Extracts from Charles Darwin

On the Origin of Species by Means of Natural Selection 1859


Causes of Variability--Effects of Habit and the use or disuse of Parts--Correlated Variation--Inheritance--Character of Domestic Varieties--Difficulty of distinguishing between Varieties and Species--Origin of Domestic Varieties from one or more Species--Domestic Pigeons, their Differences and Origin--Principles of Selection, anciently followed, their Effects--Methodical and Unconscious Selection--Unknown Origin of our Domestic Productions--Circumstances favourable to Man's power of Selection.
Variability--Individual Differences--Doubtful species--Wide ranging, much diffused, and common species, vary most--Species of the larger genera in each country vary more frequently than the species of the smaller genera--Many of the species of the larger genera resemble varieties in being very closely, but unequally, related to each other, and in having restricted ranges.
Its bearing on natural selection--The term used in a wide sense--Geometrical ratio of increase--Rapid increase of naturalised animals and plants--Nature of the checks to increase--Competition universal--Effects of climate--Protection from the number of individuals--Complex relations of all animals and plants throughout nature--Struggle for life most severe between individuals and varieties of the same species; often severe between species of the same genus--The relation of organism to organism the most important of all relations.
Natural Selection--its power compared with man's selection--its power on characters of trifling importance--its power at all ages and on both sexes--Sexual Selection--On the generality of intercrosses between individuals of the same species--Circumstances favourable and unfavourable to the results of Natural Selection, namely, intercrossing, isolation, number of individuals--Slow action--Extinction caused by Natural Selection--Divergence of Character, related to the diversity of inhabitants of any small area and to naturalisation--Action of Natural Selection, through Divergence of Character and Extinction, on the descendants from a common parent--Explains the Grouping of all organic beings--Advance in organisation--Low forms preserved--Convergence of character--Indefinite multiplication of species--Summary.
Effects of changed conditions--Use and disuse, combined with natural selection; organs of flight and of vision--Acclimatisation--Correlated variation--Compensation and economy of growth--False correlations--Multiple, rudimentary, and lowly organised structures variable--Parts developed in an unusual manner are highly variable; specific characters more variable than generic; secondary sexual characters variable--Species of the same genus vary in an analogous manner--Reversions to long-lost characters--Summary.
Difficulties of the theory of descent with modification--Absence or rarity of transitional varieties--Transitions in habits of life--Diversified habits in the same species--Species with habits widely different from those of their allies--Organs of extreme perfection--Modes of transition--Cases of difficulty--Natura non facit saltum--Organs of small importance--Organs not in all cases absolutely perfect--The law of Unity of Type and of the Conditions of Existence embraced by the theory of Natural Selection.
Longevity--Modifications not necessarily simultaneous--Modifications apparently of no direct service--Progressive development--Characters of small functional importance, the most constant--Supposed incompetence of natural selection to account for the incipient stages of useful structures--Causes which interfere with the acquisition through natural selection of useful structures--Gradations of structure with changed functions--Widely different organs in members of the same class, developed from one and the same source--Reasons for disbelieving in great and abrupt modifications.
Instincts comparable with habits, but different in their origin--Instincts graduated--Aphides and ants--Instincts variable--Domestic instincts, their origin--Natural instincts of the cuckoo, molothrus, ostrich, and parasitic bees--Slave-making ants--Hive-bee, its cell-making instinct--Changes of instinct and structure not necessarily simultaneous--Difficulties on the theory of the Natural Selection of instincts--Neuter or sterile insects--Summary.
Distinction between the sterility of first crosses and of hybrids--Sterility various in degree, not universal, affected by close interbreeding, removed by domestication--Laws governing the sterility of hybrids--Sterility not a special endowment, but incidental on other differences, not accumulated by natural selection--Causes of the sterility of first crosses and of hybrids--Parallelism between the effects of changed conditions of life and of crossing--Dimorphism and Trimorphism--Fertility of varieties when crossed and of their mongrel offspring not universal--Hybrids and mongrels compared independently of their fertility--Summary.
On the absence of intermediate varieties at the present day--On the nature of extinct intermediate varieties; on their number--On the lapse of time, as inferred from the rate of denudation and of deposition--On the lapse of time as estimated in years--On the poorness of our palaeontological collections--On the intermittence of geological formations--On the denudation of granitic areas--On the absence of intermediate varieties in any one formation--On the sudden appearance of groups of species--On their sudden appearance in the lowest known fossiliferous strata--Antiquity of the habitable earth.
On the slow and successive appearance of new species--On their different rates of change--Species once lost do not reappear--Groups of species follow the same general rules in their appearance and disappearance as do single species--On extinction--On simultaneous changes in the forms of life throughout the world--On the affinities of extinct species to each other and to living species--On the state of development of ancient forms--On the succession of the same types within the same areas--Summary of preceding and present chapter.
Present distribution cannot be accounted for by differences in physical conditions--Importance of barriers--Affinity of the productions of the same continent--Centres of creation--Means of dispersal by changes of climate and of the level of the land, and by occasional means--Dispersal during the Glacial period--Alternate Glacial periods in the north and south.
Distribution of fresh-water productions--On the inhabitants of oceanic islands--Absence of Batrachians and of terrestrial Mammals--On the relation of the inhabitants of islands to those of the nearest mainland--On colonisation from the nearest source with subsequent modification--Summary of the last and present chapter.
Classification, groups subordinate to groups--Natural system--Rules and difficulties in classification, explained on the theory of descent with modification--Classification of varieties--Descent always used in classification--Analogical or adaptive characters--Affinities, general, complex and radiating--Extinction separates and defines groups--Morphology, between members of the same class, between parts of the same individual--Embryology, laws of, explained by variations not supervening at an early age, and being inherited at a corresponding age--Rudimentary Organs; their origin explained--Summary.
Recapitulation of the objections to the theory of Natural Selection--Recapitulation of the general and special circumstances in its favour--Causes of the general belief in the immutability of species--How far the theory of Natural Selection may be extended--Effects of its adoption on the study of Natural history--Concluding remarks.


