On the Origin of Species

CHAPTER I.. Variation Under Domestication.

VARIATION UNDER DOMESTICATION.

Causes of Variability--Effects of Habit--Correlation of Growth--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--Principle of Selection anciently followed, its Effects--Methodical and Unconscious Selection--Unknown Origin of our Domestic Productions--Circumstances favourable to Man's power of Selection

7-43

CHAPTER II.. Variation Under Nature.

VARIATION UNDER NATURE.

Variability--Individual differences--Doubtful species--Wide ranging, much diffused, and common species vary most--Species of the larger genera in any country vary more 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

44-59

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CHAPTER III.. Struggle for Existence.

STRUGGLE FOR EXISTENCE.

Its bearing on natural selection--The term used in a wide sense--Geometrical powers 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

60-79

CHAPTER IV.. Natural Selection.

NATURAL SELECTION.

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 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

80-130

CHAPTER V.. Laws of Variation.

LAWS OF VARIATION.

Effects of external conditions--Use and disuse, combined with natural selection; organs of flight and of vision--Acclimatisation--Correlation of growth--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

131-170

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CHAPTER VI.. Difficulties on Theory.

DIFFICULTIES ON THEORY.

Difficulties on the theory of descent with modification--Transitions--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--Means 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

171-206

CHAPTER VII.. Instinct.

INSTINCT.

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, ostrich, and parasitic bees--Slave-making ants--Hive-bee, its cell-making instinct--Difficulties on the theory of the Natural Selection of instincts--Neuter or sterile insects--Summary

207-244

CHAPTER VIII.. Hybridism.

HYBRIDISM.

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--Causes of the sterility of first crosses and of hybrids--Parallelism between the effects of changed conditions of life and crossing--Fertility of varieties when crossed and of their mongrel offspring not universal--Hybrids and mongrels compared independently of their fertility--Summary

245-278

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CHAPTER IX.. On the Imperfection of the Geological Record.

ON THE IMPERFECTION OF THE GEOLOGICAL RECORD.

On the absence of intermediate varieties at the present day--On the nature of extinct intermediate varieties; on their number--On the vast lapse of time, as inferred from the rate of deposition and of denudation--On the poorness of our palæontological collections--On the intermittence of geological formations--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

279-311

CHAPTER X.. On the Geological Succession of Organic Beings.

ON THE GEOLOGICAL SUCCESSION OF ORGANIC BEINGS.

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 chapters

312-345

CHAPTER XI.. Geographical Distribution.

GEOGRAPHICAL DISTRIBUTION.

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 co-extensive with the world

346-382

CHAPTER XII.. Geographical Distribution--_continued_.

GEOGRAPHICAL DISTRIBUTION--continued.

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 chapters

383-410

CHAPTER XIII.

MUTUAL AFFINITIES OF ORGANIC BEINGS: MORPHOLOGY: EMBRYOLOGY: RUDIMENTARY ORGANS.

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

411-458

CHAPTER XIV.. Recapitulation and Conclusion.

RECAPITULATION AND CONCLUSION.

Recapitulation of the difficulties on 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

459-490

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ON THE ORIGIN OF SPECIES.

* * * * *

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, and in the geological relations of the present to the past inhabitants of that continent. These facts seemed to me to throw some light on the origin of species--that mystery of mysteries, as it has been called by one of our greatest philosophers. On my return home, it occurred to me, in 1837, that something might perhaps be made out on this question by patiently accumulating and reflecting on all sorts of facts which could possibly have any bearing on it. After five years' work I allowed myself to speculate on the subject, and drew up 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 steadily pursued the same object. I hope that I may be excused for entering on these personal 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 publish this Abstract. I have more especially been induced to do this, as Mr. Wallace, who is now studying the {2} natural history of the Malay archipelago, has arrived at almost exactly the same general conclusions that I have on the origin of species. Last year he sent me a memoir on this subject, with a request that I would forward it to Sir 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 authorities for my several statements; and I must trust to the reader reposing some confidence in my accuracy. No doubt errors will have crept in, though I hope I have always been cautious in trusting to good authorities alone. I can here give only the general conclusions at which I have arrived, with a few facts in illustration, but which, I 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 conclusions have been grounded; and I hope in a future work to do this. For I am well aware that scarcely a single point is discussed in this volume on which facts cannot be adduced, often apparently leading to conclusions directly 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, {3} let this opportunity pass without expressing my deep obligations to Dr. Hooker, who for the last fifteen years has aided me in every possible way by his large stores of knowledge and his excellent judgment.

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