Chapter I: Preface: To First Edition

The present work treats the problem of the Origin of Species on the same general lines as were adopted by Darwin; but from the standpoint reached after nearly thirty years of discussion, with an abundance of new facts and the advocacy of many new or old theories.

While not attempting to deal, even in outline, with the vast subject of evolution in general, an endeavour has been made to give such an account of the theory of Natural Selection as may enable any intelligent reader to obtain a clear conception of Darwin's work, and to understand something of the power and range of his great principle.

Darwin wrote for a generation which had not accepted evolution, and which poured contempt on those who upheld the derivation of species from species by any natural law of descent. He did his work so well that "descent with modification" is now universally accepted as the order of nature in the organic world; and the rising generation of naturalists can hardly realise the novelty of this idea, or that their fathers considered it a scientific heresy to be condemned rather than seriously discussed.

The objections now made to Darwin's theory apply, solely, to the particular means by which the change of species has been brought about, not to the fact of that change. The objectors seek to minimise the agency of natural selection and to subordinate it to laws of variation, of use and disuse, of intelligence, and of heredity. These views and objections are urged with much force and more confidence, and for the most part by the modern school of laboratory naturalists, to whom the peculiarities and distinctions of species, as such, their distribution and their affinities, have little interest as compared with the problems of histology and embryology, of physiology and morphology. Their work in these departments is of the greatest interest and of the highest importance, but it is not the kind of work which, by itself, enables one to form a sound judgment on the questions involved in the action of the law of natural selection. These rest mainly on the external and vital relations of species to species in a state of nature--on what has been well termed by Semper the "physiology of organisms," rather than on the anatomy or physiology of organs.

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It has always been considered a weakness in Darwin's work that he based his theory, primarily, on the evidence of variation in domesticated animals and cultivated plants. I have endeavoured to secure a firm foundation for the theory in the variations of organisms in a state of nature; and as the exact amount and precise character of these variations is of paramount importance in the numerous problems that arise when we apply the theory to explain the facts of nature, I have endeavoured, by means of a series of diagrams, to exhibit to the eye the actual variations as they are found to exist in a sufficient number of species. By doing this, not only does the reader obtain a better and more precise idea of variation than can be given by any number of tabular statements or cases of extreme individual variation, but we obtain a basis of fact by which to test the statements and objections usually put forth on the subject of specific variability; and it will be found that, throughout the work, I have frequently to appeal to these diagrams and the facts they illustrate, just as Darwin was accustomed to appeal to the facts of variation among dogs and pigeons.

I have also made what appears to me an important change in the arrangement of the subject. Instead of treating first the comparatively difficult and unfamiliar details of variation, I commence with the Struggle for Existence, which is really the fundamental phenomenon on which natural selection depends, while the particular facts which illustrate it are comparatively familiar and very interesting. It has the further advantage that, after discussing variation and the effects of artificial selection, we proceed at once to explain how natural selection acts.

Among the subjects of novelty or interest discussed in this volume, and which have important bearings on the theory of natural selection, are: (1) A proof that all _specific_ characters are (or once have been) either useful in themselves or correlated with useful characters (Chap. VI); (2) a proof that natural selection can, in certain cases, increase the sterility of crosses (Chap. VII); (3) a fuller discussion of the colour relations of animals, with additional facts and arguments on the origin of sexual differences of colour (Chaps. VIII-X); (4) an attempted solution of the difficulty presented by the occurrence of both very simple and very complex modes of securing the cross-fertilisation of plants (Chap. XI); (5) some fresh facts and arguments on the wind-carriage of seeds, and its bearing on the wide dispersal of many arctic and alpine plants (Chap. XII); (6) some new illustrations of the non-heredity of acquired characters, and a proof that the effects of use and disuse, even if inherited, must be overpowered by natural selection (Chap. XIV); and (7) a new argument as to the nature and origin of the moral and intellectual faculties of man (Chap. XV).

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Although I maintain, and even enforce, my differences from some of Darwin's views, my whole work tends forcibly to illustrate the overwhelming importance of Natural Selection over all other agencies in the production of new species. I thus take up Darwin's earlier position, from which he somewhat receded in the later editions of his works, on account of criticisms and objections which I have endeavoured to show are unsound. Even in rejecting that phase of sexual selection depending on female choice, I insist on the greater efficacy of natural selection. This is pre-eminently the Darwinian doctrine, and I therefore claim for my book the position of being the advocate of pure Darwinism.

