THE FOUNDATIONS OF THE
ORIGIN OF SPECIES

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From a photograph by Maull & Fox in 1854

THE FOUNDATIONS OF THE
ORIGIN OF SPECIES
TWO ESSAYS
WRITTEN IN 1842 AND 1844
by
CHARLES DARWIN

Edited by his son
FRANCIS DARWIN
Honorary Fellow of Christ's College

Cambridge:
at the University Press
1909

Astronomers might formerly have said that God ordered each planet to move in its particular destiny. In same manner God orders each animal created with certain form in certain country. But how much more simple and sublime power,—let attraction act according to certain law, such are inevitable consequences,—let animal«s» be created, then by the fixed laws of generation, such will be their successors.

From Darwin’s Note Book, 1837, p. 101.

TO THE MASTER AND FELLOWS
OF CHRIST’S COLLEGE, THIS
BOOK IS DEDICATED BY THE
EDITOR IN TOKEN OF RESPECT
AND GRATITUDE

CONTENTS

ESSAY OF 1842

• PAGES
  
• [Introduction] [xi]

PART I

• § i. On variation under domestication, and on the principles of selection [1]
• § ii. On variation in a state of nature and on the natural means of selection [4]
• § iii. On variation in instincts and other mental attributes [17]

PART II

• §§ iv. and v. On the evidence from Geology. (The reasons for combining the two sections are given in the Introduction) [22]
• § vi. Geographical distribution [29]
• § vii. Affinities and classification [35]
• § viii. Unity of type in the great classes [38]
• § ix. Abortive organs [45]
• § x. Recapitulation and conclusion [48]

ESSAY OF 1844

PART I

CHAPTER I
ON THE VARIATION OF ORGANIC BEINGS UNDER DOMESTICATION; AND ON THE PRINCIPLES OF SELECTION.

• Variation
• On the hereditary tendency
• Causes of Variation
• On Selection
• Crossing Breeds
• Whether our domestic races have descended from one or more wild stocks
• Limits to Variation in degree and kind
• In what consists Domestication
• Summary [57]-[80]

CHAPTER II
ON THE VARIATION OF ORGANIC BEINGS IN A WILD STATE; ON THE NATURAL MEANS OF SELECTION; AND ON THE COMPARISON OF DOMESTIC RACES AND TRUE SPECIES.

• Variation
• Natural means of Selection
• Differences between “Races” and “Species”:—first, in their trueness or variability
• Difference between “Races” and “Species” in fertility when crossed
• Causes of Sterility in Hybrids
• Infertility from causes distinct from hybridisation
• Points of Resemblance between “Races” and “Species”
• External characters of Hybrids and Mongrels
• Summary
• Limits of Variation [81]-[111]

CHAPTER III
ON THE VARIATION OF INSTINCTS AND OTHER MENTAL ATTRIBUTES UNDER DOMESTICATION AND IN A STATE OF NATURE; ON THE DIFFICULTIES IN THIS SUBJECT; AND ON ANALOGOUS DIFFICULTIES WITH RESPECT TO CORPOREAL STRUCTURES.

• Variation of mental attributes under domestication
• Hereditary habits compared with instincts
• Variation in the mental attributes of wild animals
• Principles of Selection applicable to instincts
• Difficulties in the acquirement of complex instincts by Selection
• Difficulties in the acquirement by Selection of complex corporeal structures [112]-[132]

PART II
ON THE EVIDENCE FAVOURABLE AND OPPOSED TO THE VIEW THAT SPECIES ARE NATURALLY FORMED RACES, DESCENDED FROM COMMON STOCKS.

CHAPTER IV
ON THE NUMBER OF INTERMEDIATE FORMS REQUIRED ON THE THEORY OF COMMON DESCENT; AND ON THEIR ABSENCE IN A FOSSIL STATE [133]-[143]

CHAPTER V
GRADUAL APPEARANCE AND DISAPPEARANCE OF SPECIES. [144]-[150]

• Gradual appearance of species
• Extinction of species

CHAPTER VI
ON THE GEOGRAPHICAL DISTRIBUTION OF ORGANIC BEINGS IN PAST AND PRESENT TIMES.

SECTION FIRST [151]-[174]

• Distribution of the inhabitants in the different continents
• Relation of range in genera and species
• Distribution of the inhabitants in the same continent
• Insular Faunas
• Alpine Floras
• Cause of the similarity in the floras of some distant mountains
• Whether the same species has been created more than once
• On the number of species, and of the classes to which they belong in different regions

SECOND SECTION [174]-[182]

• Geographical distribution of extinct organisms
• Changes in geographical distribution
• Summary on the distribution of living and extinct organic beings

SECTION THIRD [183]-[197]

• An attempt to explain the foregoing laws of geographical distribution, on the theory of allied species having a common descent
• Improbability of finding fossil forms intermediate between existing species

CHAPTER VII
ON THE NATURE OF THE AFFINITIES AND CLASSIFICATION
OF ORGANIC BEINGS. [198]-[213]

• Gradual appearance and disappearance of groups
• What is the Natural System?
• On the kind of relation between distinct groups
• Classification of Races or Varieties
• Classification of Races and Species similar
• Origin of genera and families

CHAPTER VIII
UNITY OF TYPE IN THE GREAT CLASSES; AND MORPHOLOGICAL STRUCTURES.

• Unity of Type
• Morphology
• Embryology
• Attempt to explain the facts of embryology
• On the graduated complexity in each great class
• Modification by selection of the forms of immature animals
• Importance of embryology in classification
• Order in time in which the great classes have first appeared [214]-[230]

CHAPTER IX
ABORTIVE OR RUDIMENTARY ORGANS.

• The abortive organs of Naturalists
• The abortive organs of Physiologists
• Abortion from gradual disuse [231]-[238]

CHAPTER X
RECAPITULATION AND CONCLUSION.

• Recapitulation
• Why do we wish to reject the Theory of Common Descent?
• Conclusion [239]-[255]
  
• Index [257]
  
• Portrait frontispiece
• Facsimile to face p. 50

INTRODUCTION

We know from the contents of Charles Darwin’s Note Book of 1837 that he was at that time a convinced Evolutionist[{1}]. Nor can there be any doubt that, when he started on board the Beagle, such opinions as he had were on the side of immutability. When therefore did the current of his thoughts begin to set in the direction of Evolution?

We have first to consider the factors that made for such a change. On his departure in 1831, Henslow gave him vol. I. of Lyell's Principles, then just published, with the warning that he was not to believe what he read[{2}]. But believe he did, and it is certain (as Huxley has forcibly pointed out[{3}]) that the doctrine of uniformitarianism when applied to Biology leads of necessity to Evolution. If the extermination of a species is no more catastrophic than the natural death of an individual, why should the birth of a species be any more miraculous than the birth of an individual? It is quite clear that this thought was vividly present to Darwin when he was writing out his early thoughts in the 1837 Note Book[{4}]:—

“Propagation explains why modern animals same type as extinct, which is law almost proved. They die, without they change, like golden pippins; it is a generation of species like generation of individuals.”

“If species generate other species their race is not utterly cut off.”

These quotations show that he was struggling to see in the origin of species a process just as scientifically comprehensible as the birth of individuals. They show, I think, that he recognised the two things not merely as similar but as identical.

It is impossible to know how soon the ferment of uniformitarianism began to work, but it is fair to suspect that in 1832 he had already begun to see that mutability was the logical conclusion of Lyell’s doctrine, though this was not acknowledged by Lyell himself.

There were however other factors of change. In his Autobiography [{5}] he wrote:—“During the voyage of the Beagle I had been deeply impressed by discovering in the Pampean formation great fossil animals covered with armour like that on the existing armadillos; secondly, by the manner in which closely allied animals replace one another in proceeding southward over the Continent; and thirdly, by the South American character of most of the productions of the Galapagos archipelago, and more especially by the manner in which they differ slightly on each island of the group; none of the islands appearing to be very ancient in a geological sense. It was evident that such facts as these, as well as many others, could only be explained on the supposition that species gradually become modified; and the subject haunted me.”

Again we have to ask: how soon did any of these influences produce an effect on Darwin’s mind? Different answers have been attempted. Huxley[{6}] held that these facts could not have produced their essential effect until the voyage had come to an end, and the “relations of the existing with the extinct species and of the species of the different geographical areas with one another were determined with some exactness.” He does not therefore allow that any appreciable advance towards evolution was made during the actual voyage of the Beagle.

Professor Judd[{7}] takes a very different view. He holds that November 1832 may be given with some confidence as the “date at which Darwin commenced that long series of observations and reasonings which eventually culminated in the preparation of the Origin of Species.”

Though I think these words suggest a more direct and continuous march than really existed between fossil-collecting in 1832 and writing the Origin of Species in 1859, yet I hold that it was during the voyage that Darwin's mind began to be turned in the direction of Evolution, and I am therefore in essential agreement with Prof. Judd, although I lay more stress than he does on the latter part of the voyage.

Let us for a moment confine our attention to the passage, above quoted, from the Autobiography and to what is said in the Introduction to the Origin, Ed. i., viz. “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 words, occurring where they do, can only mean one thing,—namely that the facts suggested an evolutionary interpretation. And this being so it must be true that his thoughts began to flow in the direction of Descent at this early date.

I am inclined to think that the “new light which was rising in his mind[{8}]” had not yet attained any effective degree of steadiness or brightness. I think so because in his Pocket Book under the date 1837 he wrote, “In July opened first note-book on ‘transmutation of species.’ Had been greatly struck from about month of previous March[{9}] on character of South American fossils, and species on Galapagos Archipelago. These facts origin (especially latter), of all my views.” But he did not visit the Galapagos till 1835 and I therefore find it hard to believe that his evolutionary views attained any strength or permanence until at any rate quite late in the voyage. The Galapagos facts are strongly against Huxley’s view, for Darwin’s attention was “thoroughly aroused[{10}]” by comparing the birds shot by himself and by others on board. The case must have struck him at once,—without waiting for accurate determinations,—as a microcosm of evolution.

It is also to be noted, in regard to the remains of extinct animals, that, in the above quotation from his Pocket Book, he speaks of March 1837 as the time at which he began to be “greatly struck on character of South American fossils,” which suggests at least that the impression made in 1832 required reinforcement before a really powerful effect was produced.

We may therefore conclude, I think, that the evolutionary current in my father's thoughts had continued to increase in force from 1832 onwards, being especially reinforced at the Galapagos in 1835 and again in 1837 when he was overhauling the results, mental and material, of his travels. And that when the above record in the Pocket Book was made he unconsciously minimised the earlier beginnings of his theorisings, and laid more stress on the recent thoughts which were naturally more vivid to him. In his letter[{11}] to Otto Zacharias (1877) he wrote, “On my return home in the autumn of 1836, I immediately began to prepare my Journal for publication, and then saw how many facts indicated the common descent of species.” This again is evidence in favour of the view that the later growths of his theory were the essentially important parts of its development.

In the same letter to Zacharias he says, “When I was on board the Beagle I believed in the permanence of species, but as far as I can remember vague doubts occasionally flitted across my mind.” Unless Prof. Judd and I are altogether wrong in believing that late or early in the voyage (it matters little which) a definite approach was made to the evolutionary standpoint, we must suppose that in 40 years such advance had shrunk in his recollection to the dimensions of “vague doubts.” The letter to Zacharias shows I think some forgetting of the past where the author says, “But I did not become convinced that species were mutable until, I think, two or three years had elapsed.” It is impossible to reconcile this with the contents of the evolutionary Note Book of 1837. I have no doubt that in his retrospect he felt that he had not been “convinced that species were mutable” until he had gained a clear conception of the mechanism of natural selection, i.e. in 1838-9.

But even on this last date there is some room, not for doubt, but for surprise. The passage in the Autobiography[{12}] is quite clear, namely that in October 1838 he read Malthus’s Essay on the principle of Population and “being well prepared to appreciate the struggle for existence ..., it at once struck me that under these circumstances favourable variations would tend to be preserved, and unfavourable ones to be destroyed. The result of this would be the formation of new species. Here then I had at last got a theory by which to work.”

It is surprising that Malthus should have been needed to give him the clue, when in the Note Book of 1837 there should occur—however obscurely expressed—the following forecast[{13}] of the importance of the survival of the fittest. “With respect to extinction, we can easily see that a variety of the ostrich (Petise [{14}]), may not be well adapted, and thus perish out; or on the other hand, like Orpheus[{15}], being favourable, many might be produced. This requires the principle that the permanent variations produced by confined breeding and changing circumstances are continued and produce«d» according to the adaptation of such circumstances, and therefore that death of species is a consequence (contrary to what would appear in America) of non-adaptation of circumstances.”

I can hardly doubt, that with his knowledge of the interdependence of organisms and the tyranny of conditions, his experience would have crystallized out into “a theory by which to work” even without the aid of Malthus.

In my father's Autobiography[{16}] he writes, “In June 1842 I first allowed myself the satisfaction of writing a very brief abstract of my theory in pencil in 35 pages; and this was enlarged during the summer of 1844 into one of 230 pages[{17}], which I had fairly copied out and still possess.” These two Essays, of 1842 and 1844, are now printed under the title The Foundations of the Origin of Species.

It will be noted that in the above passage he does not mention the MS. of 1842 as being in existence, and when I was at work on Life and Letters I had not seen it. It only came to light after my mother's death in 1896 when the house at Down was vacated. The MS. was hidden in a cupboard under the stairs which was not used for papers of any value, but rather as an overflow for matter which he did not wish to destroy.

The statement in the Autobiography that the MS. was written in 1842 agrees with an entry in my fathers Diary:—

“1842. May 18th went to Maer. June 15th to Shrewsbury, and on 18th to Capel Curig.... During my stay at Maer and Shrewsbury (five years after commencement) wrote pencil sketch of my species theory.” Again in a letter to Lyell (June 18, 1858) he speaks of his “MS. sketch written out in 1842[{18}].” In the Origin of Species, Ed. i. p. 1, he speaks of beginning his speculations in 1837 and of allowing himself to draw up some “short notes” after “five years' work,” i.e. in 1842. So far there seems no doubt as to 1842 being the date of the first sketch; but there is evidence in favour of an earlier date[{19}]. Thus across the Table of Contents of the bound copy of the 1844 MS. is written in my father's hand “This was sketched in 1839.” Again in a letter to Mr Wallace[{20}] (Jan. 25, 1859) he speaks of his own contributions to the Linnean paper[{21}] of July 1, 1858, as “written in 1839, now just twenty years ago.” This statement as it stands is undoubtedly incorrect, since the extracts are from the MS. of 1844, about the date of which no doubt exists; but even if it could be supposed to refer to the 1842 Essay, it must, I think, be rejected. I can only account for his mistake by the supposition that my father had in mind the date (1839) at which the framework of his theory was laid down. It is worth noting that in his Autobiography (p. 88) he speaks of the time “about 1839, when the theory was clearly conceived.” However this may be there can be no doubt that 1842 is the correct date. Since the publication of Life and Letters I have gained fresh evidence on this head. A small packet containing 13 pp. of MS. came to light in 1896. On the outside is written “First Pencil Sketch of Species Theory. Written at Maer and Shrewsbury during May and June 1842.” It is not however written in pencil, and it consists of a single chapter on The Principles of Variation in Domestic Organisms. A single unnumbered page is written in pencil, and is headed “Maer, May 1842, useless”; it also bears the words “This page was thought of as introduction.” It consists of the briefest sketch of the geological evidence for evolution, together with words intended as headings for discussion,—such as “Affinity,—unity of type,—fœtal state,—abortive organs.”

The back of this “useless” page is of some interest, although it does not bear on the question of date,—the matter immediately before us.

It seems to be an outline of the Essay or sketch of 1842, consisting of the titles of the three chapters of which it was to have consisted.

“I. The Principles of Var. in domestic organisms.

“II. The possible and probable application of these same principles to wild animals and consequently the possible and probable production of wild races, analogous to the domestic ones of plants and animals.

“III. The reasons for and against believing that such races have really been produced, forming what are called species.”

It will be seen that Chapter III as originally designed corresponds to Part II (p. 22) of the Essay of 1842, which is (p. 7) defined by the author as discussing “whether the characters and relations of animated things are such as favour the idea of wild species being races descended from a common stock.” Again at p. 23 the author asks “What then is the evidence in favour of it (the theory of descent) and what the evidence against it.” The generalised section of his Essay having been originally Chapter III[{22}] accounts for the curious error which occurs in pp. 18 and 22 where the second Part of the Essay is called Part III.

The division of the Essay into two parts is maintained in the enlarged Essay of 1844, in which he writes: “The Second Part of this work is devoted to the general consideration of how far the general economy of nature justifies or opposes the belief that related species and genera are descended from common stocks.” The Origin of Species however is not so divided.

We may now return to the question of the date of the Essay. I have found additional evidence in favour of 1842 in a sentence written on the back of the Table of Contents of the 1844 MS.—not the copied version but the original in my father's writing: “This was written and enlarged from a sketch in 37 pages[{23}] in Pencil (the latter written in summer of 1842 at Maer and Shrewsbury) in beginning of 1844, and finished it «sic» in July; and finally corrected the copy by Mr Fletcher in the last week in September.” On the whole it is impossible to doubt that 1842 is the date of the earlier of the two Essays.

The sketch of 1842 is written on bad paper with a soft pencil, and is in many parts extremely difficult to read, many of the words ending in mere scrawls and being illegible without context. It is evidently written rapidly, and is in his most elliptical style, the articles being frequently omitted, and the sentences being loosely composed and often illogical in structure. There is much erasure and correction, apparently made at the moment of writing, and the MS. does not give the impression of having been re-read with any care. The whole is more like hasty memoranda of what was clear to himself, than material for the convincing of others.

Many of the pages are covered with writing on the back, an instance of his parsimony in the matter of paper[{24}]. This matter consists partly of passages marked for insertion in the text, and these can generally (though by no means always) be placed where he intended. But he also used the back of one page for a preliminary sketch to be rewritten on a clean sheet. These parts of the work have been printed as footnotes, so as to allow what was written on the front of the pages to form a continuous text. A certain amount of repetition is unavoidable, but much of what is written on the backs of the pages is of too much interest to be omitted. Some of the matter here given in footnotes may, moreover, have been intended as the final text and not as the preliminary sketch.

