charles darwin

On the Origin of Species

First published 1859 · sixth edition, 1872
i
Epigrams
BUTLER: Analogy of Revealed Religion.
ii
Introduction
When on board H.M.S. 'Beagle,' as naturalist, I was much struck with certain facts in the distribution of the inhabitants of South America,…

Variation under Domestication

iii
Causes of Variability
When we look to the individuals of the same variety or sub-variety of our older cultivated plants and animals, one of the first points…
iv
Effects of Habit and the Use or Disuse of Parts; Correlated Variation; Inheritance.
Changed habits produce an inherited effect as in the period of the flowering of plants when transported from one climate to another. With…
v
Character of Domestic Varieties; difficulty of distinguishing between Varieties and Species; origin of Domestic Varieties from one or more Species.
When we look to the hereditary varieties or races of our domestic animals and plants, and compare them with closely allied species, we…
vi
Breeds of the Domestic Pigeon, their Differences and Origin.
Believing that it is always best to study some special group, I have, after deliberation, taken up domestic pigeons. I have kept every…
vii
Principles of Selection anciently followed, and their Effects.
Let us now briefly consider the steps by which domestic races have been produced, either from one or from several allied species. Some…
viii
Unconscious Selection.
At the present time, eminent breeders try by methodical selection, with a distinct object in view, to make a new strain or sub-breed,…
ix
Circumstances favourable to Man's power of Selection.
I will now say a few words on the circumstances, favourable or the reverse, to man's power of selection. A high degree of variability is…

Variation under Nature

Before applying the principles arrived at in the last chapter to organic beings in a state of nature, we must briefly discuss whether these latter are subject to any variation. To treat this subject properly, a long catalogue of dry facts ought to be given; but these I shall reserve for a future work. Nor shall I here discuss the various definitions which have been given of the term species. No one definition has satisfied all naturalists; yet every naturalist knows vaguely what he means when he speaks of a species. Generally the term includes the unknown element of a distinct act of creation. The term "variety" is almost equally difficult to define; but here community of descent is almost universally implied, though it can rarely be proved. We have also what are called monstrosities; but they graduate into varieties. By a monstrosity I presume is meant some considerable deviation of structure, generally injurious, or not useful to the species. Some authors use the term "variation" in a technical sense, as implying a modification directly due to the physical conditions of life; and "variations" in this sense are supposed not to be inherited; but who can say that the dwarfed condition of shells in the brackish waters of the Baltic, or dwarfed plants on Alpine summits, or the thicker fur of an animal from far northwards, would not in some cases be inherited for at least a few generations? And in this case I presume that the form would be called a variety.

It may be doubted whether sudden and considerable deviations of structure, such as we occasionally see in our domestic productions, more especially with plants, are ever permanently propagated in a state of nature. Almost every part of every organic being is so beautifully related to its complex conditions of life that it seems as improbable that any part should have been suddenly produced perfect, as that a complex machine should have been invented by man in a perfect state. Under domestication monstrosities sometimes occur which resemble normal structures in widely different animals. Thus pigs have occasionally been born with a sort of proboscis, and if any wild species of the same genus had naturally possessed a proboscis, it might have been argued that this had appeared as a monstrosity; but I have as yet failed to find, after diligent search, cases of monstrosities resembling normal structures in nearly allied forms, and these alone bear on the question. If monstrous forms of this kind ever do appear in a state of nature and are capable of reproduction (which is not always the case), as they occur rarely and singly, their preservation would depend on unusually favourable circumstances. They would, also, during the first and succeeding generations cross with the ordinary form, and thus their abnormal character would almost inevitably be lost. But I shall have to return in a future chapter to the preservation and perpetuation of single or occasional variations.

x
Individual Differences
The many slight differences which appear in the offspring from the same parents, or which it may be presumed have thus arisen, from being…
xi
Doubtful Species
The forms which possess in some considerable degree the character of species, but which are so closely similar to other forms, or are so…
xii
Wide-ranging, much-diffused, and common Species vary most.
Guided by theoretical considerations, I thought that some interesting results might be obtained in regard to the nature and relations of…
xiii
Species of the Larger Genera in each Country vary more frequently than the Species of the Smaller Genera.
If the plants inhabiting a country as described in any Flora, be divided into two equal masses, all those in the larger genera (i.e., those…
xiv
Many of the Species included within the Larger Genera resemble Varieties in being very closely, but unequally, related to each other, and in having restricted ranges.
There are other relations between the species of large genera and their recorded varieties which deserve notice. We have seen that there is…
xv
Summary.
Finally, varieties cannot be distinguished from species,— except, first, by the discovery of intermediate linking forms; and, secondly, by…