In considering the Origin of Species, it is quite conceivable that a naturalist, reflecting on the mutual affinities 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 descended, like varieties, from other species. Nevertheless, such a conclusion, even if well founded, would be unsatisfactory, 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, as 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, so 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 absolutely requiring the agency of certain insects to bring pollen from one flower to the other, it is equally preposterous to account for the structure of this parasite, with its relations to several distinct organic beings, by the effects of external conditions, or of habit, or of the volition of the plant itself.

The author of the Vestiges of Creation would, I presume, say that, after a certain unknown number of generations, some bird had given birth to a woodpecker, and some plant to the misseltoe, and that these had been produced perfect as we now see them; but this assumption seems to me to be no explanation, for it leaves the case of the coadaptations of organic beings to each other and to their physical conditions of life, untouched and unexplained.

It is, therefore, of the highest importance to gain a clear insight into the means of modification and coadaptation. At the commencement of my observations it seemed to me probable that a careful study of domesticated animals 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 Domestication. We shall thus see that a large amount of hereditary modification is at least possible, and, what is equally or more important, we shall see how great is the power of man in accumulating by his Selection successive slight variations. I will then pass on to the variability of species in a state of nature; but I shall, unfortunately, be compelled to treat this subject far too briefly, as it can be treated properly only by giving long catalogues of facts. We shall, however, be enabled to discuss what circumstances are most favourable to variation. In the next chapter the Struggle for Existence amongst all organic beings throughout the world, which inevitably follows from their high geometrical powers of increase, will be treated of. This is the doctrine of Malthus, applied to the whole animal and vegetable kingdoms. As many more individuals of each species are born than can possibly survive; and as, consequently, there is a frequently recurring struggle for existence, it follows that any being, if it vary however slightly in any manner profitable to itself, under the complex and sometimes varying conditions of life, will have a better chance of surviving, and thus be naturally selected. From the strong principle of inheritance, any selected variety will tend to propagate its new and modified form.


Embryology and Development.

This is one of the most important departments of natural history. Herein are included the ordinary metamorphoses of insects, with which every one is familiar. These are generally effected somewhat abruptly by a few stages and in a concealed manner; but the transformations are in reality numerous and graduated. For instance, Sir J. Lubbock has recently shown that a certain ephemerous insect (Chlöeon) during its development moults above twenty times, and each time undergoes a certain amount of change; in such cases we probably behold the act of metamorphosis in its natural or primary progress.