When a word cannot be deciphered, it is replaced by:—«illegible», the angular brackets being, as already explained, a symbol for an insertion by the editor. More commonly, however, the context makes the interpretation of a word reasonably sure although the word is not strictly legible. Such words are followed by an inserted mark of interrogation «?». Lastly, words inserted by the editor, of which the appropriateness is doubtful, are printed thus «variation?».

Two kinds of erasure occur in the MS. of 1842. One by vertical lines which seem to have been made when the 35 pp. MS. was being expanded into that of 1844, and merely imply that such a page is done with: and secondly the ordinary erasures by horizontal lines. I have not been quite consistent in regard to these: I began with the intention of printing (in square brackets) all such erasures. But I ultimately found that the confusion introduced into the already obscure sentences was greater than any possible gain; and many such erasures are altogether omitted. In the same way I have occasionally omitted hopelessly obscure and incomprehensible fragments, which if printed would only have burthened the text with a string of «illegible»s and queried words. Nor have I printed the whole of what is written on the backs of the pages, where it seemed to me that nothing but unnecessary repetition would have been the result.

In the matter of punctuation I have given myself a free hand. I may no doubt have misinterpreted the author's meaning in so doing, but without such punctuation, the number of repellantly crabbed sentences would have been even greater than at present. In dealing with the Essay of 1844, I have corrected some obvious slips without indicating such alterations, because the MS. being legible, there is no danger of changing the author's meaning.

The sections into which the Essay of 1842 is divided are in the original merely indicated by a gap in the MS. or by a line drawn across the page. No titles are given except in the case of § VIII.; and § II. is the only section which has a number in the original. I might equally well have made sections of what are now subsections, e.g. Natural Selection p. 7, or Extermination p. 28. But since the present sketch is the germ of the Essay of 1844, it seemed best to preserve the identity between the two works, by using such of the author's divisions as correspond to the chapters of the enlarged version of 1844. The geological discussion with which Part II begins corresponds to two chapters (IV and V) of the 1844 Essay. I have therefore described it as §§ IV. and V., although I cannot make sure of its having originally consisted of two sections. With this exception the ten sections of the Essay of 1842 correspond to the ten chapters of that of 1844.

The Origin of Species differs from the sketch of 1842 in not being divided into two parts. But the two volumes resemble each other in general structure. Both begin with a statement of what may be called the mechanism of evolution,—variation and selection: in both the argument proceeds from the study of domestic organisms to that of animals and plants in a state of nature. This is followed in both by a discussion of the Difficulties on Theory and this by a section Instinct which in both cases is treated as a special case of difficulty.

If I had to divide the Origin (first edition) into two parts without any knowledge of earlier MS., I should, I think, make Part II begin with Ch. VI, Difficulties on Theory. A possible reason why this part of the argument is given in Part I of the Essay of 1842 may be found in the Essay of 1844, where it is clear that the chapter on instinct is placed in Part I because the author thought it of importance to show that heredity and variation occur in mental attributes. The whole question is perhaps an instance of the sort of difficulty which made the author give up the division of his argument into two Parts when he wrote the Origin. As matters stand §§ IV. and V. of the 1842 Essay correspond to the geological chapters, IX and X, in the Origin. From this point onwards the material is grouped in the same order in both works: geographical distribution; affinities and classification; unity of type and morphology; abortive or rudimentary organs; recapitulation and conclusion.

In enlarging the Essay of 1842 into that of 1844, the author retained the sections of the sketch as chapters in the completer presentment. It follows that what has been said of the relation of the earlier Essay to the Origin is generally true of the 1844 Essay. In the latter, however, the geological discussion is, clearly instead of obscurely, divided into two chapters, which correspond roughly with Chapters IX and X of the Origin. But part of the contents of Chapter X (Origin) occurs in Chapter VI (1844) on Geographical Distribution. The treatment of distribution is particularly full and interesting in the 1844 Essay, but the arrangement of the material, especially the introduction of § III. p. 183, leads to some repetition which is avoided in the Origin. It should be noted that Hybridism, which has a separate chapter (VIII) in the Origin, is treated in Chapter II of the Essay. Finally that Chapter XIII (Origin) corresponds to Chapters VII, VIII and IX of the work of 1844.

The fact that in 1842, seventeen years before the publication of the Origin, my father should have been able to write out so full an outline of his future work, is very remarkable. In his Autobiography[{25}] he writes of the 1844 Essay, “But at that time I overlooked one problem of great importance.... This problem is the tendency in organic beings descended from the same stock to diverge in character as they become modified.” The absence of the principle of divergence is of course also a characteristic of the sketch of 1842. But at p. 37, the author is not far from this point of view. The passage referred to is: “If any species, A, in changing gets an advantage and that advantage ... is inherited, A will be the progenitor of several genera or even families in the hard struggle of nature. A will go on beating out other forms, it might come that A would people «the» earth,—we may now not have one descendant on our globe of the one or several original creations[{26}].” But if the descendants of A have peopled the earth by beating out other forms, they must have diverged in occupying the innumerable diverse modes of life from which they expelled their predecessors. What I wrote[{27}] on this subject in 1887 is I think true: “Descent with modification implies divergence, and we become so habituated to a belief in descent, and therefore in divergence, that we do not notice the absence of proof that divergence is in itself an advantage.”

The fact that there is no set discussion on the principle of divergence in the 1844 Essay, makes it clear why the joint paper read before the Linnean Society on July 1, 1858, included a letter[{28}] to Asa Gray, as well as an extract[{29}] from the Essay of 1844. It is clearly because the letter to Gray includes a discussion on divergence, and was thus, probably, the only document, including this subject, which could be appropriately made use of. It shows once more how great was the importance attached by its author to the principle of divergence.

I have spoken of the hurried and condensed manner in which the sketch of 1842 is written; the style of the later Essay (1844) is more finished. It has, however, the air of an uncorrected MS. rather than of a book which has gone through the ordeal of proof sheets. It has not all the force and conciseness of the Origin, but it has a certain freshness which gives it a character of its own. It must be remembered that the Origin was an abstract or condensation of a much bigger book, whereas the Essay of 1844 was an expansion of the sketch of 1842. It is not therefore surprising that in the Origin there is occasionally evident a chafing against the author's self-imposed limitation. Whereas in the 1844 Essay there is an air of freedom, as if the author were letting himself go, rather than applying the curb. This quality of freshness and the fact that some questions were more fully discussed in 1844 than in 1859, makes the earlier work good reading even to those who are familiar with the Origin.

The writing of this Essay “during the summer of 1844,” as stated in the Autobiography[{30}], and “from memory,” as Darwin says elsewhere[{31}], was a remarkable achievement, and possibly renders more conceivable the still greater feat of the writing of the Origin between July 1858 and September 1859.

It is an interesting subject for speculation: what influence on the world the Essay of 1844 would have exercised, had it been published in place of the Origin. The author evidently thought of its publication in its present state as an undesirable expedient, as appears clearly from the following extracts from the Life and Letters, vol. ii. pp. 16—18:

C. Darwin to Mrs Darwin.

Down, July 5, 1844.

“... I have just finished my sketch of my species theory. If, as I believe, my theory in time be accepted even by one competent judge, it will be a considerable step in science.

“I therefore write this in case of my sudden death, as my most solemn and last request, which I am sure you will consider the same as if legally entered in my will, that you will devote £400 to its publication, and further will yourself, or through Hensleigh[{32}], take trouble in promoting it. I wish that my sketch be given to some competent person, with this sum to induce him to take trouble in its improvement and enlargement. I give to him all my books on Natural History, which are either scored or have references at the end to the pages, begging him carefully to look over and consider such passages as actually bearing, or by possibility bearing, on this subject. I wish you to make a list of all such books as some temptation to an editor. I also request that you will hand over «to» him all those scraps roughly divided into eight or ten brown paper portfolios. The scraps, with copied quotations from various works, are those which may aid my editor. I also request that you, or some amanuensis, will aid in deciphering any of the scraps which the editor may think possibly of use. I leave to the editor's judgment whether to interpolate these facts in the text, or as notes, or under appendices. As the looking over the references and scraps will be a long labour, and as the correcting and enlarging and altering my sketch will also take considerable time, I leave this sum of £400 as some remuneration, and any profits from the work. I consider that for this the editor is bound to get the sketch published either at a publisher's or his own risk. Many of the scraps in the portfolios contain mere rude suggestions and early views, now useless, and many of the facts will probably turn out as having no bearing on my theory.

“With respect to editors, Mr Lyell would be the best if he would undertake it; I believe he would find the work pleasant, and he would learn some facts new to him. As the editor must be a geologist as well as a naturalist, the next best editor would be Professor Forbes of London. The next best (and quite best in many respects) would be Professor Henslow. Dr Hooker would be very good. The next, Mr Strickland[{33}]. If none of these would undertake it, I would request you to consult with Mr Lyell, or some other capable man, for some editor, a geologist and naturalist. Should one other hundred pounds make the difference of procuring a good editor, I request earnestly that you will raise £500.

“My remaining collections in Natural History may be given to any one or any museum where «they» would be accepted....”

«The following note seems to have formed part of the original letter, but may have been of later date:»

“Lyell, especially with the aid of Hooker (and of any good zoological aid), would be best of all. Without an editor will pledge himself to give up time to it, it would be of no use paying such a sum.

“If there should be any difficulty in getting an editor who would go thoroughly into the subject, and think of the bearing of the passages marked in the books and copied out of scraps of paper, then let my sketch be published as it is, stating that it was done several years ago[{34}], and from memory without consulting any works, and with no intention of publication in its present form.”

The idea that the sketch of 1844 might remain, in the event of his death, as the only record of his work, seems to have been long in his mind, for in August, 1854, when he had finished with the Cirripedes, and was thinking of beginning his “species work,” he added on the back of the above letter, “Hooker by far best man to edit my species volume. August 1854.”

I have called attention in footnotes to many points in which the Origin agrees with the Foundations. One of the most interesting is the final sentence, practically the same in the Essays of 1842 and 1844, and almost identical with the concluding words of the Origin. I have elsewhere pointed out[{35}] that the ancestry of this eloquent passage may be traced one stage further back,—to the Note Book of 1837. I have given this sentence as an appropriate motto for the Foundations in its character of a study of general laws. It will be remembered that a corresponding motto from Whewell's Bridgewater Treatise is printed opposite the title-page of the Origin of Species.

Mr Huxley who, about the year 1887, read the Essay of 1844, remarked that “much more weight is attached to the influence of external conditions in producing variation and to the inheritance of acquired habits than in the Origin.” In the Foundations the effect of conditions is frequently mentioned, and Darwin seems to have had constantly in mind the need of referring each variation to a cause. But I gain the impression that the slighter prominence given to this view in the Origin was not due to change of opinion, but rather because he had gradually come to take this view for granted; so that in the scheme of that book, it was overshadowed by considerations which then seemed to him more pressing. With regard to the inheritance of acquired characters I am not inclined to agree with Huxley. It is certain that the Foundations contains strong recognition of the importance of germinal variation, that is of external conditions acting indirectly through the “reproductive functions.” He evidently considered this as more important than the inheritance of habit or other acquired peculiarities.

Another point of interest is the weight he attached in 1842-4 to “sports” or what are now called “mutations.” This is I think more prominent in the Foundations than in the first edition of the Origin, and certainly than in the fifth and sixth editions.

Among other interesting points may be mentioned the “good effects of crossing” being “possibly analogous to good effects of change in condition,”—a principle which he upheld on experimental grounds in his Cross and Self-Fertilisation in 1876.

In conclusion, I desire to express my thanks to Mr Wallace for a footnote he was good enough to supply: and to Professor Bateson, Sir W. Thiselton-Dyer, Dr Gadow, Professor Judd, Dr Marr, Col. Prain and Dr Stapf for information on various points. I am also indebted to Mr Rutherford, of the University Library, for his careful copy of the manuscript of 1842.

Cambridge,
June 9, 1909.

EXPLANATION OF SIGNS, &c.

[ ] Means that the words so enclosed are erased in the original MS.

« » Indicates an insertion by the Editor.

Origin, Ed. vi. refers to the Popular Edition.

PART I.

§ I. «On Variation under Domestication, and on the Principles of Selection.»

An individual organism placed under new conditions [often] sometimes varies in a small degree and in very trifling respects such as stature, fatness, sometimes colour, health, habits in animals and probably disposition. Also habits of life develope certain parts. Disuse atrophies. [Most of these slight variations tend to become hereditary.]

When the individual is multiplied for long periods by buds the variation is yet small, though greater and occasionally a single bud or individual departs widely from its type (example)[{36}] and continues steadily to propagate, by buds, such new kind.

When the organism is bred for several generations under new or varying conditions, the variation is greater in amount and endless in kind [especially[{37}] holds good when individuals have long been exposed to new conditions]. The nature of the external conditions tends to effect some definite change in all or greater part of offspring,—little food, small size—certain foods harmless &c. &c. organs affected and diseases—extent unknown. A certain degree of variation (Müller's twins)[{38}] seems inevitable effect of process of reproduction. But more important is that simple «?» generation, especially under new conditions [when no crossing] «causes» infinite variation and not direct effect of external conditions, but only in as much as it affects the reproductive functions[{39}]. There seems to be no part (beau ideal of liver)[{40}] of body, internal or external, or mind or habits, or instincts which does not vary in some small degree and [often] some «?» to a great amount.

[All such] variations [being congenital] or those very slowly acquired of all kinds [decidedly evince a tendency to become hereditary], when not so become simple variety, when it does a race. Each[{41}] parent transmits its peculiarities, therefore if varieties allowed freely to cross, except by the chance of two characterized by same peculiarity happening to marry, such varieties will be constantly demolished[{42}]. All bisexual animals must cross, hermaphrodite plants do cross, it seems very possible that hermaphrodite animals do cross,—conclusion strengthened: ill effects of breeding in and in, good effects of crossing possibly analogous to good effects of change in condition «?»[{43}].

Therefore if in any country or district all animals of one species be allowed freely to cross, any small tendency in them to vary will be constantly counteracted. Secondly reversion to parent form—analogue of vis medicatrix[{44}]. But if man selects, then new races rapidly formed,—of late years systematically followed,—in most ancient times often practically followed[{45}]. By such selection make race-horse, dray-horse—one cow good for tallow, another for eating &c.—one plant's good lay «illegible» in leaves another in fruit &c. &c.: the same plant to supply his wants at different times of year. By former means animals become adapted, as a direct effect to a cause, to external conditions, as size of body to amount of food. By this latter means they may also be so adapted, but further they may be adapted to ends and pursuits, which by no possibility can affect growth, as existence of tallow-chandler cannot tend to make fat. In such selected races, if not removed to new conditions, and «if» preserved from all cross, after several generations become very true, like each other and not varying. But man[{46}] selects only «?» what is useful and curious—has bad judgment, is capricious,—grudges to destroy those that do not come up to his pattern,—has no [knowledge] power of selecting according to internal variations,—can hardly keep his conditions uniform,—[cannot] does not select those best adapted to the conditions under which «the» form «?» lives, but those most useful to him. This might all be otherwise.

§ II. «On Variation in a State of Nature and on the Natural Means of Selection.»

Let us see how far above principles of variation apply to wild animals. Wild animals vary exceedingly little—yet they are known as individuals[{47}]. British Plants, in many genera number quite uncertain of varieties and species: in shells chiefly external conditions[{48}]. Primrose and cowslip. Wild animals from different [countries can be recognized]. Specific character gives some organs as varying. Variations analogous in kind, but less in degree with domesticated animals—chiefly external and less important parts.

Our experience would lead us to expect that any and every one of these organisms would vary if «the organism were» taken away «?» and placed under new conditions. Geology proclaims a constant round of change, bringing into play, by every possible «?» change of climate and the death of pre-existing inhabitants, endless variations of new conditions. These «?» generally very slow, doubtful though «illegible» how far the slowness «?» would produce tendency to vary. But Geolog«ists» show change in configuration which, together with the accidents of air and water and the means of transportal which every being possesses, must occasionally bring, rather suddenly, organism to new conditions and «?» expose it for several generations. Hence «?» we should expect every now and then a wild form to vary[{49}]; possibly this may be cause of some species varying more than others.

According to nature of new conditions, so we might expect all or majority of organisms born under them to vary in some definite way. Further we might expect that the mould in which they are cast would likewise vary in some small degree. But is there any means of selecting those offspring which vary in the same manner, crossing them and keeping their offspring separate and thus producing selected races: otherwise as the wild animals freely cross, so must such small heterogeneous varieties be constantly counter-balanced and lost, and a uniformity of character [kept up] preserved. The former variation as the direct and necessary effects of causes, which we can see can act on them, as size of body from amount of food, effect of certain kinds of food on certain parts of bodies &c. &c.; such new varieties may then become adapted to those external [natural] agencies which act on them. But can varieties be produced adapted to end, which cannot possibly influence their structure and which it is absurd to look «at» as effects of chance. Can varieties like some vars of domesticated animals, like almost all wild species be produced adapted by exquisite means to prey on one animal or to escape from another,—or rather, as it puts out of question effects of intelligence and habits, can a plant become adapted to animals, as a plant which cannot be impregnated without agency of insect; or hooked seeds depending on animal“s existence: woolly animals cannot have any direct effect on seeds of plant. This point which all theories about climate adapting woodpecker[{50}] to crawl «?» up trees, «illegible» miseltoe, «sentence incomplete». But if every part of a plant or animal was to vary «illegible», and if a being infinitely more sagacious than man (not an omniscient creator) during thousands and thousands of years were to select all the variations which tended towards certain ends ([or were to produce causes «?» which tended to the same end]), for instance, if he foresaw a canine animal would be better off, owing to the country producing more hares, if he were longer legged and keener sight,—greyhound produced[{51}]. If he saw that aquatic «animal would need» skinned toes. If for some unknown cause he found it would advantage a plant, which «?» like most plants is occasionally visited by bees &c.: if that plant’s seed were occasionally eaten by birds and were then carried on to rotten trees, he might select trees with fruit more agreeable to such birds as perched, to ensure their being carried to trees; if he perceived those birds more often dropped the seeds, he might well have selected a bird who would «illegible» rotten trees or [gradually select plants which «he» had proved to live on less and less rotten trees]. Who, seeing how plants vary in garden, what blind foolish man has done[{52}] in a few years, will deny an all-seeing being in thousands of years could effect (if the Creator chose to do so), either by his own direct foresight or by intermediate means,—which will represent «?» the creator of this universe. Seems usual means. Be it remembered I have nothing to say about life and mind and all forms descending from one common type[{53}]. I speak of the variation of the existing great divisions of the organised kingdom, how far I would go, hereafter to be seen.