Struggle for Existence

xvi
The Bearing of Struggle for Existence on Natural Selection
Before entering on the subject of this chapter, I must make a few preliminary remarks to show how the struggle for existence bears on…
xvii
The Term, Struggle for Existence, Used in a Large Sense
I should premise that I use this term in a large and metaphorical sense, including dependence of one being on another, and including (which…
xviii
Geometrical Ratio of Increase.
A struggle for existence inevitably follows from the high rate at which all organic beings tend to increase. Every being, which during its…
xix
Nature of the Checks to Increase.
The causes which check the natural tendency of each species to increase are most obscure. Look at the most vigorous species; by as much as…
xx
Complex Relations of all Animals and Plants to Each Other in the Struggle for Existence.
Many cases are on record showing how complex and unexpected are the checks and relations between organic beings, which have to struggle…
xxi
Struggle for Life most severe between Individuals and Varieties of the same Species.
As the species of the same genus usually have, though by no means invariably, much similarity in habits and constitution, and always in…

Natural Selection; or the Survival of the Fittest

xxii
Natural Selection: Its Power Compared with Man’s Selection
How will the struggle for existence, briefly discussed in the last chapter, act in regard to variation? Can the principle of selection,…
xxiii
Sexual Selection.
Inasmuch as peculiarities often appear under domestication in one sex and become hereditarily attached to that sex, so no doubt it will be…
xxiv
Illustrations of the Action of Natural Selection, or the Survival of the Fittest.
In order to make it clear how, as I believe, natural selection acts, I must beg permission to give one or two imaginary illustrations. Let…
xxv
On the Intercrossing of Individuals.
I must here introduce a short digression. In the case of animals and plants with separated sexes, it is of course obvious that two…
xxvi
Circumstances favourable for the production of new forms through Natural Selection.
This is an extremely intricate subject. A great amount of variability, under which term individual differences are always included, will…
xxvii
Extinction caused by Natural Selection.
This subject will be more fully discussed in our chapter on Geology; but it must here be alluded to from being intimately connected with…
xxviii
Divergence of Character.
The principle, which I have designated by this term, is of high importance, and explains, as I believe, several important facts. In the…
xxix
The Probable Effects of the Action of Natural Selection through Divergence of Character and Extinction, on the Descendants of a Common Ancestor.
After the foregoing discussion, which has been much compressed, we may assume that the modified descendants of any one species will succeed…
xxx
On the degree to which organisation tends to advance.
Natural selection acts exclusively by the preservation and accumulation of variations, which are beneficial under the organic and inorganic…
xxxi
Convergence of Character.
Mr. H. C. Watson thinks that I have overrated the importance of divergence of character (in which, however, he apparently believes), and…
xxxii
Summary of Chapter.
If under changing conditions of life organic beings present individual differences in almost every part of their structure, and this cannot…

Laws of Variation

xxxiii
Effects of Changed Conditions
I have hitherto sometimes spoken as if the variations — so common and multiform with organic beings under domestication, and in a lesser…
xxxiv
Effects of the increased use and disuse of parts, as controlled by natural selection.
From the facts alluded to in the first chapter, I think there can be no doubt that use in our domestic animals has strengthened and…
xxxv
Acclimatisation.
Habit is hereditary with plants, as in the period of flowering, in the time of sleep, in the amount of rain requisite for seeds to…
xxxvi
Correlated variation.
I mean by this expression that the whole organisation is so tied together, during its growth and development, that when slight variations…
xxxvii
Compensation and Economy of Growth.
The elder Geoffroy and Goethe propounded, at about the same time, their law of compensation or balancement of growth; or, as Goethe…
xxxviii
Multiple, Rudimentary, and Lowly-organised Structures are Variable.
It seems to be a rule, as remarked by Is. Geoffroy St. Hilaire, both with varieties and species, that when any part or organ is repeated…
xxxix
A Part developed in any Species in an extraordinary degree or manner, in comparison with the same Part in allied Species, tends to be highly variable.
Several years ago I was much struck by a remark to the above effect made by Mr. Waterhouse. Professor Owen, also, seems to have come to a…
xl
Specific characters more variable than generic characters.
The principle discussed under the last heading may be applied to our present subject. It is notorious that specific characters are more…
xli
Summary.
Our ignorance of the laws of variation is profound. Not in one case out of a hundred can we pretend to assign any reason why this or that…