What great changes of structure are effected during the development of some animals is seen in the case of insects, but still more plainly with many crustaceans. When, however, we read of the several wonderful cases, recently discovered, of the so-called alternate generations of animals, we come to the climax of developmental transformation. What fact can be more astonishing than that a delicate branching coralline, studded with polypi and attached to a submarine rock, should produce, first by budding and then by transverse division, a host of huge floating jelly-fishes; and that these should produce eggs, from which are hatched swimming animalcules, which attach themselves to rocks and become developed into branching corallines; and so on in an endless cycle?

Hence it will be seen that I follow those naturalists who look at all cases of alternate generation, as essentially modifications of the process of budding, which may supervene at any stage of development. This view of the close connection between alternate generations and ordinary metamorphoses has recently been much strengthened by Wagner's discovery of the larva of a Cecidomyia,-that is of the maggot of a fly, -producing asexually within its body other and similar larv‘; these again repeating the process

Extracts from Charles Darwin

Descent of Man, and Selection in relation to Sex 1871

Part Three. Sexual selection in relation to man, and conclusion

Chapter 19.

Secondary Sexual Characters of Man.

Differences between man and woman--Causes of such differences, and of certain characters common to both sexes--Law of battle--Differences in mental powers, and voice--On the influence of beauty in determining the marriages of mankind--Attention paid by savages to ornaments--Their ideas of beauty in women--The tendency to exaggerate each natural peculiarity.

Chapter 20.

Secondary Sexual Characters of Man--continued.

On the effects of the continued selection of women according to a different standard of beauty in each race - On the causes which interfere with sexual selection in civilised and savage nations - Conditions favourable to sexual selection during primeval times--On the manner of action of sexual selection with mankind--On the women in savage tribes having some power to choose their husbands--Absence of hair on the body, and development of the beard--Colour of the skin--Summary.

Chapter 21.

General Summary and Conclusion.

Main conclusion that man is descended from some lower form--Manner of development--Genealogy of man--Intellectual and moral faculties--Sexual selection--Concluding remarks.

On the causes which interfere with sexual selection in civilised and savage nations

The chief causes are, first, so-called communal marriages or promiscuous intercourse; secondly, the consequences of female infanticide; thirdly, early betrothals; and lastly, the low estimation in which women are held, as mere slaves.

Early betrothals and slavery of women

With many savages it is the custom to betroth the females whilst mere infants; and this would effectually prevent preference being exerted on either side according to personal appearance. But it would not prevent the more attractive women from being afterwards stolen or taken by force from their husbands by the more powerful men; and this often happens in Australia, America, and elsewhere. The same consequences with reference to sexual selection would to a certain extent follow, when women are valued almost solely as slaves or beasts of burden, as is the case with many savages. The men, however, at all times would prefer the handsomest slaves according to their standard of beauty.

We thus see that several customs prevail with savages which must greatly interfere with, or completely stop, the action of sexual selection. On the other hand, the conditions of life to which savages are exposed, and some of their habits, are favourable to natural selection; and this comes into play at the same time with sexual selection. Savages are known to suffer severely from recurrent famines; they do not increase their food by artificial means; they rarely refrain from marriage (16. Burchell says ('Travels in S. Africa,' vol. ii. 1824, p. 58), that among the wild nations of Southern Africa, neither men nor women ever pass their lives in a state of celibacy. Azara ('Voyages dans l'Amerique Merid.' tom. ii. 1809, p. 21) makes precisely the same remark in regard to the wild Indians of South America.), and generally marry whilst young. Consequently they must be subjected to occasional hard struggles for existence, and the favoured individuals will alone survive.