Before considering whether «there» be any natural means of selection, and secondly (which forms the 2nd Part of this sketch) the far more important point whether the characters and relations of animated «things» are such as favour the idea of wild species being races «?» descended from a common stock, as the varieties of potato or dahlia or cattle having so descended, let us consider probable character of [selected races] wild varieties.

Natural Selection. De Candolle’s war of nature,—seeing contented face of nature,—may be well at first doubted; we see it on borders of perpetual cold[{54}]. But considering the enormous geometrical power of increase in every organism and as «?» every country, in ordinary cases «countries» must be stocked to full extent, reflection will show that this is the case. Malthus on man,—in animals no moral [check] restraint «?»—they breed in time of year when provision most abundant, or season most favourable, every country has its seasons,—calculate robins,—oscillating from years of destruction[{55}]. If proof were wanted let any singular change of climate «occur» here «?», how astoundingly some tribes «?» increase, also introduced animals[{56}], the pressure is always ready,—capacity of alpine plants to endure other climates,—think of endless seeds scattered abroad,—forests regaining their percentage[{57}],—a thousand wedges[{58}] are being forced into the œconomy of nature. This requires much reflection; study Malthus and calculate rates of increase and remember the resistance,—only periodical.

The unavoidable effect of this «is» that many of every species are destroyed either in egg or [young or mature (the former state the more common)]. In the course of a thousand generations infinitesimally small differences must inevitably tell[{59}]; when unusually cold winter, or hot or dry summer comes, then out of the whole body of individuals of any species, if there be the smallest differences in their structure, habits, instincts [senses], health &c., «it» will on an average tell; as conditions change a rather larger proportion will be preserved: so if the chief check to increase falls on seeds or eggs, so will, in the course of 1000 generations or ten thousand, those seeds (like one with down to fly[{60}]) which fly furthest and get scattered most ultimately rear most plants, and such small differences tend to be hereditary like shades of expression in human countenance. So if one parent «?» fish deposits its egg in infinitesimally different circumstances, as in rather shallower or deeper water &c., it will then «?» tell.

Let hares[{61}] increase very slowly from change of climate affecting peculiar plants, and some other «illegible» rabbit decrease in same proportion [let this unsettle organisation of], a canine animal, who formerly derived its chief sustenance by springing on rabbits or running them by scent, must decrease too and might thus readily become exterminated. But if its form varied very slightly, the long legged fleet ones, during a thousand years being selected, and the less fleet rigidly destroyed must, if no law of nature be opposed to it, alter forms.

Remember how soon Bakewell on the same principle altered cattle and Western, sheep,—carefully avoiding a cross (pigeons) with any breed. We cannot suppose that one plant tends to vary in fruit and another in flower, and another in flower and foliage,—some have been selected for both fruit and flower: that one animal varies in its covering and another not,—another in its milk. Take any organism and ask what is it useful for and on that point it will be found to vary,—cabbages in their leaf,—corn in size «and» quality of grain, both in times of year,—kidney beans for young pod and cotton for envelope of seeds &c. &c.: dogs in intellect, courage, fleetness and smell «?»: pigeons in peculiarities approaching to monsters. This requires consideration,—should be introduced in first chapter if it holds, I believe it does. It is hypothetical at best[{62}].

Nature’s variation far less, but such selection far more rigid and scrutinising. Man’s races not [even so well] only not better adapted to conditions than other races, but often not «?» one race adapted to its conditions, as man keeps and propagates some alpine plants in garden. Nature lets «an» animal live, till on actual proof it is found less able to do the required work to serve the desired end, man judges solely by his eye, and knows not whether nerves, muscles, arteries, are developed in proportion to the change of external form.

Besides selection by death, in bisexual animals «illegible» the selection in time of fullest vigour, namely struggle of males; even in animals which pair there seems a surplus «?» and a battle, possibly as in man more males produced than females, struggle of war or charms[{63}]. Hence that male which at that time is in fullest vigour, or best armed with arms or ornaments of its species, will gain in hundreds of generations some small advantage and transmit such characters to its offspring. So in female rearing its young, the most vigorous and skilful and industrious, «whose» instincts «are» best developed, will rear more young, probably possessing her good qualities, and a greater number will thus «be» prepared for the struggle of nature. Compared to man using a male alone of good breed. This latter section only of limited application, applies to variation of [specific] sexual characters. Introduce here contrast with Lamarck,—absurdity of habit, or chance?? or external conditions, making a woodpecker adapted to tree[{64}].

Before considering difficulties of theory of selection let us consider character of the races produced, as now explained, by nature. Conditions have varied slowly and the organisms best adapted in their whole course of life to the changed conditions have always been selected,—man selects small dog and afterwards gives it profusion of food,—selects a long-backed and short-legged breed and gives it no particular exercise to suit this function &c. &c. In ordinary cases nature has not allowed her race to be contaminated with a cross of another race, and agriculturists know how difficult they find always to prevent this,—effect would be trueness. This character and sterility when crossed, and generally a greater amount of difference, are two main features, which distinguish domestic races from species.

[Sterility not universal admitted by all[{65}]. Gladiolus, Crinum, Calceolaria[{66}] must be species if there be such a thing. Races of dogs and oxen: but certainly very general; indeed a gradation of sterility most perfect[{67}] very general. Some nearest species will not cross (crocus, some heath «?»), some genera cross readily (fowls[{68}] and grouse, peacock &c.). Hybrids no ways monstrous quite perfect except secretions[{69}] hence even the mule has bred,—character of sterility, especially a few years ago «?» thought very much more universal than it now is, has been thought the distinguishing character; indeed it is obvious if all forms freely crossed, nature would be a chaos. But the very gradation of the character, even if it always existed in some degree which it does not, renders it impossible as marks «?» those «?» suppose distinct as species[{70}]]. Will analogy throw any light on the fact of the supposed races of nature being sterile, though none of the domestic ones are? Mr Herbert «and» Koelreuter have shown external differences will not guide one in knowing whether hybrids will be fertile or not, but the chief circumstance is constitutional differences[{71}], such as being adapted to different climate or soil, differences which [must] probably affect the whole body of the organism and not any one part. Now wild animals, taken out of their natural conditions, seldom breed. I do not refer to shows or to Zoological Societies where many animals unite, but «do not?» breed, and others will never unite, but to wild animals caught and kept quite tame left loose and well fed about houses and living many years. Hybrids produced almost as readily as pure breds. St Hilaire great distinction of tame and domestic,—elephants,—ferrets[{72}]. Reproductive organs not subject to disease in Zoological Garden. Dissection and microscope show that hybrid is in exactly same condition as another animal in the intervals of breeding season, or those animals which taken wild and not bred in domesticity, remain without breeding their whole lives. It should be observed that so far from domesticity being unfavourable in itself «it» makes more fertile: [when animal is domesticated and breeds, productive power increased from more food and selection of fertile races]. As far as animals go might be thought «an» effect on their mind and a special case.

But turning to plants we find same class of facts. I do not refer to seeds not ripening, perhaps the commonest cause, but to plants not setting, which either is owing to some imperfection of ovule or pollen. Lindley says sterility is the [curse] bane of all propagators,—Linnæus about alpine plants. American bog plants,—pollen in exactly same state as in hybrids,—same in geraniums. Persian and Chinese[{73}] lilac will not seed in Italy and England. Probably double plants and all fruits owe their developed parts primarily «?» to sterility and extra food thus «?» applied[{74}]. There is here gradation «in» sterility and then parts, like diseases, are transmitted hereditarily. We cannot assign any cause why the Pontic Azalea produces plenty of pollen and not American[{75}], why common lilac seeds and not Persian, we see no difference in healthiness. We know not on what circumstances these facts depend, why ferret breeds, and cheetah[{76}], elephant and pig in India will not.

Now in crossing it is certain every peculiarity in form and constitution is transmitted: an alpine plant transmits its alpine tendency to its offspring, an American plant its American-bog constitution, and «with» animals, those peculiarities, on which[{77}] when placed out of their natural conditions they are incapable of breeding; and moreover they transmit every part of their constitution, their respiration, their pulse, their instinct, which are all suddenly modified, can it be wondered at that they are incapable of breeding? I think it may be truly said it would be more wonderful if they did. But it may be asked why have not the recognised varieties, supposed to have been produced through the means of man, [not refused to breed] have all bred[{78}]. Variation depends on change of condition and selection[{79}], as far as man’s systematic or unsystematic selection «has» gone; he takes external form, has little power from ignorance over internal invisible constitutional differences. Races which have long been domesticated, and have much varied, are precisely those which were capable of bearing great changes, whose constitutions were adapted to a diversity of climates. Nature changes slowly and by degrees. According to many authors probably breeds of dogs are another case of modified species freely crossing. There is no variety which «illegible» has been «illegible» adapted to peculiar soil or situation for a thousand years and another rigorously adapted to another, till such can be produced, the question is not tried[{80}]. Man in past ages, could transport into different climates, animals and plants which would freely propagate in such new climates. Nature could effect, with selection, such changes slowly, so that precisely those animals which are adapted to submit to great changes have given rise to diverse races,—and indeed great doubt on this head[{81}].

Before leaving this subject well to observe that it was shown that a certain amount of variation is consequent on mere act of reproduction, both by buds and sexually,—is vastly increased when parents exposed for some generations to new conditions[{82}], and we now find that many animals when exposed for first time to very new conditions, are «as» incapable of breeding as hybrids. It [probably] bears also on supposed fact of crossed animals when not infertile, as in mongrels, tending to vary much, as likewise seems to be the case, when true hybrids possess just sufficient fertility to propagate with the parent breeds and inter se for some generations. This is Koelreuter’s belief. These facts throw light on each other and support the truth of each other, we see throughout a connection between the reproductive faculties and exposure to changed conditions of life whether by crossing or exposure of the individuals[{83}].

Difficulties on theory of selection[{84}]. It may be objected such perfect organs as eye and ear, could never be formed, in latter less difficulty as gradations more perfect; at first appears monstrous and to «the» end appears difficulty. But think of gradation, even now manifest, (Tibia and Fibula). Everyone will allow if every fossil preserved, gradation infinitely more perfect; for possibility of selection a perfect «?» gradation is required. Different groups of structure, slight gradation in each group,—every analogy renders it probable that intermediate forms have existed. Be it remembered what strange metamorphoses; part of eye, not directly connected with vision, might come to be [thus used] gradually worked in for this end,—swimming bladder by gradation of structure is admitted to belong to the ear system,—rattlesnake. [Woodpecker best adapted to climb.] In some cases gradation not possible,—as vertebræ,—actually vary in domestic animals,—less difficult if growth followed. Looking to whole animals, a bat formed not for flight[{85}]. Suppose we had flying fish[{86}] and not one of our now called flying fish preserved, who would have guessed intermediate habits. Woodpeckers and tree-frogs both live in countries where no trees[{87}].

The gradations by which each individual organ has arrived at its present state, and each individual animal with its aggregate of organs has arrived, probably never could be known, and all present great difficulties. I merely wish to show that the proposition is not so monstrous as it at first appears, and that if good reason can be advanced for believing the species have descended from common parents, the difficulty of imagining intermediate forms of structure not sufficient to make one at once reject the theory.

§ III. «On Variation in instincts and other mental attributes.»

The mental powers of different animals in wild and tame state [present still greater difficulties] require a separate section. Be it remembered I have nothing to do with origin of memory, attention, and the different faculties of the mind[{88}], but merely with their differences in each of the great divisions of nature. Disposition, courage, pertinacity «?», suspicion, restlessness, ill-temper, sagacity and «the» reverse unquestionably vary in animals and are inherited (Cuba wildness dogs, rabbits, fear against particular object as man Galapagos[{89}]). Habits purely corporeal, breeding season &c., time of going to rest &c., vary and are hereditary, like the analogous habits of plants which vary and are inherited. Habits of body, as manner of movement d^o. and d^o. Habits, as pointing and setting on certain occasions d^o. Taste for hunting certain objects and manner of doing so,—sheep-dog. These are shown clearly by crossing and their analogy with true instinct thus shown,—retriever. Do not know objects for which they do it. Lord Brougham’s definition[{90}]. Origin partly habit, but the amount necessarily unknown, partly selection. Young pointers pointing stones and sheep—tumbling pigeons—sheep[{91}] going back to place where born. Instinct aided by reason, as in the taylor-bird[{92}]. Taught by parents, cows choosing food, birds singing. Instincts vary in wild state (birds get wilder) often lost[{93}]; more perfect,—nest without roof. These facts [only clear way] show how incomprehensibly brain has power of transmitting intellectual operations.

Faculties[{94}] distinct from true instincts,—finding [way]. It must I think be admitted that habits whether congenital or acquired by practice [sometimes] often become inherited[{95}]; instincts, influence, equally with structure, the preservation of animals; therefore selection must, with changing conditions tend to modify the inherited habits of animals. If this be admitted it will be found possible that many of the strangest instincts may be thus acquired. I may observe, without attempting definition, that an inherited habit or trick (trick because may be born) fulfils closely what we mean by instinct. A habit is often performed unconsciously, the strangest habits become associated, d^o. tricks, going in certain spots &c. &c., even against will, is excited by external agencies, and looks not to the end,—a person playing a pianoforte. If such a habit were transmitted it would make a marvellous instinct. Let us consider some of the most difficult cases of instincts, whether they could be possibly acquired. I do not say probably, for that belongs to our 3rd Part[{96}], I beg this may be remembered, nor do I mean to attempt to show exact method. I want only to show that whole theory ought not at once to be rejected on this score.

Every instinct must, by my theory, have been acquired gradually by slight changes «illegible» of former instinct, each change being useful to its then species. Shamming death struck me at first as remarkable objection. I found none really sham death[{97}], and that there is gradation; now no one doubts that those insects which do it either more or less, do it for some good, if then any species was led to do it more, and then «?» escaped &c. &c.

Take migratory instincts, faculty distinct from instinct, animals have notion of time,—like savages. Ordinary finding way by memory, but how does savage find way across country,—as incomprehensible to us, as animal to them,—geological changes,—fishes in river,—case of sheep in Spain[{98}]. Architectural instincts,—a manufacturer’s employee in making single articles extraordinary skill,—often said seem to make it almost «illegible», child born with such a notion of playing[{99}],—we can fancy tailoring acquired in same perfection,—mixture of reason,—water-ouzel,—taylor-bird,—gradation of simple nest to most complicated.

Bees again, distinction of faculty,—how they make a hexagon,—Waterhouse’s theory[{100}],—the impulse to use whatever faculty they possess,—the taylor-bird has the faculty of sewing with beak, instinct impels him to do it.

Last case of parent feeding young with different food (take case of Galapagos birds, gradation from Hawfinch to Sylvia) selection and habit might lead old birds to vary taste «?» and form, leaving their instinct of feeding their young with same food[{101}],—or I see no difficulty in parents being forced or induced to vary the food brought, and selection adapting the young ones to it, and thus by degree any amount of diversity might be arrived at. Although we can never hope to see the course revealed by which different instincts have been acquired, for we have only present animals (not well known) to judge of the course of gradation, yet once grant the principle of habits, whether congenital or acquired by experience, being inherited and I can see no limit to the [amount of variation] extraordinariness «?» of the habits thus acquired.

Summing up this Division. If variation be admitted to occur occasionally in some wild animals, and how can we doubt it, when we see [all] thousands «of» organisms, for whatever use taken by man, do vary. If we admit such variations tend to be hereditary, and how can we doubt it when we «remember» resemblances of features and character,—disease and monstrosities inherited and endless races produced (1200 cabbages). If we admit selection is steadily at work, and who will doubt it, when he considers amount of food on an average fixed and reproductive powers act in geometrical ratio. If we admit that external conditions vary, as all geology proclaims, they have done and are now doing,—then, if no law of nature be opposed, there must occasionally be formed races, [slightly] differing from the parent races. So then any such law[{102}], none is known, but in all works it is assumed, in «?» flat contradiction to all known facts, that the amount of possible variation is soon acquired. Are not all the most varied species, the oldest domesticated: who «would» think that horses or corn could be produced? Take dahlia and potato, who will pretend in 5000 years[{103}] «that great changes might not be effected»: perfectly adapted to conditions and then again brought into varying conditions. Think what has been done in few last years, look at pigeons, and cattle. With the amount of food man can produce he may have arrived at limit of fatness or size, or thickness of wool «?», but these are the most trivial points, but even in these I conclude it is impossible to say we know the limit of variation. And therefore with the [adapting] selecting power of nature, infinitely wise compared to those of man, «I conclude» that it is impossible to say we know the limit of races, which would be true «to their» kind; if of different constitutions would probably be infertile one with another, and which might be adapted in the most singular and admirable manner, according to their wants, to external nature and to other surrounding organisms,—such races would be species. But is there any evidence «that» species «have» been thus produced, this is a question wholly independent of all previous points, and which on examination of the kingdom of nature «we» ought to answer one way or another.

PART II[{104}].

§§ IV. & V. «On the evidence from Geology.»

I may premise, that according to the view ordinarily received, the myriads of organisms peopling this world have been created by so many distinct acts of creation. As we know nothing of the «illegible» will of a Creator,—we can see no reason why there should exist any relation between the organisms thus created; or again, they might be created according to any scheme. But it would be marvellous if this scheme should be the same as would result from the descent of groups of organisms from [certain] the same parents, according to the circumstances, just attempted to be developed.