Difficulties of the Theory

xlii
Difficulties of the Theory of Descent with Modification
Long before the reader has arrived at this part of my work, a crowd of difficulties will have occurred to him. Some of them are so serious…
xliii
On the absence or rarity of transitional varieties.
As natural selection acts solely by the preservation of profitable modifications, each new form will tend in a fully-stocked country to…
xliv
On the Origin and Transition of Organic Beings with peculiar Habits and Structure.
It has been asked by the opponents of such views as I hold, how, for instance, could a land carnivorous animal have been converted into one…
xlv
Organs of extreme Perfection and Complication.
To suppose that the eye with all its inimitable contrivances for adjusting the focus to different distances, for admitting different…
xlvi
Modes of transition.
If it could be demonstrated that any complex organ existed, which could not possibly have been formed by numerous, successive, slight…
xlvii
Special difficulties of the theory of natural selection.
Although we must be extremely cautious in concluding that any organ could not have been produced by successive, small, transitional…
xlviii
Organs of little apparent Importance, as affected by Natural Selection.
As natural selection acts by life and death,— by the survival of the fittest, and by the destruction of the less well-fitted individuals,—…
xlix
Utilitarian Doctrine, how far true: Beauty, how acquired.
The foregoing remarks lead me to say a few words on the protest lately made by some naturalists against the utilitarian doctrine that every…
l
Summary: The Law of Unity of Type and of the Conditions of Existence embraced by the Theory of Natural Selection.
We have in this chapter discussed some of the difficulties and objections which may be urged against the theory. Many of them are serious;…
li
Miscellaneous Objections to the Theory of Natural Selection
I will devote this chapter to the consideration of various miscellaneous objections which have been advanced against my views, as some of…

Instinct

lii
Instincts Comparable with Habits, but Different in Their Origin
Many instincts are so wonderful that their development will probably appear to the reader a difficulty sufficient to overthrow my whole…
liii
Inherited Changes of Habit or Instinct in Domesticated Animals.
The possibility, or even probability, of inherited variations of instinct in a state of nature will be strengthened by briefly considering…
liv
Special Instincts.
We shall, perhaps, best understand how instincts in a state of nature have become modified by selection by considering a few cases. I will…
lv
Objections to the Theory of Natural Selection as applied to Instincts: Neuter and Sterile Insects.
It has been objected to the foregoing view of the origin of instincts that "the variations of structure and of instinct must have been…
lvi
Summary.
I have endeavoured in this chapter briefly to show that the mental qualities of our domestic animals vary, and that the variations are…

Hybridism

lvii
Distinction between the Sterility of First Crosses and of Hybrids
The view commonly entertained by naturalists is that species, when intercrossed, have been specially endowed with sterility, in order to…
lviii
Laws governing the Sterility of first Crosses and of Hybrids.
We will now consider a little more in detail the laws governing the sterility of first crosses and of hybrids. Our chief object will be to…
lix
Origin and Causes of the Sterility of first Crosses and of Hybrids.
At one time it appeared to me probable, as it has to others, that the sterility of first crosses and of hybrids might have been slowly…
lx
Reciprocal Dimorphism and Trimorphism.
This subject may be here briefly discussed, and will be found to throw some light on hybridism. Several plants belonging to distinct orders…
lxi
Fertility of Varieties when Crossed, and of their Mongrel Offspring, not universal.
It may be urged as an overwhelming argument that there must be some essential distinction between species and varieties inasmuch as the…
lxii
Hybrids and Mongrels compared, independently of their fertility.
Independently of the question of fertility, the offspring of species and of varieties when crossed may be compared in several other…
lxiii
Summary of Chapter.
First crosses between forms, sufficiently distinct to be ranked as species, and their hybrids, are very generally, but not universally,…

On the Imperfection of the Geological Record

lxiv
On the Absence of Intermediate Varieties at the Present Day.
In the sixth chapter I enumerated the chief objections which might be justly urged against the views maintained in this volume. Most of…
lxv
On the Lapse of Time, as inferred from the rate of Deposition and extent of Denudatym.
Independently of our not finding fossil remains of such infinitely numerous connecting links, it may be objected that time cannot have…
lxvi
On the Poorness of our Palæontological Collections.
Now let us turn to our richest museums, and what a paltry display we behold! That our collections are imperfect is admitted by every one.…
lxvii
On the Absence of Numerous Intermediate Varieties in any Single Formation.
From these several considerations it cannot be doubted that the geological record, viewed as a whole, is extremely imperfect; but if we…
lxviii
On the sudden Appearance of whole Groups of allied Species.
The abrupt manner in which whole groups of species suddenly appear in certain formations, has been urged by several palæontologists — for…
lxix
On the sudden Appearance of Groups of allied Species in the lowest known Fossiliferous Strata.
There is another and allied difficulty, which is much more serious. I allude to the manner in which species belonging to several of the…

