced, often apparently leading to conclusions di- rectly opposite to those at which I have arrived. A fair result can be obtained only by fully stating and balancing the facts and arguments on both sides of each question ; and this cannot possibly be here done. I much regret that want of space prevents my having the satisfaction of acknowledging the generous assistance which I have received from very many naturalists, some of them personally unknown to ‘me. I cannot, however, let this opportunity pass without expressing my deep obligationd to Dr. Hobker, who for the last fiftecs years has aided me in every possible way by his large stores of knowledge and his excellent judgment. In considering the Origin. of Species, it is quite con- ceivable that a naturalist, reflecting on the mutual affini- ties of organic beings, on their embryological relations, their geographical distribution, geological succession, and other such facts, might come to the conclusion that each species had not been independently created, but had de-INTRODUCTION. uu ascended, like varieties, from other i Nevertheless such a conclusion, even if well founded, would be unsat- isfactory, until it could be shown how the innumerable species inhabiting this world have been modified, so as to acquire that perfection of structure and coadaptation which most justly excites our admiration. Naturalists continually refer to external conditions, such as climate, food, &c., as the only possible cause of variation. In one very limited sense, a8 we shall hereafter see, this may be true ; but it is preposterous to attribute to mere external conditions, the structure, for instance, of the woodpecker, with its feet, tail, beak, and tongue, 0 admirably adapted to catch insects under the bark of trees. In the case of the misseltoe, which draws its nourishment from certain trees, which has seeds that must be transported by certain birds, and which has flowers with separate sexes abso- Iutely requiring the agency of certain insects to bring pol- len from one flower to the other, it is equally preposterous to account for the structure of this parasite, with its rela- tions to several distinct organic beings, by the effects of external conditions, or of habit, or of the volition of the Piatt ie therefore, of the highest importance to gat it fore, e importance a clear inright into the means of modification and coudapta- tion. At the commencement of my observations it seemed to me probable that a careful study of domesticated ani- mals and of cultivated plants would offer the best chance of making out this obscure problem. Nor have I been disappointed ; in this and in all other perplexing cases I have invariably found that our knowledge, imperfect though it be, of variation under domestication, afforded the best and safest clue. I may venture to express my conviction of the high value of such studies, although they have been very commonly neglected by naturalists. From these considerations, I shall devote the first chapter of this Abstract to Variation under Domestica- tion, We shall thus see that a large amount of hereditary modification is at least possible; and, what is equally or more important, we shall seo how great is the wer of man in accumulating by his Selection successive slight