With equal probability did old cosmogonists say fossils were created, as we now see them, with a false resemblance to living beings[{105}]; what would the Astronomer say to the doctrine that the planets moved «not» according to the law of gravitation, but from the Creator having willed each separate planet to move in its particular orbit? I believe such a proposition (if we remove all prejudices) would be as legitimate as to admit that certain groups of living and extinct organisms, in their distribution, in their structure and in their relations one to another and to external conditions, agreed with the theory and showed signs of common descent, and yet were created distinct. As long as it was thought impossible that organisms should vary, or should anyhow become adapted to other organisms in a complicated manner, and yet be separated from them by an impassable barrier of sterility[{106}], it was justifiable, even with some appearance in favour of a common descent, to admit distinct creation according to the will of an Omniscient Creator; or, for it is the same thing, to say with Whewell that the beginnings of all things surpass the comprehension of man. In the former sections I have endeavoured to show that such variation or specification is not impossible, nay, in many points of view is absolutely probable. What then is the evidence in favour of it and what the evidence against it. With our imperfect knowledge of past ages [surely there will be some] it would be strange if the imperfection did not create some unfavourable evidence.

Give sketch of the Past,—beginning with facts appearing hostile under present knowledge,—then proceed to geograph. distribution,—order of appearance,—affinities,—morphology &c., &c.

Our theory requires a very gradual introduction of new forms[{107}], and extermination of the old (to which we shall revert). The extermination of old may sometimes be rapid, but never the introduction. In the groups descended from common parent, our theory requires a perfect gradation not differing more than breed«s» of cattle, or potatoes, or cabbages in forms. I do not mean that a graduated series of animals must have existed, intermediate between horse, mouse, tapir[{108}], elephant [or fowl and peacock], but that these must have had a common parent, and between horse and this «?» parent &c., &c., but the common parent may possibly have differed more from either than the two do now from each other. Now what evidence of this is there? So perfect gradation in some departments, that some naturalists have thought that in some large divisions, if all existing forms were collected, a near approach to perfect gradation would be made. But such a notion is preposterous with respect to all, but evidently so with mammals. Other naturalists have thought this would be so if all the specimens entombed in the strata were collected[{109}]. I conceive there is no probability whatever of this; nevertheless it is certain all the numerous fossil forms fall in«to», as Buckland remarks, not present classes, families and genera, they fall between them: so is it with new discoveries of existing forms. Most ancient fossils, that is most separated «by» space of time, are most apt to fall between the classes—(but organisms from those countries most separated by space also fall between the classes «e.g.» Ornithorhyncus?). As far as geological discoveries «go» they tend towards such gradation[{110}]. Illustrate it with net. Toxodon,—tibia and fibula,—dog and otter,—but so utterly improbable is «it», in ex. gr. Pachydermata, to compose series as perfect as cattle, that if, as many geologists seem to infer, each separate formation presents even an approach to a consecutive history, my theory must be given up. Even if it were consecutive, it would only collect series of one district in our present state of knowledge; but what probability is there that any one formation during the immense period which has elapsed during each period will generally present a consecutive history. [Compare number living at one period to fossils preserved—look at enormous periods of time.]

Referring only to marine animals, which are obviously most likely to be preserved, they must live where «?» sediment (of a kind favourable for preservation, not sand and pebble)[{111}] is depositing quickly and over large area and must be thickly capped, «illegible» littoral deposits: for otherwise denudation «will destroy them»,—they must live in a shallow space which sediment will tend to fill up,—as movement is «in?» progress if soon brought «?» up «?» subject to denudation,—[if] as during subsidence favourable, accords with facts of European deposits[{112}], but subsidence apt to destroy agents which produce sediment[{113}].

I believe safely inferred «that» groups of marine «?» fossils only preserved for future ages where sediment goes on long «and» continuous«ly» and with rapid but not too rapid deposition in «an» area of subsidence. In how few places in any one region like Europe will «?» these contingencies be going on? Hence «?» in past ages mere [gaps] pages preserved[{114}]. Lyell's doctrine carried to extreme,—we shall understand difficulty if it be asked:—what chance of series of gradation between cattle by «illegible» at age «illegible» as far back as Miocene[{115}]? We know then cattle existed. Compare number of living,—immense duration of each period,—fewness of fossils.

This only refers to consecutiveness of history of organisms of each formation.

The foregoing argument will show firstly, that formations are distinct merely from want of fossils «of intermediate beds», and secondly, that each formation is full of gaps, has been advanced to account for fewness of preserved organisms compared to what have lived on the world. The very same argument explains why in older formations the organisms appear to come on and disappear suddenly,—but in [later] tertiary not quite suddenly[{116}], in later tertiary gradually,—becoming rare and disappearing,—some have disappeared within man’s time. It is obvious that our theory requires gradual and nearly uniform introduction, possibly more sudden extermination,—subsidence of continent of Australia &c., &c.

Our theory requires that the first form which existed of each of the great divisions would present points intermediate between existing ones, but immensely different. Most geologists believe Silurian[{117}] fossils are those which first existed in the whole world, not those which have chanced to be the oldest not destroyed,—or the first which existed in profoundly deep seas in progress of conversion from sea to land: if they are first they «? we» give up. Not so Hutton or Lyell: if first reptile[{118}] of Red Sandstone «?» really was first which existed: if Pachyderm[{119}] of Paris was first which existed: fish of Devonian: dragon fly of Lias: for we cannot suppose them the progenitors: they agree too closely with existing divisions. But geologists consider Europe as «?» a passage from sea to island «?» to continent (except Wealden, see Lyell). These animals therefore, I consider then mere introduction «?» from continents long since submerged.

Finally, if views of some geologists be correct, my theory must be given up. [Lyell’s views, as far as they go, are in favour, but they go so little in favour, and so much more is required, that it may «be» viewed as objection.] If geology present us with mere pages in chapters, towards end of «a» history, formed by tearing out bundles of leaves, and each page illustrating merely a small portion of the organisms of that time, the facts accord perfectly with my theory[{120}].

Extermination. We have seen that in later periods the organisms have disappeared by degrees and [perhaps] probably by degrees in earlier, and I have said our theory requires it. As many naturalists seem to think extermination a most mysterious circumstance[{121}] and call in astonishing agencies, it is well to recall what we have shown concerning the struggle of nature. An exterminating agency is at work with every organism: we scarcely see it: if robins would increase to thousands in ten years how severe must the process be. How imperceptible a small increase: fossils become rare: possibly sudden extermination as Australia, but as present means very slow and many means of escape, I shall doubt very sudden exterminations. Who can explain why some species abound more,—why does marsh titmouse, or ring-ouzel, now little change,—why is one sea-slug rare and another common on our coasts,—why one species of Rhinoceros more than another,—why is «illegible» tiger of India so rare? Curious and general sources of error, the place of an organism is instantly filled up.

We know state of earth has changed, and as earthquakes and tides go on, the state must change,—many geologists believe a slow gradual cooling. Now let us see in accordance with principles of [variation] specification explained in Sect. II. how species would probably be introduced and how such results accord with what is known.

The first fact geology proclaims is immense number of extinct forms, and new appearances. Tertiary strata leads to belief, that forms gradually become rare and disappear and are gradually supplied by others. We see some forms now becoming rare and disappearing, we know of no sudden creation: in older periods the forms appear to come in suddenly, scene shifts: but even here Devonian, Permian &c. [keep on supplying new links in chain]—Genera and higher forms come on and disappear, in same way leaving a species on one or more stages below that in which the form abounded.

«Geographical Distribution.»

§ VI. Let us consider the absolute state of distribution of organisms of earth's face.

Referring chiefly, but not exclusively (from difficulty of transport, fewness, and the distinct characteristics of groups) to Mammalia; and first considering the three or four main [regions] divisions; North America, Europe, Asia, including greater part of E. Indian Archipelago and Africa are intimately allied. Africa most distinct, especially most southern parts. And the Arctic regions, which unite N. America, Asia and Europe, only separated (if we travel one way by Behring’s St.) by a narrow strait, is most intimately allied, indeed forms but one restricted group. Next comes S. America,—then Australia, Madagascar (and some small islands which stand very remote from the land). Looking at these main divisions separately, the organisms vary according to changes in condition[{122}] of different parts. But besides this, barriers of every kind seem to separate regions in a greater degree than proportionally to the difference of climates on each side. Thus great chains of mountains, spaces of sea between islands and continents, even great rivers and deserts. In fact the amount «of» difference in the organisms bears a certain, but not invariable relation to the amount of physical difficulties to transit[{123}].

There are some curious exceptions, namely, similarity of fauna of mountains of Europe and N. America and Lapland. Other cases just «the» reverse, mountains of eastern S. America, Altai «?», S. India «?»[{124}]: mountain summits of islands often eminently peculiar. Fauna generally of some islands, even when close, very dissimilar, in others very similar. [I am here led to observe one or more centres of creation[{125}].]

The simple geologist can explain many of the foregoing cases of distribution. Subsidence of a continent in which free means of dispersal, would drive the lowland plants up to the mountains, now converted into islands, and the semi-alpine plants would take place of alpine, and alpine be destroyed, if mountains originally were not of great height. So we may see, during gradual changes[{126}] of climate on a continent, the propagation of species would vary and adapt themselves to small changes causing much extermination[{127}]. The mountains of Europe were quite lately covered with ice, and the lowlands probably partaking of the Arctic climate and Fauna. Then as climate changed, arctic fauna would take place of ice, and an inundation of plants from different temperate countries «would» seize the lowlands, leaving islands of arctic forms. But if this had happened on an island, whence could the new forms have come,—here the geologist calls in creationists. If island formed, the geologist will suggest «that» many of the forms might have been borne from nearest land, but if peculiar, he calls in creationist,—as such island rises in height &c., he still more calls in creation. The creationist tells one, on a «illegible» spot the American spirit of creation makes Orpheus and Tyrannus and American doves, and in accordance with past and extinct forms, but no persistent relation between areas and distribution, Geologico-Geograph.-Distribution.

Now according to analogy of domesticated animals let us see what would result. Let us take case of farmer on Pampas, where everything approaches nearer to state of nature. He works on organisms having strong tendency to vary: and he knows «that the» only way to make a distinct breed is to select and separate. It would be useless to separate the best bulls and pair with best cows if their offspring run loose and bred with the other herds, and tendency to reversion not counteracted; he would endeavour therefore to get his cows on islands and then commence his work of selection. If several farmers in different rincons[{128}] were to set to work, especially if with different objects, several breeds would soon be produced. So would it be with horticulturist and so history of every plant shows; the number of varieties[{129}] increase in proportion to care bestowed on their selection and, with crossing plants, separation. Now, according to this analogy, change of external conditions, and isolation either by chance landing «of» a form on an island, or subsidence dividing a continent, or great chain of mountains, and the number of individuals not being numerous will best favour variation and selection[{130}]. No doubt change could be effected in same country without any barrier by long continued selection on one species: even in case of a plant not capable of crossing would easier get possession and solely occupy an island[{131}]. Now we can at once see that «if» two parts of a continent isolated, new species thus generated in them, would have closest affinities, like cattle in counties of England: if barrier afterwards destroyed one species might destroy the other or both keep their ground. So if island formed near continent, let it be ever so different, that continent would supply inhabitants, and new species (like the old) would be allied with that continent. An island generally very different soil and climate, and number and order of inhabitants supplied by chance, no point so favourable for generation of new species[{132}],—especially the mountains, hence, so it is. As isolated mountains formed in a plain country (if such happens) is an island. As other islands formed, the old species would spread and thus extend and the fauna of distant island might ultimately meet and a continent formed between them. No one doubts continents formed by repeated elevations and depressions[{133}]. In looking backwards, but not so far that all geographical boundaries are destroyed, we can thus at once see why existing forms are related to the extinct in the same manner as existing ones are in some part of existing continent. By chance we might even have one or two absolute parent fossils.

The detection of transitional forms would be rendered more difficult on rising point of land.

The distribution therefore in the above enumerated points, even the trivial ones, which on any other «theory?» can be viewed as so many ultimate facts, all follow «in» a simple manner on the theory of the occurrence of species by «illegible» and being adapted by selection to «illegible», conjoined with their power of dispersal, and the steady geographico-geological changes which are now in progress and which undoubtedly have taken place. Ought to state the opinion of the immutability of species and the creation by so many separate acts of will of the Creator[{134}].

§ VII. «Affinities and Classification.»

Looking now to the affinities of organisms, without relation to their distribution, and taking all fossil and recent, we see the degrees of relationship are of different degrees and arbitrary,—sub-genera,—genera,—sub-families, families, orders and classes and kingdoms. The kind of classification which everyone feels is most correct is called the natural system, but no can define this. If we say with Whewell «that we have an» undefined instinct of the importance of organs[{135}], we have no means in lower animals of saying which is most important, and yet everyone feels that some one system alone deserves to be called natural. The true relationship of organisms is brought before one by considering relations of analogy, an otter-like animal amongst mammalia and an otter amongst marsupials. In such cases external resemblance and habit of life and the final end of whole organization very strong, yet no relation[{136}]. Naturalists cannot avoid these terms of relation and affinity though they use them metaphorically. If used in simple earnestness the natural system ought to be a genealogical «one»; and our knowledge of the points which are most easily affected in transmission are those which we least value in considering the natural system, and practically when we find they do vary we regard them of less value[{137}]. In classifying varieties the same language is used and the same kind of division: here also (in pine-apple)[{138}] we talk of the natural classification, overlooking similarity of the fruits, because whole plant differs. The origin of sub-genera, genera, &c., &c., is not difficult on notion of genealogical succession, and accords with what we know of similar gradations of affinity in domesticated organisms. In the same region the organic beings are «illegible» related to each other and the external conditions in many physical respects are allied[{139}] and their differences of same kind, and therefore when a new species has been selected and has obtained a place in the economy of nature, we may suppose that generally it will tend to extend its range during geographical changes, and thus, becoming isolated and exposed to new conditions, will slightly alter and its structure by selection become slightly remodified, thus we should get species of a sub-genus and genus,—as varieties of merino-sheep,—varieties of British and Indian cattle. Fresh species might go on forming and others become extinct and all might become extinct, and then we should have «an» extinct genus; a case formerly mentioned, of which numerous cases occur in Palæontology. But more often the same advantages which caused the new species to spread and become modified into several species would favour some of the species being preserved: and if two of the species, considerably different, each gave rise to group of new species, you would have two genera; the same thing will go on. We may look at case in other way, looking to future. According to mere chance every existing species may generate another, but if any species, A, in changing gets an advantage and that advantage (whatever it may be, intellect, &c., &c., or some particular structure or constitution) is inherited[{140}], A will be the progenitor of several genera or even families in the hard struggle of nature. A will go on beating out other forms, it might come that A would people earth,—we may now not have one descendant on our globe of the one or several original creations[{141}]. External conditions air, earth, water being same[{142}] on globe, and the communication not being perfect, organisms of widely different descent might become adapted to the same end and then we should have cases of analogy[{143}], [they might even tend to become numerically representative]. From this often happening each of the great divisions of nature would have their representative eminently adapted to earth, to «air»[{144}], to water, and to these in «illegible» and then these great divisions would show numerical relations in their classification.

§ VIII. Unity [or similarity] of type in the great classes.

Nothing more wonderful in Nat. Hist. than looking at the vast number of organisms, recent and fossil, exposed to the most diverse conditions, living in the most distant climes, and at immensely remote periods, fitted to wholely different ends, yet to find large groups united by a similar type of structure. When we for instance see bat, horse, porpoise-fin, hand, all built on same structure[{145}], having bones[{146}] with same name, we see there is some deep bond of union between them[{147}], to illustrate this is the foundation and objects «?» «of» what is called the Natural System; and which is foundation of distinction «?» of true and adaptive characters[{148}]. Now this wonderful fact of hand, hoof, wing, paddle and claw being the same, is at once explicable on the principle of some parent-forms, which might either be «illegible» or walking animals, becoming through infinite number of small selections adapted to various conditions. We know that proportion, size, shape of bones and their accompanying soft parts vary, and hence constant selection would alter, to almost any purpose «?» the framework of an organism, but yet would leave a general, even closest similarity in it.

[We know the number of similar parts, as vertebræ and ribs can vary, hence this also we might expect.] Also «if» the changes carried on to a certain point, doubtless type will be lost, and this is case with Plesiosaurus[{149}]. The unity of type in past and present ages of certain great divisions thus undoubtedly receives the simplest explanation.

There is another class of allied and almost identical facts, admitted by the soberest physiologists, [from the study of a certain set of organs in a group of organisms] and refers «? referring» to a unity of type of different organs in the same individual, denominated the science of “Morphology.” The «? this» discovered by beautiful and regular series, and in the case of plants from monstrous changes, that certain organs in an individual are other organs metamorphosed. Thus every botanist considers petals, nectaries, stamens, pistils, germen as metamorphosed leaf. They thus explain, in the most lucid manner, the position and number of all parts of the flower, and the curious conversion under cultivation of one part into another. The complicated double set of jaws and palpi of crustaceans[{150}], and all insects are considered as metamorphosed «limbs» and to see the series is to admit this phraseology. The skulls of the vertebrates are undoubtedly composed of three metamorphosed vertebræ; thus we can understand the strange form of the separate bones which compose the casket holding man’s brain. These[{151}] facts differ but slightly from those of last section, if with wing, paddle, hand and hoof, some common structure was yet visible, or could be made out by a series of occasional monstrous conversions, and if traces could be discovered of «the» whole having once existed as walking or swimming instruments, these organs would be said to be metamorphosed, as it is they are only said to exhibit a common type.

This distinction is not drawn by physiologists, and is only implied by some by their general manner of writing. These facts, though affecting every organic being on the face of the globe, which has existed, or does exist, can only be viewed by the Creationist as ultimate and inexplicable facts. But this unity of type through the individuals of a group, and this metamorphosis of the same organ into other organs, adapted to diverse use, necessarily follows on the theory of descent[{152}]. For let us take case of Vertebrata, which if[{153}] they descended from one parent and by this theory all the Vertebrata have been altered by slow degrees, such as we see in domestic animals. We know that proportions alter, and even that occasionally numbers of vertebræ alter, that parts become soldered, that parts are lost, as tail and toes, but we know «that?» here we can see that possibly a walking organ might «?» be converted into swimming or into a gliding organ and so on to a flying organ. But such gradual changes would not alter the unity of type in their descendants, as parts lost and soldered and vertebræ. But we can see that if this carried to extreme, unity lost,—Plesiosaurus. Here we have seen the same organ is formed «?» «for» different purposes «ten words illegible»: and if, in several orders of vertebrata, we could trace origin «of» spinous processes and monstrosities &c. we should say, instead of there existing a unity of type, morphology[{154}], as we do when we trace the head as being the vertebræ metamorphosed. Be it observed that Naturalists, as they use terms of affinity without attaching real meaning, here also they are obliged to use metamorphosis, without meaning that any parent of crustacean was really an animal with as many legs as crustacean has jaws. The theory of descent at once explains these wonderful facts.