On the Geological Succession of Organic Beings

lxx
On the Slow and Successive Appearance of New Species
Let us now see whether the several facts and laws relating to the geological succession of organic beings accord best with the common view…
lxxi
On Extinction.
We have as yet only spoken incidentally of the disappearance of species and of groups of species. On the theory of natural selection, the…
lxxii
On the Farms of Life changing almost simultaneously throughout the World.
Scarcely any palæontological discovery is more striking than the fact that the forms of life change almost simultaneously throughout the…
lxxiii
On the Affinities of Extinct Species to each other, and to Living Forms.
Let us now look to the mutual affinities of extinct and living species. All fall into a few grand classes; and this fact is at once…
lxxiv
On the State of Development of Ancient compared with Living Forms.
We have seen in the fourth chapter that the degree of differentiation and specialisation of the parts in organic beings, when arrived at…
lxxv
On the Succession of the same Types within the same Areas, during the later Tertiary periods.
Mr. Clift many years ago showed that the fossil mammals from the Australian caves were closely allied to the living marsupials of that…
lxxvi
Summary of the preceding and present Chapters.
I have attempted to show that the geological record is extremely imperfect; that only a small portion of the globe has been geologically…

Geographical Distribution

lxxvii
Present Distribution Cannot Be Accounted for by Differences in Physical Conditions
In considering the distribution of organic beings over the face of the globe, the first great fact which strikes us is, that neither the…
lxxviii
Means of Dispersal.
Sir C. Lyell and other authors have ably treated this subject. I can give here only the briefest abstract of the more important facts.…
lxxix
Dispersal during the Glacial Period.
The identity of many plants and animals, on mountain-summits, separated from each other by hundreds of miles of lowlands, where Alpine…
lxxx
Alternate Glacial Periods in the North and South.
But we must return to our more immediate subject. I am convinced that Forbes's view may be largely extended. In Europe we meet with the…

Geographical Distribution—Continued

lxxxi
Fresh-water Productions
As lakes and river-systems are separated from each other by barriers of land, it might have been thought that fresh-water productions would…
lxxxii
On the Inhabitants of Oceanic Islands.
We now come to the last of the three classes of facts, which I have selected as presenting the greatest amount of difficulty with respect…
lxxxiii
Absence of Batrachians and Terrestrial Mammals on Oceanic Islands.
With respect to the absence of whole orders of animals on oceanic islands, Bory St. Vincent long ago remarked that Batrachians (frogs,…
lxxxiv
On the Relation of the Inhabitants of Islands to those of the nearest Mainland.
The most striking and important fact for us is the affinity of the species which inhabit islands to those of the nearest mainland, without…
lxxxv
Summary of the last and present Chapters.
In these chapters I have endeavoured to show that if we make due allowance for our ignorance of the full effects of changes of climate and…

Mutual Affinities of Organic Beings: Morphology—Embryology—Rudimentary Organs

lxxxvi
Classification.
From the most remote period in the history of the world organic beings have been found to resemble each other in descending degrees, so…
lxxxvii
Analogical Resemblances.
We can understand why a species or a group of species may depart from its allies, in several of its most important characteristics, and yet…
lxxxviii
On the Nature of the Affinities Connecting Organic Beings.
As the modified descendants of dominant species, belonging to the larger genera, tend to inherit the advantages which made the groups to…
lxxxix
Morphology.
We have seen that the members of the same class, independently of their habits of life, resemble each other in the general plan of their…
xc
Development and Embryology.
This is one of the most important subjects in the whole round of natural history. The metamorphoses of insects, with which every one is…
xci
Rudimentary, Atrophied, and Aborted Organs.
Organs or parts in this strange condition, bearing the plain stamp of inutility, are extremely common, or even general, throughout nature.…
xcii
Summary.
In this chapter I have attempted to show that the arrangement of all organic beings throughout all time in groups under groups — that the…
xciii
Recapitulation and Conclusion
As this whole volume is one long argument, it may be convenient to the reader to have the leading facts and inferences briefly…