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