Now few of the physiologists who use this language really suppose that the parent of insect with the metamorphosed jaw, was an insect with [more] so many legs, or that the parent of flowering plants, originally had no stamens, or pistils or petals, but some other means of propagation,—and so in other cases. Now according to our theory during the infinite number of changes, we might expect that an organ used for a purpose might be used for a different one by his descendant, as must have been the case by our theory with the bat, porpoise, horse, &c., which are descended from one parent. And if it so chanced that traces of the former use and structure of the part should be retained, which is manifestly possible if not probable, then we should have the organs, on which morphology is founded and which instead of being metaphorical becomes plain and «and instead of being» utterly unintelligible becomes simple matter of fact[{155}].

«Embryology.» This general unity of type in great groups of organisms (including of course these morphological cases) displays itself in a most striking manner in the stages through which the fœtus passes[{156}]. In early stage, the wing of bat, hoof, hand, paddle are not to be distinguished. At a still earlier «stage» there is no difference between fish, bird, &c. &c. and mammal. It is not that they cannot be distinguished, but the arteries[{157}] «illegible». It is not true that one passes through the form of a lower group, though no doubt fish more nearly related to fœtal state[{158}].

This similarity at the earliest stage is remarkably shown in the course of the arteries which become greatly altered, as fœtus advances in life and assumes the widely different course and number which characterize full-grown fish and mammals. How wonderful that in egg, in water or air, or in womb of mother, artery[{159}] should run in same course.

Light can be thrown on this by our theory. The structure of each organism is chiefly adapted to the sustension of its life, when full-grown, when it has to feed itself and propagate[{160}]. The structure of a kitten is quite in secondary degree adapted to its habits, whilst fed by its mother’s milk and prey. Hence variation in the structure of the full-grown species will chiefly determine the preservation of a species now become ill-suited to its habitat, or rather with a better place opened to it in the economy of Nature. It would not matter to the full-grown cat whether in its young state it was more or less eminently feline, so that it become so when full-grown. No doubt most variation, (not depending on habits of life of individual) depends on early change[{161}] and we must suspect that at whatever time of life the alteration of fœtus is effected, it tends to appear at same period. When we «see» a tendency to particular disease in old age transmitted by the male, we know some effect is produced during conception, on the simple cell of ovule, which will not produce its effect till half a century afterwards and that effect is not visible[{162}]. So we see in grey-hound, bull-dog, in race-horse and cart-horse, which have been selected for their form in full-life, there is much less «?» difference in the few first days after birth[{163}], than when full-grown: so in cattle, we see it clearly in cases of cattle, which differ obviously in shape and length of horns. If man were during 10,000 years to be able to select, far more diverse animals from horse or cow, I should expect there would be far less differences in the very young and fœtal state: and this, I think, throws light on above marvellous fact. In larvæ, which have long life selection, perhaps, does much,—in the pupa not so much[{164}] There is no object gained in varying form &c. of fœtus (beyond certain adaptations to mother’s womb) and therefore selection will not further act on it, than in giving to its changing tissues a tendency to certain parts afterwards to assume certain forms.

Thus there is no power to change the course of the arteries, as long as they nourish the fœtus; it is the selection of slight changes which supervene at any time during «illegible» of life.

The less differences of fœtus,—this has obvious meaning on this view: otherwise how strange that a [monkey] horse, a man, a bat should at one time of life have arteries, running in a manner, which is only intelligibly useful in a fish! The natural system being on theory genealogical, we can at once see, why fœtus, retaining traces of the ancestral form, is of the highest value in classification.

§ IX. «Abortive organs.»

There is another grand class of facts relating to what are called abortive organs. These consist of organs which the same reasoning power that shows us how beautifully these organs in some cases are adapted to certain end, declares in other cases are absolutely useless. Thus teeth in Rhinoceros[{165}], whale, narwhal,—bone on tibia, muscles which do not move,—little bone of wing of Apteryx,—bone representing extremities in some snake,—little wings within «?» soldered cover of beetles,—men and bulls, mammæ: filaments without anthers in plants, mere scales representing petals in others, in feather-hyacinth whole flower. Almost infinitely numerous. No one can reflect on these without astonishment, can anything be clearer than that wings are to fly and teeth «to bite», and yet we find these organs perfect in every detail in situations where they cannot possibly be of their normal use[{166}].

The term abortive organ has been thus applied to above structure (as invariable as all other parts[{167}]) from their absolute similarity to monstrous cases, where from accident, certain organs are not developed; as infant without arms or fingers with mere stump representing them: teeth represented by mere points of ossification: headless children with mere button,—viscera represented by small amorphous masses, &c.,—the tail by mere stump,—a solid horn by minute hanging one[{168}]. There is a tendency in all these cases, when life is preserved, for such structures to become hereditary. We see it in tailless dogs and cats. In plants we see this strikingly,—in Thyme, in Linum flavum,—stamen in Geranium pyrenaicum[{169}]. Nectaries abort into petals in Columbine «Aquilegia», produced from some accident and then become hereditary, in some cases only when propagated by buds, in other cases by seed. These cases have been produced suddenly by accident in early growth, but it is part of law of growth that when any organ is not used it tends to diminish (duck’s wing[{170}]?) muscles of dog’s ears, «and of» rabbits, muscles wither, arteries grow up. When eye born defective, optic nerve (Tuco Tuco) is atrophied. As every part whether useful or not (diseases, double flowers) tends to be transmitted to offspring, the origin of abortive organs whether produced at the birth or slowly acquired is easily understood in domestic races of organisms: [a struggle between the atrophy and hereditariness. Abortive organs in domestic races.] There will always be a struggle between atrophy of an organ rendered useless, and hereditariness[{171}]. Because we can understand the origin of abortive organs in certain cases, it would be wrong to conclude absolutely that all must have had same origin, but the strongest analogy is in favour of it. And we can by our theory, for during infinite changes some organ, we might have anticipated, would have become useless. «We can» readily explain the fact, so astounding on any other view, namely that organs possibly useless have been formed often with the same exquisite care as when of vital importance.

Our theory, I may remark would permit an organ «to» become abortive with respect to its primary use, to be turned to any other purpose, (as the buds in a cauliflower) thus we can see no difficulty in bones of male marsupials being used as fulcrum of muscles, or style of marygold[{172}],—indeed in one point of view, the heads of [vertebrated] animal may be said to be abortive vertebræ turned into other use: legs of some crustacea abortive jaws, &c., &c. De Candolle's analogy of table covered with dishes[{173}].

«The following passage was possibly intended to be inserted here.» Degradation and complication see Lamarck: no tendency to perfection: if room, [even] high organism would have greater power in beating lower one, thought «?» to be selected for a degraded end.

§ X. Recapitulation and conclusion.

Let us recapitulate the whole «?» «of» these latter sections by taking case of the three species of Rhinoceros, which inhabit Java, Sumatra, and mainland of Malacca or India. We find these three close neighbours, occupants of distinct but neighbouring districts, as a group having a different aspect from the Rhinoceros of Africa, though some of these latter inhabit very similar countries, but others most diverse stations. We find them intimately related [scarcely «?» differences more than some breeds of cattle] in structure to the Rhinoceros, which for immense periods have inhabited this one, out of three main zoological divisions of the world. Yet some of these ancient animals were fitted to very different stations: we find all three «illegible» of the generic character of the Rhinoceros, which form a [piece of net][{174}] set of links in the broken chain representing the Pachydermata, as the chain likewise forms a portion in other and longer chains. We see this wonderfully in dissecting the coarse leg of all three and finding nearly the same bones as in bat’s wings or man’s hand, but we see the clear mark in solid tibia of the fusion into it of the fibula. In all three we find their heads composed of three altered vertebræ, short neck, same bones as giraffe. In the upper jaws of all three we find small teeth like rabbit’s. In dissecting them in fœtal state we find at a not very early stage their form exactly alike the most different animals, and even with arteries running as in a fish: and this similarity holds when the young one is produced in womb, pond, egg or spawn. Now these three undoubted species scarcely differ more than breeds of cattle, are probably subject to many the same contagious diseases; if domesticated these forms would vary, and they might possibly breed together, and fuse into something[{175}] different «from» their aboriginal forms; might be selected to serve different ends.

Now the Creationist believes these three Rhinoceroses were created[{176}] with their deceptive appearance of true, not «illegible» relationship; as well can I believe the planets revolve in their present courses not from one law of gravity but from distinct volition of Creator.

If real species, sterile one with another, differently adapted, now inhabiting different countries, with different structures and instincts, are admitted to have common descent, we can only legitimately stop where our facts stop. Look how far in some case a chain of species will lead us. «This probably refers to the Crustacea, where the two ends of the series have “hardly a character in common.” Origin, Ed. i. p. 419.» May we not jump (considering how much extermination, and how imperfect geological records) from one sub-genus to another sub-genus. Can genera restrain us; many of the same arguments, which made us give up species, inexorably demand genera and families and orders to fall, and classes tottering. We ought to stop only when clear unity of type, independent of use and adaptation, ceases.

Be it remembered no naturalist pretends to give test from external characters of species; in many genera the distinction is quite arbitrary[{177}]. But there remains one other way of comparing species with races; it is to compare the effects of crossing them. Would it not be wonderful, if the union of two organisms, produced by two separate acts of Creation, blended their characters together when crossed according to the same rules, as two races which have undoubtedly descended from same parent stock; yet this can be shown to be the case. For sterility, though a usual «?», is not an invariable concomitant, it varies much in degree and has been shown to be probably dependent on causes closely analogous with those which make domesticated organisms sterile. Independent of sterility there is no difference between mongrels and hybrids, as can be shown in a long series of facts. It is strikingly seen in cases of instincts, when the minds of the two species or races become blended together[{178}]. In both cases if the half-breed be crossed with either parent for a few generations, all traces of the one parent form is lost (as Kölreuter in two tobacco species almost sterile together), so that the Creationist in the case of a species, must believe that one act of creation is absorbed into another!

Facsimile of the original manuscript of the paragraph on p. 50.

Conclusion.

Such are my reasons for believing that specific forms are not immutable. The affinity of different groups, the unity of types of structure, the representative forms through which fœtus passes, the metamorphosis of organs, the abortion of others cease to be metaphorical expressions and become intelligible facts. We no longer look «an» on animal as a savage does at a ship[{179}], or other great work of art, as a thing wholly beyond comprehension, but we feel far more interest in examining it. How interesting is every instinct, when we speculate on their origin as an hereditary or congenital habit or produced by the selection of individuals differing slightly from their parents. We must look at every complicated mechanism and instinct, as the summary of a long history, «as the summing up» of[{180}] useful contrivances, much like a work of art. How interesting does the distribution of all animals become, as throwing light on ancient geography. [We see some seas bridged over.] Geology loses in its glory from the imperfection of its archives[{181}], but how does it gain in the immensity of the periods of its formations and of the gaps separating these formations. There is much grandeur in looking at the existing animals either as the lineal descendants of the forms buried under thousand feet of matter, or as the coheirs of some still more ancient ancestor. It accords with what we know of the law impressed on matter by the Creator, that the creation and extinction of forms, like the birth and death of individuals should be the effect of secondary [laws] means[{182}]. It is derogatory that the Creator of countless systems of worlds should have created each of the myriads of creeping parasites and [slimy] worms which have swarmed each day of life on land and water «on» [this] one globe. We cease being astonished, however much we may deplore, that a group of animals should have been directly created to lay their eggs in bowels and flesh of other,—that some organisms should delight in cruelty,—that animals should be led away by false instincts,—that annually there should be an incalculable waste of eggs and pollen. From death, famine, rapine, and the concealed war of nature we can see that the highest good, which we can conceive, the creation of the higher animals has directly come. Doubtless it at first transcends our humble powers, to conceive laws capable of creating individual organisms, each characterised by the most exquisite workmanship and widely-extended adaptations. It accords better with [our modesty] the lowness of our faculties to suppose each must require the fiat of a creator, but in the same proportion the existence of such laws should exalt our notion of the power of the omniscient Creator[{183}]. There is a simple grandeur in the view of life with its powers of growth, assimilation and reproduction, being originally breathed into matter under one or a few forms, and that whilst this our planet has gone circling on according to fixed laws, and land and water, in a cycle of change, have gone on replacing each other, that from so simple an origin, through the process of gradual selection of infinitesimal changes, endless forms most beautiful and most wonderful have been evolved[{184}].

N.B.—There ought somewhere to be a discussion from Lyell to show that external conditions do vary, or a note to Lyell's works «work?».

Besides other difficulties in ii. Part, non-acclimatisation of plants. Difficulty when asked how did white and negro become altered from common intermediate stock: no facts. We do NOT know that species are immutable, on the contrary. What arguments against this theory, except our not perceiving every step, like the erosion of valleys[{185}].

THE ESSAY OF 1844

PART I

CHAPTER I
ON THE VARIATION OF ORGANIC BEINGS UNDER DOMESTICATION; AND ON THE PRINCIPLES OF SELECTION

The most favourable conditions for variation seem to be when organic beings are bred for many generations under domestication[{186}]: one may infer this from the simple fact of the vast number of races and breeds of almost every plant and animal, which has long been domesticated. Under certain conditions organic beings even during their individual lives become slightly altered from their usual form, size, or other characters: and many of the peculiarities thus acquired are transmitted to their offspring. Thus in animals, the size and vigour of body, fatness, period of maturity, habits of body or consensual movements, habits of mind and temper, are modified or acquired during the life of the individual[{187}], and become inherited. There is reason to believe that when long exercise has given to certain muscles great development, or disuse has lessened them, that such development is also inherited. Food and climate will occasionally produce changes in the colour and texture of the external coverings of animals; and certain unknown conditions affect the horns of cattle in parts of Abyssinia; but whether these peculiarities, thus acquired during individual lives, have been inherited, I do not know. It appears certain that malconformation and lameness in horses, produced by too much work on hard roads,—that affections of the eyes in this animal probably caused by bad ventilation,—that tendencies towards many diseases in man, such as gout, caused by the course of life and ultimately producing changes of structure, and that many other diseases produced by unknown agencies, such as goitre, and the idiotcy resulting from it, all become hereditary.

It is very doubtful whether the flowers and leaf-buds, annually produced from the same bulb, root, or tree, can properly be considered as parts of the same individual, though in some respects they certainly seem to be so. If they are parts of an individual, plants also are subject to considerable changes during their individual lives. Most florist-flowers if neglected degenerate, that is, they lose some of their characters; so common is this, that trueness is often stated, as greatly enhancing the value of a variety[{188}]: tulips break their colours only after some years’ culture; some plants become double and others single, by neglect or care: these characters can be transmitted by cuttings or grafts, and in some cases by true or seminal propagation. Occasionally a single bud on a plant assumes at once a new and widely different character: thus it is certain that nectarines have been produced on peach trees and moss roses on provence roses; white currants on red currant bushes; flowers of a different colour from that of the stock, in Chrysanthemums, Dahlias, sweet-williams, Azaleas, &c., &c.; variegated leaf-buds on many trees, and other similar cases. These new characters appearing in single buds, can, like those lesser changes affecting the whole plant, be multiplied not only by cuttings and such means, but often likewise by true seminal generation.

The changes thus appearing during the lives of individual animals and plants are extremely rare compared with those which are congenital or which appear soon after birth. Slight differences thus arising are infinitely numerous: the proportions and form of every part of the frame, inside and outside, appear to vary in very slight degrees: anatomists dispute what is the “beau ideal” of the bones, the liver and kidneys, like painters do of the proportions of the face: the proverbial expression that no two animals or plants are born absolutely alike, is much truer when applied to those under domestication, than to those in a state of nature[{189}]. Besides these slight differences, single individuals are occasionally born considerably unlike in certain parts or in their whole structure to their parents: these are called by horticulturists and breeders “sports”; and are not uncommon except when very strongly marked. Such sports are known in some cases to have been parents of some of our domestic races; and such probably have been the parents of many other races, especially of those which in some senses may be called hereditary monsters; for instance where there is an additional limb, or where all the limbs are stunted (as in the Ancon sheep), or where a part is wanting, as in rumpless fowls and tailless dogs or cats[{190}]. The effects of external conditions on the size, colour and form, which can rarely and obscurely be detected during one individual life, become apparent after several generations: the slight differences, often hardly describable, which characterize the stock of different countries, and even of districts in the same country, seem to be due to such continued action.

On the hereditary tendency.

A volume might be filled with facts showing what a strong tendency there is to inheritance, in almost every case of the most trifling, as well as of the most remarkable congenital peculiarities[{191}]. The term congenital peculiarity, I may remark, is a loose expression and can only mean a peculiarity apparent when the part affected is nearly or fully developed: in the Second Part, I shall have to discuss at what period of the embryonic life connatal peculiarities probably first appear; and I shall then be able to show from some evidence, that at whatever period of life a new peculiarity first appears, it tends hereditarily to appear at a corresponding period[{192}]. Numerous though slight changes, slowly supervening in animals during mature life (often, though by no means always, taking the form of disease), are, as stated in the first paragraphs, very often hereditary. In plants, again, the buds which assume a different character from their stock likewise tend to transmit their new peculiarities. There is not sufficient reason to believe that either mutilations[{193}] or changes of form produced by mechanical pressure, even if continued for hundreds of generations, or that any changes of structure quickly produced by disease, are inherited; it would appear as if the tissue of the part affected must slowly and freely grow into the new form, in order to be inheritable. There is a very great difference in the hereditary tendency of different peculiarities, and of the same peculiarity, in different individuals and species; thus twenty thousand seeds of the weeping ash have been sown and not one come up true;—out of seventeen seeds of the weeping yew, nearly all came up true. The ill-formed and almost monstrous “Niata” cattle of S. America and Ancon sheep, both when bred together and when crossed with other breeds, seem to transmit their peculiarities to their offspring as truly as the ordinary breeds. I can throw no light on these differences in the power of hereditary transmission. Breeders believe, and apparently with good cause, that a peculiarity generally becomes more firmly implanted after having passed through several generations; that is if one offspring out of twenty inherits a peculiarity from its parents, then its descendants will tend to transmit this peculiarity to a larger proportion than one in twenty; and so on in succeeding generations. I have said nothing about mental peculiarities being inheritable for I reserve this subject for a separate chapter.