I wish to express my obligation to Mr. Francis Darwin for lending me some of his father's unused notes, and to many other friends for facts or information, which have, I believe, been acknowledged either in the text or footnotes. Mr. James Sime has kindly read over the proofs and given me many useful suggestions; and I have to thank Professor Meldola, Mr. Hemsley, and Mr. E.B. Poulton for valuable notes or corrections in the later chapters in which their special subjects are touched upon.

GODALMING, _March 1889_.

CONTENTS

CHAPTER I

WHAT ARE "SPECIES" AND WHAT IS MEANT BY THEIR "ORIGIN"

Definition of species--Special creation--The early
transmutationists--Scientific opinion before Darwin--The problem
before Darwin--The change of opinion effected by Darwin--The
Darwinian theory--Proposed mode of treatment of the subject

CHAPTER II

THE STRUGGLE FOR EXISTENCE

Its importance--The struggle among plants--Among
animals--Illustrative cases--Succession of trees in forests of
Denmark--The struggle for existence on the Pampas--Increase of
organisms in a geometrical ratio--Examples of rapid increase of
animals--Rapid increase and wide spread of plants--Great
fertility not essential to rapid increase--Struggle between
closely allied species most severe--The ethical aspect of the
struggle for existence

CHAPTER III

THE VARIABILITY OF SPECIES IN A STATE OF NATURE

Importance of variability--Popular ideas regarding
it--Variability of the lower animals--The variability of
insects--Variation among lizards--Variation among
birds--Diagrams of bird-variation--Number of varying
individuals--Variation in the mammalia--Variation in internal
organs--Variations in the skull--Variations in the habits of
animals--The variability of plants--Species which vary
little--Concluding remarks

CHAPTER IV

VARIATION OF DOMESTICATED ANIMALS AND CULTIVATED PLANTS

The facts of variation and artificial selection--Proofs of the
generality of variation--Variations of apples and
melons--Variations of flowers--Variations of domestic
animals--Domestic pigeons--Acclimatisation--Circumstances
favourable to selection by man--Conditions favourable to
variation--Concluding remarks

CHAPTER V

NATURAL SELECTION BY VARIATION AND SURVIVAL OF THE FITTEST

Effect of struggle for existence under unchanged conditions--The
effect under change of conditions--Divergence of character--In
insects--In birds--In mammalia--Divergence leads to a maximum of
life in each area--Closely allied species inhabit distinct
areas--Adaptation to conditions at various periods of life--The
continued existence of low forms of life--Extinction of low
types among the higher animals--Circumstances favourable to the
origin of new species--Probable origin of the dippers--The
importance of isolation--On the advance of organisation by
natural selection--Summary of the first five chapters

CHAPTER VI

DIFFICULTIES AND OBJECTIONS

Difficulty as to smallness of variations--As to the right
variations occurring when required--The beginnings of important
organs--The mammary glands--The eyes of flatfish--Origin of the
eye--Useless or non-adaptive characters--Recent extension of the
region of utility in plants--The same in animals--Uses of
tails--Of the horns of deer--Of the scale-ornamentation of
reptiles--Instability of non-adaptive characters--Delboeuf's
law--No "specific" character proved to be useless--The swamping
effects of intercrossing--Isolation as preventing
intercrossing--Gulick on the effects of isolation--Cases in
which isolation is ineffective

CHAPTER VII

ON THE INFERTILITY OF CROSSES BETWEEN DISTINCT SPECIES AND THE USUAL STERILITY OF THEIR HYBRID OFFSPRING

Statement of the problem--Extreme susceptibility of the
reproductive functions--Reciprocal crosses--Individual
differences in respect to cross-fertilisation--Dimorphism and
trimorphism among plants--Cases of the fertility of hybrids and
of the infertility of mongrels--The effects of close
interbreeding--Mr. Huth's objections--Fertile hybrids among
animals--Fertility of hybrids among plants--Cases of sterility
of mongrels--Parallelism between crossing and change of
conditions--Remarks on the facts of hybridity--Sterility due to
changed conditions and usually correlated with other
characters--Correlation of colour with constitutional
peculiarities--The isolation of varieties by selective
association--The influence of natural selection upon sterility
and fertility--Physiological selection--Summary and concluding
remarks