At a very early period, before man attained to his present rank in the scale, many of his conditions would be different from what now obtains amongst savages. Judging from the analogy of the lower animals, he would then either live with a single female, or be a polygamist. The most powerful and able males would succeed best in obtaining attractive females. They would also succeed best in the general struggle for life, and in defending their females, as well as their offspring, from enemies of all kinds. At this early period the ancestors of man would not be sufficiently advanced in intellect to look forward to distant contingencies; they would not foresee that the rearing of all their children, especially their female children, would make the struggle for life severer for the tribe. They would be governed more by their instincts and less by their reason than are savages at the present day. They would not at that period have partially lost one of the strongest of all instincts, common to all the lower animals, namely the love of their young offspring; and consequently they would not have practised female infanticide. Women would not have been thus rendered scarce, and polyandry would not have been practised; for hardly any other cause, except the scarcity of women seems sufficient to break down the natural and widely prevalent feeling of jealousy, and the desire of each male to possess a female for himself. Polyandry would be a natural stepping-stone to communal marriages or almost promiscuous intercourse; though the best authorities believe that this latter habit preceded polyandry. During primordial times there would be no early betrothals, for this implies foresight. Nor would women be valued merely as useful slaves or beasts of burthen. Both sexes, if the females as well as the males were permitted to exert any choice, would choose their partners not for mental charms, or property, or social position, but almost solely from external appearance. All the adults would marry or pair, and all the offspring, as far as that was possible, would be reared; so that the struggle for existence would be periodically excessively severe. Thus during these times all the conditions for sexual selection would have been more favourable than at a later period, when man had advanced in his intellectual powers but had retrograded in his instincts. Therefore, whatever influence sexual selection may have had in producing the differences between the races of man, and between man and the higher Quadrumana, this influence would have been more powerful at a remote period than at the present day, though probably not yet wholly lost.


With primeval man under the favourable conditions just stated, and with those savages who at the present time enter into any marriage tie, sexual selection has probably acted in the following manner, subject to greater or less interference from female infanticide, early betrothals, etc. The strongest and most vigorous men--those who could best defend and hunt for their families, who were provided with the best weapons and possessed the most property, such as a large number of dogs or other animals,--would succeed in rearing a greater average number of offspring than the weaker and poorer members of the same tribes. There can, also, be no doubt that such men would generally be able to select the more attractive women. At present the chiefs of nearly every tribe throughout the world succeed in obtaining more than one wife. I hear from Mr. Mantell that, until recently, almost every girl in New Zealand who was pretty, or promised to be pretty, was tapu to some chief. With the Kafirs, as Mr. C. Hamilton states (17. 'Anthropological Review,' Jan. 1870, p. xvi.), "the chiefs generally have the pick of the women for many miles round, and are most persevering in establishing or confirming their privilege." We have seen that each race has its own style of beauty, and we know that it is natural to man to admire each characteristic point in his domestic animals, dress, ornaments, and personal appearance, when carried a little beyond the average. If then the several foregoing propositions be admitted, and I cannot see that they are doubtful, it would be an inexplicable circumstance if the selection of the more attractive women by the more powerful men of each tribe, who would rear on an average a greater number of children, did not after the lapse of many generations somewhat modify the character of the tribe.

When a foreign breed of our domestic animals is introduced into a new country, or when a native breed is long and carefully attended to, either for use or ornament, it is found after several generations to have undergone a greater or less amount of change whenever the means of comparison exist. This follows from unconscious selection during a long series of generations--that is, the preservation of the most approved individuals--without any wish or expectation of such a result on the part of the breeder. So again, if during many years two careful breeders rear animals of the same family, and do not compare them together or with a common standard, the animals are found to have become, to the surprise of their owners, slightly different. (18. The 'Variation of Animals and Plants under Domestication,' vol. ii. pp. 210-217.) Each breeder has impressed, as von Nathusius well expresses it, the character of his own mind--his own taste and judgment--on his animals. What reason, then, can be assigned why similar results should not follow from the long-continued selection of the most admired women by those men of each tribe who were able to rear the greatest number of children? This would be unconscious selection, for an effect would be produced, independently of any wish or expectation on the part of the men who preferred certain women to others.

Let us suppose the members of a tribe, practising some form of marriage, to spread over an unoccupied continent, they would soon split up into distinct hordes, separated from each other by various barriers, and still more effectually by the incessant wars between all barbarous nations. The hordes would thus be exposed to slightly different conditions and habits of life, and would sooner or later come to differ in some small degree. As soon as this occurred, each isolated tribe would form for itself a slightly different standard of beauty (19. An ingenious writer argues, from a comparison of the pictures of Raphael, Rubens, and modern French artists, that the idea of beauty is not absolutely the same even throughout Europe: see the 'Lives of Haydn and Mozart,' by Bombet (otherwise M. Beyle), English translation, p. 278.); and then unconscious selection would come into action through the more powerful and leading men preferring certain women to others. Thus the differences between the tribes, at first very slight, would gradually and inevitably be more or less increased.

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Freud: Totem and Taboo
Durkheim: Elementary Forms