Causes of Variation.

Attention must here be drawn to an important distinction in the first origin or appearance of varieties: when we see an animal highly kept producing offspring with an hereditary tendency to early maturity and fatness; when we see the wild-duck and Australian dog always becoming, when bred for one or a few generations in confinement, mottled in their colours; when we see people living in certain districts or circumstances becoming subject to an hereditary taint to certain organic diseases, as consumption or plica polonica,—we naturally attribute such changes to the direct effect of known or unknown agencies acting for one or more generations on the parents. It is probable that a multitude of peculiarities may be thus directly caused by unknown external agencies. But in breeds, characterized by an extra limb or claw, as in certain fowls and dogs; by an extra joint in the vertebræ; by the loss of a part, as the tail; by the substitution of a tuft of feathers for a comb in certain poultry; and in a multitude of other cases, we can hardly attribute these peculiarities directly to external influences, but indirectly to the laws of embryonic growth and of reproduction. When we see a multitude of varieties (as has often been the case, where a cross has been carefully guarded against) produced from seeds matured in the very same capsule[{194}], with the male and female principle nourished from the same roots and necessarily exposed to the same external influences; we cannot believe that the endless slight differences between seedling varieties thus produced, can be the effect of any corresponding difference in their exposure. We are led (as Müller has remarked) to the same conclusion, when we see in the same litter, produced by the same act of conception, animals considerably different.

As variation to the degree here alluded to has been observed only in organic beings under domestication, and in plants amongst those most highly and long cultivated, we must attribute, in such cases, the varieties (although the difference between each variety cannot possibly be attributed to any corresponding difference of exposure in the parents) to the indirect effects of domestication on the action of the reproductive system[{195}]. It would appear as if the reproductive powers failed in their ordinary function of producing new organic beings closely like their parents; and as if the entire organization of the embryo, under domestication, became in a slight degree plastic[{196}]. We shall hereafter have occasion to show, that in organic beings, a considerable change from the natural conditions of life, affects, independently of their general state of health, in another and remarkable manner the reproductive system. I may add, judging from the vast number of new varieties of plants which have been produced in the same districts and under nearly the same routine of culture, that probably the indirect effects of domestication in making the organization plastic, is a much more efficient source of variation than any direct effect which external causes may have on the colour, texture, or form of each part. In the few instances in which, as in the Dahlia[{197}], the course of variation has been recorded, it appears that domestication produces little effect for several generations in rendering the organization plastic; but afterwards, as if by an accumulated effect, the original character of the species suddenly gives way or breaks.

On Selection.

We have hitherto only referred to the first appearance in individuals of new peculiarities; but to make a race or breed, something more is generally[{198}] requisite than such peculiarities (except in the case of the peculiarities being the direct effect of constantly surrounding conditions) should be inheritable,—namely the principle of selection, implying separation. Even in the rare instances of sports, with the hereditary tendency very strongly implanted, crossing must be prevented with other breeds, or if not prevented the best characterized of the half-bred offspring must be carefully selected. Where the external conditions are constantly tending to give some character, a race possessing this character will be formed with far greater ease by selecting and breeding together the individuals most affected. In the case of the endless slight variations produced by the indirect effects of domestication on the action of the reproductive system, selection is indispensable to form races; and when carefully applied, wonderfully numerous and diverse races can be formed. Selection, though so simple in theory, is and has been important to a degree which can hardly be overrated. It requires extreme skill, the results of long practice, in detecting the slightest difference in the forms of animals, and it implies some distinct object in view; with these requisites and patience, the breeder has simply to watch for every the smallest approach to the desired end, to select such individuals and pair them with the most suitable forms, and so continue with succeeding generations. In most cases careful selection and the prevention of accidental crosses will be necessary for several generations, for in new breeds there is a strong tendency to vary and especially to revert to ancestral forms: but in every succeeding generation less care will be requisite for the breed will become truer; until ultimately only an occasional individual will require to be separated or destroyed. Horticulturalists in raising seeds regularly practise this, and call it “roguing,” or destroying the “rogues” or false varieties. There is another and less efficient means of selection amongst animals: namely repeatedly procuring males with some desirable qualities, and allowing them and their offspring to breed freely together; and this in the course of time will affect the whole lot. These principles of selection have been methodically followed for scarcely a century; but their high importance is shown by the practical results, and is admitted in the writings of the most celebrated agriculturalists and horticulturalists;—I need only name Anderson, Marshall, Bakewell, Coke, Western, Sebright and Knight.

Even in well-established breeds the individuals of which to an unpractised eye would appear absolutely similar, which would give, it might have been thought, no scope to selection, the whole appearance of the animal has been changed in a few years (as in the case of Lord Western’s sheep), so that practised agriculturalists could scarcely credit that a change had not been effected by a cross with other breeds. Breeders both of plants and animals frequently give their means of selection greater scope, by crossing different breeds and selecting the offspring; but we shall have to recur to this subject again.

The external conditions will doubtless influence and modify the results of the most careful selection; it has been found impossible to prevent certain breeds of cattle from degenerating on mountain pastures; it would probably be impossible to keep the plumage of the wild-duck in the domesticated race; in certain soils, no care has been sufficient to raise cauliflower seed true to its character; and so in many other cases. But with patience it is wonderful what man has effected. He has selected and therefore in one sense made one breed of horses to race and another to pull; he has made sheep with fleeces good for carpets and other sheep good for broadcloth; he has, in the same sense, made one dog to find game and give him notice when found, and another dog to fetch him the game when killed; he has made by selection the fat to lie mixed with the meat in one breed and in another to accumulate in the bowels for the tallow-chandler[{199}]; he has made the legs of one breed of pigeons long, and the beak of another so short, that it can hardly feed itself; he has previously determined how the feathers on a bird’s body shall be coloured, and how the petals of many flowers shall be streaked or fringed, and has given prizes for complete success;—by selection, he has made the leaves of one variety and the flower-buds of another variety of the cabbage good to eat, at different seasons of the year; and thus has he acted on endless varieties. I do not wish to affirm that the long-and short-wooled sheep, or that the pointer and retriever, or that the cabbage and cauliflower have certainly descended from one and the same aboriginal wild stock; if they have not so descended, though it lessens what man has effected, a large result must be left unquestioned.

In saying as I have done that man makes a breed, let it not be confounded with saying that man makes the individuals, which are given by nature with certain desirable qualities; man only adds together and makes a permanent gift of nature’s bounties. In several cases, indeed, for instance in the “Ancon” sheep, valuable from not getting over fences, and in the turnspit dog, man has probably only prevented crossing; but in many cases we positively know that he has gone on selecting, and taking advantage of successive small variations.

Selection[{200}] has been methodically followed, as I have said, for barely a century; but it cannot be doubted that occasionally it has been practised from the remotest ages, in those animals completely under the dominion of man. In the earliest chapters of the Bible there are rules given for influencing the colours of breeds, and black and white sheep are spoken of as separated. In the time of Pliny the barbarians of Europe and Asia endeavoured by cross-breeding with a wild stock to improve the races of their dogs and horses. The savages of Guyana now do so with their dogs: such care shows at least that the characters of individual animals were attended to. In the rudest times of English history, there were laws to prevent the exportation of fine animals of established breeds, and in the case of horses, in Henry VIII’s time, laws for the destruction of all horses under a certain size. In one of the oldest numbers of the Phil. Transactions, there are rules for selecting and improving the breeds of sheep. Sir H. Bunbury, in 1660, has given rules for selecting the finest seedling plants, with as much precision as the best recent horticulturalist could. Even in the most savage and rude nations, in the wars and famines which so frequently occur, the most useful of their animals would be preserved: the value set upon animals by savages is shown by the inhabitants of Tierra del Fuego devouring their old women before their dogs, which as they asserted are useful in otter-hunting[{201}]: who can doubt but that in every case of famine and war, the best otter-hunters would be preserved, and therefore in fact selected for breeding. As the offspring so obviously take after their parents, and as we have seen that savages take pains in crossing their dogs and horses with wild stocks, we may even conclude as probable that they would sometimes pair the most useful of their animals and keep their offspring separate. As different races of men require and admire different qualities in their domesticated animals, each would thus slowly, though unconsciously, be selecting a different breed. As Pallas has remarked, who can doubt but that the ancient Russian would esteem and endeavour to preserve those sheep in his flocks which had the thickest coats. This kind of insensible selection by which new breeds are not selected and kept separate, but a peculiar character is slowly given to the whole mass of the breed, by often saving the life of animals with certain characteristics, we may feel nearly sure, from what we see has been done by the more direct method of separate selection within the last 50 years in England, would in the course of some thousand years produce a marked effect.

Crossing Breeds.

When once two or more races are formed, or if more than one race, or species fertile inter se, originally existed in a wild state, their crossing becomes a most copious source of new races[{202}]. When two well-marked races are crossed the offspring in the first generation take more or less after either parent or are quite intermediate between them, or rarely assume characters in some degree new. In the second and several succeeding generations, the offspring are generally found to vary exceedingly, one compared with another, and many revert nearly to their ancestral forms. This greater variability in succeeding generations seems analogous to the breaking or variability of organic beings after having been bred for some generations under domestication[{203}]. So marked is this variability in cross-bred descendants, that Pallas and some other naturalists have supposed that all variation is due to an original cross; but I conceive that the history of the potato, Dahlia, Scotch Rose, the guinea-pig, and of many trees in this country, where only one species of the genus exists, clearly shows that a species may vary where there can have been no crossing. Owing to this variability and tendency to reversion in cross-bred beings, much careful selection is requisite to make intermediate or new permanent races: nevertheless crossing has been a most powerful engine, especially with plants, where means of propagation exist by which the cross-bred varieties can be secured without incurring the risk of fresh variation from seminal propagation: with animals the most skilful agriculturalists now greatly prefer careful selection from a well-established breed, rather than from uncertain cross-bred stocks.

Although intermediate and new races may be formed by the mingling of others, yet if the two races are allowed to mingle quite freely, so that none of either parent race remain pure, then, especially if the parent races are not widely different, they will slowly blend together, and the two races will be destroyed, and one mongrel race left in its place. This will of course happen in a shorter time, if one of the parent races exists in greater number than the other. We see the effect of this mingling, in the manner in which the aboriginal breeds of dogs and pigs in the Oceanic Islands and the many breeds of our domestic animals introduced into S. America, have all been lost and absorbed in a mongrel race. It is probably owing to the freedom of crossing, that, in uncivilised countries, where inclosures do not exist, we seldom meet with more than one race of a species: it is only in enclosed countries, where the inhabitants do not migrate, and have conveniences for separating the several kinds of domestic animals, that we meet with a multitude of races. Even in civilised countries, want of care for a few years has been found to destroy the good results of far longer periods of selection and separation.

This power of crossing will affect the races of all terrestrial animals; for all terrestrial animals require for their reproduction the union of two individuals. Amongst plants, races will not cross and blend together with so much freedom as in terrestrial animals; but this crossing takes place through various curious contrivances to a surprising extent. In fact such contrivances exist in so very many hermaphrodite flowers by which an occasional cross may take place, that I cannot avoid suspecting (with Mr Knight) that the reproductive action requires, at intervals, the concurrence of distinct individuals[{204}]. Most breeders of plants and animals are firmly convinced that benefit is derived from an occasional cross, not with another race, but with another family of the same race; and that, on the other hand, injurious consequences follow from long-continued close interbreeding in the same family. Of marine animals, many more, than was till lately believed, have their sexes on separate individuals; and where they are hermaphrodite, there seems very generally to be means through the water of one individual occasionally impregnating another: if individual animals can singly propagate themselves for perpetuity, it is unaccountable that no terrestrial animal, where the means of observation are more obvious, should be in this predicament of singly perpetuating its kind. I conclude, then, that races of most animals and plants, when unconfined in the same country, would tend to blend together.

Whether our domestic races have descended from one or more wild stocks.

Several naturalists, of whom Pallas[{205}] regarding animals, and Humboldt regarding certain plants, were the first, believe that the breeds of many of our domestic animals such as of the horse, pig, dog, sheep, pigeon, and poultry, and of our plants have descended from more than one aboriginal form. They leave it doubtful, whether such forms are to be considered wild races, or true species, whose offspring are fertile when crossed inter se. The main arguments for this view consist, firstly, of the great difference between such breeds, as the Race-and Cart-Horse, or the Greyhound and Bull-dog, and of our ignorance of the steps or stages through which these could have passed from a common parent; and secondly that in the most ancient historical periods, breeds resembling some of those at present most different, existed in different countries. The wolves of N. America and of Siberia are thought to be different species; and it has been remarked that the dogs belonging to the savages in these two countries resemble the wolves of the same country; and therefore that they have probably descended from two different wild stocks. In the same manner, these naturalists believe that the horse of Arabia and of Europe have probably descended from two wild stocks both apparently now extinct. I do not think the assumed fertility of these wild stocks any very great difficulty on this view; for although in animals the offspring of most cross-bred species are infertile, it is not always remembered that the experiment is very seldom fairly tried, except when two near species both breed freely (which does not readily happen, as we shall hereafter see) when under the dominion of man. Moreover in the case of the China[{206}] and common goose, the canary and siskin, the hybrids breed freely; in other cases the offspring from hybrids crossed with either pure parent are fertile, as is practically taken advantage of with the yak and cow; as far as the analogy of plants serves, it is impossible to deny that some species are quite fertile inter se; but to this subject we shall recur.

On the other hand, the upholders of the view that the several breeds of dogs, horses, &c., &c., have descended each from one stock, may aver that their view removes all difficulty about fertility, and that the main argument from the high antiquity of different breeds, somewhat similar to the present breeds, is worth little without knowing the date of the domestication of such animals, which is far from being the case. They may also with more weight aver that, knowing that organic beings under domestication do vary in some degree, the argument from the great difference between certain breeds is worth nothing, without we know the limits of variation during a long course of time, which is far from the case. They may argue that almost every county in England, and in many districts of other countries, for instance in India, there are slightly different breeds of the domestic animals; and that it is opposed to all that we know of the distribution of wild animals to suppose that these have descended from so many different wild races or species: if so, they may argue, is it not probable that countries quite separate and exposed to different climates would have breeds not slightly, but considerably, different? Taking the most favourable case, on both sides, namely that of the dog; they might urge that such breeds as the bull-dog and turnspit have been reared by man, from the ascertained fact that strictly analogous breeds (namely the Niata ox and Ancon sheep) in other quadrupeds have thus originated. Again they may say, seeing what training and careful selection has effected for the greyhound, and seeing how absolutely unfit the Italian greyhound is to maintain itself in a state of nature, is it not probable that at least all greyhounds,—from the rough deerhound, the smooth Persian, the common English, to the Italian,—have descended from one stock[{207}]? If so, is it so improbable that the deerhound and long-legged shepherd dog have so descended? If we admit this, and give up the bull-dog, we can hardly dispute the probable common descent of the other breeds.

The evidence is so conjectural and balanced on both sides that at present I conceive that no one can decide: for my own part, I lean to the probability of most of our domestic animals having descended from more than one wild stock; though from the arguments last advanced and from reflecting on the slow though inevitable effect of different races of mankind, under different circumstances, saving the lives of and therefore selecting the individuals most useful to them, I cannot doubt but that one class of naturalists have much overrated the probable number of the aboriginal wild stocks. As far as we admit the difference of our races «to be» due to the differences of their original stocks, so much must we give up of the amount of variation produced under domestication. But this appears to me unimportant, for we certainly know in some few cases, for instance in the Dahlia, and potato, and rabbit, that a great number of varieties have proceeded from one stock; and, in many of our domestic races, we know that man, by slowly selecting and by taking advantage of sudden sports, has considerably modified old races and produced new ones. Whether we consider our races as the descendants of one or several wild stocks, we are in far the greater number of cases equally ignorant what these stocks were.

Limits to Variation in degree and kind.

Man’s power in making races deends, in the first instance, on the stock on which he works being variable; but his labours are modified and limited, as we have seen, by the direct effects of the external conditions,—by the deficient or imperfect hereditariness of new peculiarities,—and by the tendency to continual variation and especially to reversion to ancestral forms. If the stock is not variable under domestication, of course he can do nothing; and it appears that species differ considerably in this tendency to variation, in the same way as even sub-varieties from the same variety differ greatly in this respect, and transmit to their offspring this difference in tendency. Whether the absence of a tendency to vary is an unalterable quality in certain species, or depends on some deficient condition of the particular state of domestication to which they are exposed, there is no evidence. When the organization is rendered variable, or plastic, as I have expressed it, under domestication, different parts of the frame vary more or less in different species: thus in the breeds of cattle it has been remarked that the horns are the most constant or least variable character, for these often remain constant, whilst the colour, size, proportions of the body, tendency to fatten &c., vary; in sheep, I believe, the horns are much more variable. As a general rule the less important parts of the organization seem to vary most, but I think there is sufficient evidence that every part occasionally varies in a slight degree. Even when man has the primary requisite variability he is necessarily checked by the health and life of the stock he is working on: thus he has already made pigeons with such small beaks that they can hardly eat and will not rear their own young; he has made families of sheep with so strong a tendency to early maturity and to fatten, that in certain pastures they cannot live from their extreme liability to inflammation; he has made (i.e. selected) sub-varieties of plants with a tendency to such early growth that they are frequently killed by the spring frosts; he has made a breed of cows having calves with such large hinder quarters that they are born with great difficulty, often to the death of their mothers[{208}]; the breeders were compelled to remedy this by the selection of a breeding stock with smaller hinder quarters; in such a case, however, it is possible by long patience and great loss, a remedy might have been found in selecting cows capable of giving birth to calves with large hinder quarters, for in human kind there «are» no doubt hereditary bad and good confinements. Besides the limits already specified, there can be little doubt that the variation of different parts of the frame are connected together by many laws[{209}]: thus the two sides of the body, in health and disease, seem almost always to vary together: it has been asserted by breeders that if the head is much elongated, the bones of the extremities will likewise be so; in seedling-apples large leaves and fruit generally go together, and serve the horticulturalist as some guide in his selection; we can here see the reason, as the fruit is only a metamorphosed leaf. In animals the teeth and hair seem connected, for the hairless Chinese dog is almost toothless. Breeders believe that one part of the frame or function being increased causes other parts to decrease: they dislike great horns and great bones as so much flesh lost; in hornless breeds of cattle certain bones of the head become more developed: it is said that fat accumulating in one part checks its accumulation in another, and likewise checks the action of the udder. The whole organization is so connected that it is probable there are many conditions determining the variation of each part, and causing other parts to vary with it; and man in making new races must be limited and ruled by all such laws.