CHAPTER VIII

THE ORIGIN AND USES OF COLOUR IN ANIMALS

The Darwinian theory threw new light on organic colour--The
problem to be solved--The constancy of animal colour indicates
utility--Colour and environment--Arctic animals
white--Exceptions prove the rule--Desert, forest, nocturnal, and
oceanic animals--General theories of animal colour--Variable
protective colouring--Mr. Poulton's experiments--Special or
local colour adaptations--Imitation of particular objects--How
they have been produced--Special protective colouring of
butterflies--Protective resemblance among marine
animals--Protection by terrifying enemies--Alluring
coloration--The coloration of birds' eggs--Colour as a means of
recognition--Summary of the preceding exposition--Influence of
locality or of climate on colour--Concluding remarks

CHAPTER IX

WARNING COLORATION AND MIMICRY

The skunk as an example of warning coloration--Warning colours
among insects--Butterflies--Caterpillars--Mimicry--How mimicry
has been produced--Heliconidae--Perfection of the
imitation--Other cases of mimicry among Lepidoptera--Mimicry
among protected groups--Its explanation--Extension of the
principle--Mimicry in other orders of insects--Mimicry among the
vertebrata--Snakes--The rattlesnake and the cobra--Mimicry among
birds--Objections to the theory of mimicry--Concluding remarks
on warning colours and mimicry

CHAPTER X

COLOURS AND ORNAMENTS CHARACTERISTIC OF SEX

Sex colours in the mollusca and crustacea--In insects--In
butterflies and moths--Probable causes of these colours--Sexual
selection as a supposed cause--Sexual coloration of birds--Cause
of dull colours of female birds--Relation of sex colour to
nesting habits--Sexual colours of other vertebrates--Sexual
selection by the struggles of males--Sexual characters due to
natural selection--Decorative plumage of males and its effect on
the females--Display of decorative plumage by the males--A
theory of animal coloration--The origin of accessory
plumes--Development of accessory plumes and their display--The
effect of female preference will be neutralised by natural
selection--General laws of animal coloration--Concluding remarks

CHAPTER XI

THE SPECIAL COLOURS OF PLANTS: THEIR ORIGIN AND PURPOSE

The general colour relations of plants--Colours of fruits--The
meaning of nuts--Edible or attractive fruits--The colours of
flowers--Modes of securing cross-fertilisation--The
interpretation of the facts--Summary of additional facts
bearing on insect fertilisation--Fertilisation of flowers by
birds--Self-fertilisation of flowers--Difficulties and
contradictions--Intercrossing not necessarily
advantageous--Supposed evil results of close interbreeding--How
the struggle for existence acts among flowers--Flowers the
product of insect agency--Concluding remarks on colour in nature

CHAPTER XII

THE GEOGRAPHICAL DISTRIBUTION OF ORGANISMS

The facts to be explained--The conditions which have determined
distribution--The permanence of oceans--Oceanic and continental
areas--Madagascar and New Zealand--The teachings of the
thousand-fathom line--The distribution of marsupials--The
distribution of tapirs--Powers of dispersal as illustrated by
insular organisms--Birds and insects at sea--Insects at great
altitudes--The dispersal of plants--Dispersal of seeds by the
wind--Mineral matter carried by the wind--Objections to the
theory of wind-dispersal answered--Explanation of north
temperate plants in the southern hemisphere--No proof of
glaciation in the tropics--Lower temperature not needed to
explain the facts--Concluding remarks

CHAPTER XIII

THE GEOLOGICAL EVIDENCES OF EVOLUTION

What we may expect--The number of known species of extinct
animals--Causes of the imperfection of the geological
record--Geological evidences of
evolution--Shells--Crocodiles--The rhinoceros tribe--The
pedigree of the horse tribe--Development of deer's horns--Brain
development--Local relations of fossil and living animals--Cause
of extinction of large animals--Indications of general progress
in plants and animals--The progressive development of
plants--Possible cause of sudden late appearance of
exogens--Geological distribution of insects--Geological
succession of vertebrata--Concluding remarks

CHAPTER XIV

FUNDAMENTAL PROBLEMS IN RELATION TO VARIATION AND HEREDITY

Fundamental difficulties and objections--Mr. Herbert Spencer's
factors of organic evolution--Disuse and effects of withdrawal
of natural selection--Supposed effects of disuse among wild
animals--Difficulty as to co-adaptation of parts by variation
and selection--Direct action of the environment--The American
school of evolutionists--Origin of the feet of the
ungulates--Supposed action of animal intelligence--Semper on the
direct influence of the environment--Professor Geddes's theory
of variation in plants--Objections to the theory--On the origin
of spines--Variation and selection overpower the effects of use
and disuse--Supposed action of the environment in imitating
variations--Weismann's theory of heredity--The cause of
variation--The non-heredity of acquired characters--The theory
of instinct--Concluding remarks