In what consists Domestication.

In this chapter we have treated of variation under domestication, and it now remains to consider in what does this power of domestication consist[{210}], a subject of considerable difficulty. Observing that organic beings of almost every class, in all climates, countries, and times, have varied when long bred under domestication, we must conclude that the influence is of some very general nature[{211}]. Mr Knight alone, as far as I know, has tried to define it; he believes it consists of an excess of food, together with transport to a more genial climate, or protection from its severities. I think we cannot admit this latter proposition, for we know how many vegetable products, aborigines of this country, here vary, when cultivated without any protection from the weather; and some of our variable trees, as apricots, peaches, have undoubtedly been derived from a more genial climate. There appears to be much more truth in the doctrine of excess of food being the cause, though I much doubt whether this is the sole cause, although it may well be requisite for the kind of variation desired by man, namely increase of size and vigour. No doubt horticulturalists, when they wish to raise new seedlings, often pluck off all the flower-buds, except a few, or remove the whole during one season, so that a great stock of nutriment may be thrown into the flowers which are to seed. When plants are transported from high-lands, forests, marshes, heaths, into our gardens and greenhouses, there must be a considerable change of food, but it would be hard to prove that there was in every case an excess of the kind proper to the plant. If it be an excess of food, compared with that which the being obtained in its natural state[{212}], the effects continue for an improbably long time; during how many ages has wheat been cultivated, and cattle and sheep reclaimed, and we cannot suppose their amount of food has gone on increasing, nevertheless these are amongst the most variable of our domestic productions. It has been remarked (Marshall) that some of the most highly kept breeds of sheep and cattle are truer or less variable than the straggling animals of the poor, which subsist on commons, and pick up a bare subsistence[{213}]. In the case of forest-trees raised in nurseries, which vary more than the same trees do in their aboriginal forests, the cause would seem simply to lie in their not having to struggle against other trees and weeds, which in their natural state doubtless would limit the conditions of their existence. It appears to me that the power of domestication resolves itself into the accumulated effects of a change of all or some of the natural conditions of the life of the species, often associated with excess of food. These conditions moreover, I may add, can seldom remain, owing to the mutability of the affairs, habits, migrations, and knowledge of man, for very long periods the same. I am the more inclined to come to this conclusion from finding, as we shall hereafter show, that changes of the natural conditions of existence seem peculiarly to affect the action of the reproductive system[{214}]. As we see that hybrids and mongrels, after the first generation, are apt to vary much, we may at least conclude that variability does not altogether depend on excess of food.

After these views, it may be asked how it comes that certain animals and plants, which have been domesticated for a considerable length of time, and transported from very different conditions of existence, have not varied much, or scarcely at all; for instance, the ass, peacock, guinea-fowl, asparagus, Jerusalem artichoke[{215}]. I have already said that probably different species, like different sub-varieties, possess different degrees of tendency to vary; but I am inclined to attribute in these cases the want of numerous races less to want of variability than to selection not having been practised on them. No one will take the pains to select without some corresponding object, either of use or amusement; the individuals raised must be tolerably numerous, and not so precious, but that he may freely destroy those not answering to his wishes. If guinea-fowls or peacocks[{216}] became “fancy” birds, I cannot doubt that after some generations several breeds would be raised. Asses have not been worked on from mere neglect; but they differ in some degree in different countries. The insensible selection, due to different races of mankind preserving those individuals most useful to them in their different circumstances, will apply only to the oldest and most widely domesticated animals. In the case of plants, we must put entirely out of the case those exclusively (or almost so) propagated by cuttings, layers or tubers, such as the Jerusalem artichoke and laurel; and if we put on one side plants of little ornament or use, and those which are used at so early a period of their growth that no especial characters signify, as asparagus[{217}] and seakale, I can think of none long cultivated which have not varied. In no case ought we to expect to find as much variation in a race when it alone has been formed, as when several have been formed, for their crossing and recrossing will greatly increase their variability.

Summary of first Chapter.

To sum up this chapter. Races are made under domestication: 1st, by the direct effects of the external conditions to which the species is exposed: 2nd, by the indirect effects of the exposure to new conditions, often aided by excess of food, rendering the organization plastic, and by man’s selecting and separately breeding certain individuals, or introducing to his stock selected males, or often preserving with care the life of the individuals best adapted to his purposes: 3rd, by crossing and recrossing races already made, and selecting their offspring. After some generations man may relax his care in selection: for the tendency to vary and to revert to ancestral forms will decrease, so that he will have only occasionally to remove or destroy one of the yearly offspring which departs from its type. Ultimately, with a large stock, the effects of free crossing would keep, even without this care, his breed true. By these means man can produce infinitely numerous races, curiously adapted to ends, both most important and most frivolous; at the same time that the effects of the surrounding conditions, the laws of inheritance, of growth, and of variation, will modify and limit his labours.

CHAPTER II
ON THE VARIATION OF ORGANIC BEINGS IN A WILD STATE; ON THE NATURAL MEANS OF SELECTION; AND ON THE COMPARISON OF DOMESTIC RACES AND TRUE SPECIES

Having treated of variation under domestication, we now come to it in a state of nature.

Most organic beings in a state of nature vary exceedingly little[{218}]: I put out of the case variations (as stunted plants &c., and sea-shells in brackish water[{219}]) which are directly the effect of external agencies and which we do not know are in the breed[{220}], or are hereditary. The amount of hereditary variation is very difficult to ascertain, because naturalists (partly from the want of knowledge, and partly from the inherent difficulty of the subject) do not all agree whether certain forms are species or races[{221}]. Some strongly marked races of plants, comparable with the decided sports of horticulturalists, undoubtedly exist in a state of nature, as is actually known by experiment, for instance in the primrose and cowslip[{222}], in two so-called species of dandelion, in two of foxglove[{223}], and I believe in some pines. Lamarck has observed that, as long as we confine our attention to one limited country, there is seldom much difficulty in deciding what forms to call species and what varieties; and that it is when collections flow in from all parts of the world that naturalists often feel at a loss to decide the limit of variation. Undoubtedly so it is, yet amongst British plants (and I may add land shells), which are probably better known than any in the world, the best naturalists differ very greatly in the relative proportions of what they call species and what varieties. In many genera of insects, and shells, and plants, it seems almost hopeless to establish which are which. In the higher classes there are less doubts; though we find considerable difficulty in ascertaining what deserve to be called species amongst foxes and wolves, and in some birds, for instance in the case of the white barn-owl. When specimens are brought from different parts of the world, how often do naturalists dispute this same question, as I found with respect to the birds brought from the Galapagos islands. Yarrell has remarked that the individuals of the same undoubted species of birds, from Europe and N. America, usually present slight, indefinable though perceptible differences. The recognition indeed of one animal by another of its kind seems to imply some difference. The disposition of wild animals undoubtedly differs. The variation, such as it is, chiefly affects the same parts in wild organisms as in domestic breeds; for instance, the size, colour, and the external and less important parts. In many species the variability of certain organs or qualities is even stated as one of the specific characters: thus, in plants, colour, size, hairiness, the number of the stamens and pistils, and even their presence, the form of the leaves; the size and form of the mandibles of the males of some insects; the length and curvature of the beak in some birds (as in Opetiorynchus) are variable characters in some species and quite fixed in others. I do not perceive that any just distinction can be drawn between this recognised variability of certain parts in many species and the more general variability of the whole frame in domestic races.

Although the amount of variation be exceedingly small in most organic beings in a state of nature, and probably quite wanting (as far as our senses serve) in the majority of cases; yet considering how many animals and plants, taken by mankind from different quarters of the world for the most diverse purposes, have varied under domestication in every country and in every age, I think we may safely conclude that all organic beings with few exceptions, if capable of being domesticated and bred for long periods, would vary. Domestication seems to resolve itself into a change from the natural conditions of the species [generally perhaps including an increase of food]; if this be so, organisms in a state of nature must occasionally, in the course of ages, be exposed to analogous influences; for geology clearly shows that many places must, in the course of time, become exposed to the widest range of climatic and other influences; and if such places be isolated, so that new and better adapted organic beings cannot freely emigrate, the old inhabitants will be exposed to new influences, probably far more varied, than man applies under the form of domestication. Although every species no doubt will soon breed up to the full number which the country will support, yet it is easy to conceive that, on an average, some species may receive an increase of food; for the times of dearth may be short, yet enough to kill, and recurrent only at long intervals. All such changes of conditions from geological causes would be exceedingly slow; what effect the slowness might have we are ignorant; under domestication it appears that the effects of change of conditions accumulate, and then break out. Whatever might be the result of these slow geological changes, we may feel sure, from the means of dissemination common in a lesser or greater degree to every organism taken conjointly with the changes of geology, which are steadily (and sometimes suddenly, as when an isthmus at last separates) in progress, that occasionally organisms must suddenly be introduced into new regions, where, if the conditions of existence are not so foreign as to cause its extermination, it will often be propagated under circumstances still more closely analogous to those of domestication; and therefore we expect will evince a tendency to vary. It appears to me quite inexplicable if this has never happened; but it can happen very rarely. Let us then suppose that an organism by some chance (which might be hardly repeated in 1000 years) arrives at a modern volcanic island in process of formation and not fully stocked with the most appropriate organisms; the new organism might readily gain a footing, although the external conditions were considerably different from its native ones. The effect of this we might expect would influence in some small degree the size, colour, nature of covering &c., and from inexplicable influences even special parts and organs of the body. But we might further (and «this» is far more important) expect that the reproductive system would be affected, as under domesticity, and the structure of the offspring rendered in some degree plastic. Hence almost every part of the body would tend to vary from the typical form in slight degrees, and in no determinate way, and therefore without selection the free crossing of these small variations (together with the tendency to reversion to the original form) would constantly be counteracting this unsettling effect of the extraneous conditions on the reproductive system. Such, I conceive, would be the unimportant result without selection. And here I must observe that the foregoing remarks are equally applicable to that small and admitted amount of variation which has been observed in some organisms in a state of nature; as well as to the above hypothetical variation consequent on changes of condition.

Let us now suppose a Being[{224}] with penetration sufficient to perceive differences in the outer and innermost organization quite imperceptible to man, and with forethought extending over future centuries to watch with unerring care and select for any object the offspring of an organism produced under the foregoing circumstances; I can see no conceivable reason why he could not form a new race (or several were he to separate the stock of the original organism and work on several islands) adapted to new ends. As we assume his discrimination, and his forethought, and his steadiness of object, to be incomparably greater that those qualities in man, so we may suppose the beauty and complications of the adaptations of the new races and their differences from the original stock to be greater than in the domestic races produced by man’s agency: the ground-work of his labours we may aid by supposing that the external conditions of the volcanic island, from its continued emergence and the occasional introduction of new immigrants, vary; and thus to act on the reproductive system of the organism, on which he is at work, and so keep its organization somewhat plastic. With time enough, such a Being might rationally (without some unknown law opposed him) aim at almost any result.

For instance, let this imaginary Being wish, from seeing a plant growing on the decaying matter in a forest and choked by other plants, to give it power of growing on the rotten stems of trees, he would commence selecting every seedling whose berries were in the smallest degree more attractive to tree-frequenting birds, so as to cause a proper dissemination of the seeds, and at the same time he would select those plants which had in the slightest degree more and more power of drawing nutriment from rotten wood; and he would destroy all other seedlings with less of this power. He might thus, in the course of century after century, hope to make the plant by degrees grow on rotten wood, even high up on trees, wherever birds dropped the non-digested seeds. He might then, if the organization of the plant was plastic, attempt by continued selection of chance seedlings to make it grow on less and less rotten wood, till it would grow on sound wood[{225}]. Supposing again, during these changes the plant failed to seed quite freely from non-impregnation, he might begin selecting seedlings with a little sweeter «or» differently tasted honey or pollen, to tempt insects to visit the flowers regularly: having effected this, he might wish, if it profited the plant, to render abortive the stamens and pistils in different flowers, which he could do by continued selection. By such steps he might aim at making a plant as wonderfully related to other organic beings as is the mistletoe, whose existence absolutely depends on certain insects for impregnation, certain birds for transportal, and certain trees for growth. Furthermore, if the insect which had been induced regularly to visit this hypothetical plant profited much by it, our same Being might wish by selection to modify by gradual selection the insect’s structure, so as to facilitate its obtaining the honey or pollen: in this manner he might adapt the insect (always presupposing its organization to be in some degree plastic) to the flower, and the impregnation of the flower to the insect; as is the case with many bees and many plants.

Seeing what blind capricious man has actually effected by selection during the few last years, and what in a ruder state he has probably effected without any systematic plan during the last few thousand years, he will be a bold person who will positively put limits to what the supposed Being could effect during whole geological periods. In accordance with the plan by which this universe seems governed by the Creator, let us consider whether there exists any secondary means in the economy of nature by which the process of selection could go on adapting, nicely and wonderfully, organisms, if in ever so small a degree plastic, to diverse ends. I believe such secondary means do exist[{226}].

Natural means of Selection[{227}].

De Candolle, in an eloquent passage, has declared that all nature is at war, one organism with another, or with external nature. Seeing the contented face of nature, this may at first be well doubted; but reflection will inevitably prove it is too true. The war, however, is not constant, but only recurrent in a slight degree at short periods and more severely at occasional more distant periods; and hence its effects are easily overlooked. It is the doctrine of Malthus applied in most cases with ten-fold force. As in every climate there are seasons for each of its inhabitants of greater and less abundance, so all annually breed; and the moral restraint, which in some small degree checks the increase of mankind, is entirely lost. Even slow-breeding mankind has doubled in 25 years[{228}], and if he could increase his food with greater ease, he would double in less time. But for animals, without artificial means, on an average the amount of food for each species must be constant; whereas the increase of all organisms tends to be geometrical, and in a vast majority of cases at an enormous ratio. Suppose in a certain spot there are eight pairs of [robins] birds, and that only four pairs of them annually (including double hatches) rear only four young; and that these go on rearing their young at the same rate: then at the end of seven years (a short life, excluding violent deaths, for any birds) there will be 2048 robins, instead of the original sixteen; as this increase is quite impossible, so we must conclude either that robins do not rear nearly half their young or that the average life of a robin when reared is from accident not nearly seven years. Both checks probably concur. The same kind of calculation applied to all vegetables and animals produces results either more or less striking, but in scarcely a single instance less striking than in man[{229}].

Many practical illustrations of this rapid tendency to increase are on record, namely during peculiar seasons, in the extraordinary increase of certain animals, for instance during the years 1826 to 1828, in La Plata, when from drought, some millions of cattle perished, the whole country swarmed with innumerable mice: now I think it cannot be doubted that during the breeding season all the mice (with the exception of a few males or females in excess) ordinarily pair; and therefore that this astounding increase during three years must be attributed to a greater than usual number surviving the first year, and then breeding, and so on, till the third year, when their numbers were brought down to their usual limits on the return of wet weather. Where man has introduced plants and animals into a new country favourable to them, there are many accounts in how surprisingly few years the whole country has become stocked with them. This increase would necessarily stop as soon as the country was fully stocked; and yet we have every reason to believe from what is known of wild animals that all would pair in the spring. In the majority of cases it is most difficult to imagine where the check falls, generally no doubt on the seeds, eggs, and young; but when we remember how impossible even in mankind (so much better known than any other animal) it is to infer from repeated casual observations what the average of life is, or to discover how different the percentage of deaths to the births in different countries, we ought to feel no legitimate surprise at not seeing where the check falls in animals and plants. It should always be remembered that in most cases the checks are yearly recurrent in a small regular degree, and in an extreme degree during occasionally unusually cold, hot, dry, or wet years, according to the constitution of the being in question. Lighten any check in the smallest degree, and the geometrical power of increase in every organism will instantly increase the average numbers of the favoured species. Nature may be compared to a surface, on which rest ten thousand sharp wedges touching each other and driven inwards by incessant blows[{230}]. Fully to realise these views much reflection is requisite; Malthus on man should be studied; and all such cases as those of the mice in La Plata, of the cattle and horses when first turned out in S. America, of the robins by our calculation, &c., should be well considered: reflect on the enormous multiplying power inherent and annually in action in all animals; reflect on the countless seeds scattered by a hundred ingenious contrivances, year after year, over the whole face of the land; and yet we have every reason to suppose that the average percentage of every one of the inhabitants of a country will ordinarily remain constant. Finally, let it be borne in mind that this average number of individuals (the external conditions remaining the same) in each country is kept up by recurrent struggles against other species or against external nature (as on the borders of the arctic regions[{231}], where the cold checks life); and that ordinarily each individual of each species holds its place, either by its own struggle and capacity of acquiring nourishment in some period (from the egg upwards) of its life, or by the struggle of its parents (in short lived organisms, when the main check occurs at long intervals) against and compared with other individuals of the same or different species.

But let the external conditions of a country change; if in a small degree, the relative proportions of the inhabitants will in most cases simply be slightly changed; but let the number of inhabitants be small, as in an island[{232}], and free access to it from other countries be circumscribed; and let the change of condition continue progressing (forming new stations); in such case the original inhabitants must cease to be so perfectly adapted to the changed conditions as they originally were. It has been shown that probably such changes of external conditions would, from acting on the reproductive system, cause the organization of the beings most affected to become, as under domestication, plastic. Now can it be doubted from the struggle each individual (or its parents) has to obtain subsistence that any minute variation in structure, habits, or instincts, adapting that individual better to the new conditions, would tell upon its vigour and health? In the struggle it would have a better chance of surviving, and those of its offspring which inherited the variation, let it be ever so slight, would have a better chance to survive. Yearly more are bred than can survive; the smallest grain in the balance, in the long run, must tell on which death shall fall, and which shall survive[{233}]. Let this work of selection, on the one hand, and death on the other, go on for a thousand generations; who would pretend to affirm that it would produce no effect, when we remember what in a few years Bakewell effected in cattle and Western in sheep, by this identical principle of selection.

To give an imaginary example, from changes in progress on an island, let the organization[{234}] of a canine animal become slightly plastic, which animal preyed chiefly on rabbits, but sometimes on hares; let these same changes cause the number of rabbits very slowly to decrease and the number of hares to increase; the effect of this would be that the fox or dog would be driven to try to catch more hares, and his numbers would tend to decrease; his organization, however, being slightly plastic, those individuals with the lightest forms, longest limbs, and best eye-sight (though perhaps with less cunning or scent) would be slightly favoured, let the difference be ever so small, and would tend to live longer and to survive during that time of the year when food was shortest; they would also rear more young, which young would tend to inherit these slight peculiarities. The less fleet ones would be rigidly destroyed. I can see no more reason to doubt but that these causes in a thousand generations would produce a marked effect, and adapt the form of the fox to catching hares instead of rabbits, than that greyhounds can be improved by selection and careful breeding. So would it be with plants under similar circumstances; if the number of individuals of a species with plumed seeds could be increased by greater powers of dissemination within its own area (that is if the check to increase fell chiefly on the seeds), those seeds which were provided with ever so little more down, or with a plume placed so as to be slightly more acted on by the winds, would in the long run tend to be most disseminated; and hence a greater number of seeds thus formed would germinate, and would tend to produce plants inheriting this slightly better adapted down.

Besides this natural means of selection, by which those individuals are preserved, whether in their egg or seed or in their mature state, which are best adapted to the place they fill in nature, there is a second agency at work in most bisexual animals tending to produce the same effect, namely the struggle of the males for the females. These struggles are generally decided by the law of battle; but in the case of birds, apparently, by the charms of their song[{235}], by their beauty or their power of courtship, as in the dancing rock-thrush of Guiana. Even in the animals which pair there seems to be an excess of males which would aid in causing a struggle: in the polygamous animals[{236}], however, as in deer, oxen, poultry, we might expect there would be severest struggle: is it not in the polygamous animals that the males are best formed for mutual war? The most vigorous males, implying perfect adaptation, must generally gain the victory in their several contests. This kind of selection, however, is less rigorous than the other; it does not require the death of the less successful, but gives to them fewer descendants. This struggle falls, moreover, at a time of year when food is generally abundant, and perhaps the effect chiefly produced would be the alteration of sexual characters, and the selection of individual forms, no way related to their power of obtaining food, or of defending themselves from their natural enemies, but of fighting one with another. This natural struggle amongst the males may be compared in effect, but in a less degree, to that produced by those agriculturalists who pay less attention to the careful selection of all the young animals which they breed and more to the occasional use of a choice male[{237}].

Differences between “Races” and “Species”:—first, in their trueness or variability.

Races[{238}] produced by these natural means of selection[{239}] we may expect would differ in some respects from those produced by man. Man selects chiefly by the eye, and is not able to perceive the course of every vessel and nerve, or the form of the bones, or whether the internal structure corresponds to the outside shape. He[{240}] is unable to select shades of constitutional differences, and by the protection he affords and his endeavours to keep his property alive, in whatever country he lives, he checks, as much as lies in his power, the selecting action of nature, which will, however, go on to a lesser degree with all living things, even if their length of life is not determined by their own powers of endurance. He has bad judgment, is capricious, he does not, or his successors do not, wish to select for the same exact end for hundreds of generations. He cannot always suit the selected form to the properest conditions; nor does he keep those conditions uniform: he selects that which is useful to him, not that best adapted to those conditions in which each variety is placed by him: he selects a small dog, but feeds it highly; he selects a long-backed dog, but does not exercise it in any peculiar manner, at least not during every generation. He seldom allows the most vigorous males to struggle for themselves and propagate, but picks out such as he possesses, or such as he prefers, and not necessarily those best adapted to the existing conditions. Every agriculturalist and breeder knows how difficult it is to prevent an occasional cross with another breed. He often grudges to destroy an individual which departs considerably from the required type. He often begins his selection by a form or sport considerably departing from the parent form. Very differently does the natural law of selection act; the varieties selected differ only slightly from the parent forms[{241}]; the conditions are constant for long periods and change slowly; rarely can there be a cross; the selection is rigid and unfailing, and continued through many generations; a selection can never be made without the form be better adapted to the conditions than the parent form; the selecting power goes on without caprice, and steadily for thousands of years adapting the form to these conditions. The selecting power is not deceived by external appearances, it tries the being during its whole life; and if less well «?» adapted than its congeners, without fail it is destroyed; every part of its structure is thus scrutinised and proved good towards the place in nature which it occupies.

We have every reason to believe that in proportion to the number of generations that a domestic race is kept free from crosses, and to the care employed in continued steady selection with one end in view, and to the care in not placing the variety in conditions unsuited to it; in such proportion does the new race become “true” or subject to little variation[{242}]. How incomparably “truer” then would a race produced by the above rigid, steady, natural means of selection, excellently trained and perfectly adapted to its conditions, free from stains of blood or crosses, and continued during thousands of years, be compared with one produced by the feeble, capricious, misdirected and ill-adapted selection of man. Those races of domestic animals produced by savages, partly by the inevitable conditions of their life, and partly unintentionally by their greater care of the individuals most valuable to them, would probably approach closest to the character of a species; and I believe this is the case. Now the characteristic mark of a species, next, if not equal in importance to its sterility when crossed with another species, and indeed almost the only other character (without we beg the question and affirm the essence of a species, is its not having descended from a parent common to any other form), is the similarity of the individuals composing the species, or in the language of agriculturalists their “trueness.”

Difference between “Races” and “Species” in fertility when crossed.

The sterility of species, or of their offspring, when crossed has, however, received more attention than the uniformity in character of the individuals composing the species. It is exceedingly natural that such sterility[{243}] should have been long thought the certain characteristic of species. For it is obvious that if the allied different forms which we meet with in the same country could cross together, instead of finding a number of distinct species, we should have a confused and blending series. The fact however of a perfect gradation in the degree of sterility between species, and the circumstance of some species most closely allied (for instance many species of crocus and European heaths) refusing to breed together, whereas other species, widely different, and even belonging to distinct genera, as the fowl and the peacock, pheasant and grouse[{244}], Azalea and Rhododendron, Thuja and Juniperus, breeding together ought to have caused a doubt whether the sterility did not depend on other causes, distinct from a law, coincident with their creation. I may here remark that the fact whether one species will or will not breed with another is far less important than the sterility of the offspring when produced; for even some domestic races differ so greatly in size (as the great stag-greyhound and lap-dog, or cart-horse and Burmese ponies) that union is nearly impossible; and what is less generally known is, that in plants Kölreuter has shown by hundreds of experiments that the pollen of one species will fecundate the germen of another species, whereas the pollen of this latter will never act on the germen of the former; so that the simple fact of mutual impregnation certainly has no relation whatever to the distinctness in creation of the two forms. When two species are attempted to be crossed which are so distantly allied that offspring are never produced, it has been observed in some cases that the pollen commences its proper action by exserting its tube, and the germen commences swelling, though soon afterwards it decays. In the next stage in the series, hybrid offspring are produced though only rarely and few in number, and these are absolutely sterile: then we have hybrid offspring more numerous, and occasionally, though very rarely, breeding with either parent, as is the case with the common mule. Again, other hybrids, though infertile inter se, will breed quite freely with either parent, or with a third species, and will yield offspring generally infertile, but sometimes fertile; and these latter again will breed with either parent, or with a third or fourth species: thus Kölreuter blended together many forms. Lastly it is now admitted by those botanists who have longest contended against the admission, that in certain families the hybrid offspring of many of the species are sometimes perfectly fertile in the first generation when bred together: indeed in some few cases Mr Herbert[{245}] found that the hybrids were decidedly more fertile than either of their pure parents. There is no way to escape from the admission that the hybrids from some species of plants are fertile, except by declaring that no form shall be considered as a species, if it produces with another species fertile offspring: but this is begging the question[{246}]. It has often been stated that different species of animals have a sexual repugnance towards each other; I can find no evidence of this; it appears as if they merely did not excite each others passions. I do not believe that in this respect there is any essential distinction between animals and plants; and in the latter there cannot be a feeling of repugnance.

Causes of Sterility in Hybrids.

The difference in nature between species which causes the greater or lesser degree of sterility in their offspring appears, according to Herbert and Kölreuter, to be connected much less with external form, size, or structure, than with constitutional peculiarities; by which is meant their adaptation to different climates, food and situation, &c.: these peculiarities of constitution probably affect the entire frame, and no one part in particular[{247}].

From the foregoing facts I think we must admit that there exists a perfect gradation in fertility between species which when crossed are quite fertile (as in Rhododendron, Calceolaria, &c.), and indeed in an extraordinary degree fertile (as in Crinum), and those species which never produce offspring, but which by certain effects (as the exsertion of the pollen-tube) evince their alliance. Hence, I conceive, we must give up sterility, although undoubtedly in a lesser or greater degree of very frequent occurrence, as an unfailing mark by which species can be distinguished from races, i.e. from those forms which have descended from a common stock.

Infertility from causes distinct from hybridisation.

Let us see whether there are any analogous facts which will throw any light on this subject, and will tend to explain why the offspring of certain species, when crossed, should be sterile, and not others, without requiring a distinct law connected with their creation to that effect. Great numbers, probably a large majority of animals when caught by man and removed from their natural conditions, although taken very young, rendered quite tame, living to a good old age, and apparently quite healthy, seem incapable under these circumstances of breeding[{248}]. I do not refer to animals kept in menageries, such as at the Zoological Gardens, many of which, however, appear healthy and live long and unite but do not produce; but to animals caught and left partly at liberty in their native country. Rengger[{249}] enumerates several caught young and rendered tame, which he kept in Paraguay, and which would not breed: the hunting leopard or cheetah and elephant offer other instances; as do bears in Europe, and the 25 species of hawks, belonging to different genera, thousands of which have been kept for hawking and have lived for long periods in perfect vigour. When the expense and trouble of procuring a succession of young animals in a wild state be borne in mind, one may feel sure that no trouble has been spared in endeavours to make them breed. So clearly marked is this difference in different kinds of animals, when captured by man, that St Hilaire makes two great classes of animals useful to man:—the tame, which will not breed, and the domestic which will breed in domestication. From certain singular facts we might have supposed that the non-breeding of animals was owing to some perversion of instinct. But we meet with exactly the same class of facts in plants: I do not refer to the large number of cases where the climate does not permit the seed or fruit to ripen, but where the flowers do not “set,” owing to some imperfection of the ovule or pollen. The latter, which alone can be distinctly examined, is often manifestly imperfect, as any one with a microscope can observe by comparing the pollen of the Persian and Chinese lilacs[{250}] with the common lilac; the two former species (I may add) are equally sterile in Italy as in this country. Many of the American bog plants here produce little or no pollen, whilst the Indian species of the same genera freely produce it. Lindley observes that sterility is the bane of the horticulturist[{251}]: Linnæus has remarked on the sterility of nearly all alpine flowers when cultivated in a lowland district[{252}]. Perhaps the immense class of double flowers chiefly owe their structure to an excess of food acting on parts rendered slightly sterile and less capable of performing their true function, and therefore liable to be rendered monstrous, which monstrosity, like any other disease, is inherited and rendered common. So far from domestication being in itself unfavourable to fertility, it is well known that when an organism is once capable of submission to such conditions «its» fertility is increased[{253}] beyond the natural limit. According to agriculturists, slight changes of conditions, that is of food or habitation, and likewise crosses with races slightly different, increase the vigour and probably the fertility of their offspring. It would appear also that even a great change of condition, for instance, transportal from temperate countries to India, in many cases does not in the least affect fertility, although it does health and length of life and the period of maturity. When sterility is induced by domestication it is of the same kind, and varies in degree, exactly as with hybrids: for be it remembered that the most sterile hybrid is no way monstrous; its organs are perfect, but they do not act, and minute microscopical investigations show that they are in the same state as those of pure species in the intervals of the breeding season. The defective pollen in the cases above alluded to precisely resembles that of hybrids. The occasional breeding of hybrids, as of the common mule, may be aptly compared to the most rare but occasional reproduction of elephants in captivity. The cause of many exotic Geraniums producing (although in vigorous health) imperfect pollen seems to be connected with the period when water is given them[{254}]; but in the far greater majority of cases we cannot form any conjecture on what exact cause the sterility of organisms taken from their natural conditions depends. Why, for instance, the cheetah will not breed whilst the common cat and ferret (the latter generally kept shut up in a small box) do,—why the elephant will not whilst the pig will abundantly—why the partridge and grouse in their own country will not, whilst several species of pheasants, the guinea-fowl from the deserts of Africa and the peacock from the jungles of India, will. We must, however, feel convinced that it depends on some constitutional peculiarities in these beings not suited to their new condition; though not necessarily causing an ill state of health. Ought we then to wonder much that those hybrids which have been produced by the crossing of species with different constitutional tendencies (which tendencies we know to be eminently inheritable) should be sterile: it does not seem improbable that the cross from an alpine and lowland plant should have its constitutional powers deranged, in nearly the same manner as when the parent alpine plant is brought into a lowland district. Analogy, however, is a deceitful guide, and it would be rash to affirm, although it may appear probable, that the sterility of hybrids is due to the constitutional peculiarities of one parent being disturbed by being blended with those of the other parent in exactly the same manner as it is caused in some organic beings when placed by man out of their natural conditions[{255}]. Although this would be rash, it would, I think, be still rasher, seeing that sterility is no more incidental to all cross-bred productions than it is to all organic beings when captured by man, to assert that the sterility of certain hybrids proved a distinct creation of their parents.

But it may be objected[{256}] (however little the sterility of certain hybrids is connected with the distinct creations of species), how comes it, if species are only races produced by natural selection, that when crossed they so frequently produce sterile offspring, whereas in the offspring of those races confessedly produced by the arts of man there is no one instance of sterility. There is not much difficulty in this, for the races produced by the natural means above explained will be slowly but steadily selected; will be adapted to various and diverse conditions, and to these conditions they will be rigidly confined for immense periods of time; hence we may suppose that they would acquire different constitutional peculiarities adapted to the stations they occupy; and on the constitutional differences between species their sterility, according to the best authorities, depends. On the other hand man selects by external appearance[{257}]; from his ignorance, and from not having any test at least comparable in delicacy to the natural struggle for food, continued at intervals through the life of each individual, he cannot eliminate fine shades of constitution, dependent on invisible differences in the fluids or solids of the body; again, from the value which he attaches to each individual, he asserts his utmost power in contravening the natural tendency of the most vigorous to survive. Man, moreover, especially in the earlier ages, cannot have kept his conditions of life constant, and in later ages his stock pure. Until man selects two varieties from the same stock, adapted to two climates or to other different external conditions, and confines each rigidly for one or several thousand years to such conditions, always selecting the individuals best adapted to them, he cannot be said to have even commenced the experiment. Moreover, the organic beings which man has longest had under domestication have been those which were of the greatest use to him, and one chief element of their usefulness, especially in the earlier ages, must have been their capacity to undergo sudden transportals into various climates, and at the same time to retain their fertility, which in itself implies that in such respects their constitutional peculiarities were not closely limited. If the opinion already mentioned be correct, that most of the domestic animals in their present state have descended from the fertile commixture of wild races or species, we have indeed little reason now to expect infertility between any cross of stock thus descended.

It is worthy of remark, that as many organic beings, when taken by man out of their natural conditions, have their reproductive system «so» affected as to be incapable of propagation, so, we saw in the first chapter, that although organic beings when taken by man do propagate freely, their offspring after some generations vary or sport to a degree which can only be explained by their reproductive system being «in» some way affected. Again, when species cross, their offspring are generally sterile; but it was found by Kölreuter that when hybrids are capable of breeding with either parent, or with other species, that their offspring are subject after some generations to excessive variation[{258}]. Agriculturists, also, affirm that the offspring from mongrels, after the first generation, vary much. Hence we see that both sterility and variation in the succeeding generations are consequent both on the removal of individual species from their natural states and on species crossing. The connection between these facts may be accidental, but they certainly appear to elucidate and support each other,—on the principle of the reproductive system of all organic beings being eminently sensitive to any disturbance, whether from removal or commixture, in their constitutional relations to the conditions to which they are exposed.

Points of Resemblance between “Races” and “Species[{259}].”

Races and reputed species agree in some respects, although differing from causes which, we have seen, we can in some degree understand, in the fertility and “trueness” of their offspring. In the first place, there is no clear sign by which to distinguish races from species, as is evident from the great difficulty experienced by naturalists in attempting to discriminate them. As far as external characters are concerned, many of the races which are descended from the same stock differ far more than true species of the same genus; look at the willow-wrens, some of which skilful ornithologists can hardly distinguish from each other except by their nests; look at the wild swans, and compare the distinct species of these genera with the races of domestic ducks, poultry, and pigeons; and so again with plants, compare the cabbages, almonds, peaches and nectarines, &c. with the species of many genera. St Hilaire has even remarked that there is a greater difference in size between races, as in dogs (for he believes all have descended from one stock), than between the species of any one genus; nor is this surprising, considering that amount of food and consequently of growth is the element of change over which man has most power. I may refer to a former statement, that breeders believe the growth of one part or strong action of one function causes a decrease in other parts; for this seems in some degree analogous to the law of “organic compensation[{260}],” which many naturalists believe holds good. To give an instance of this law of compensation,—those species of Carnivora which have the canine teeth greatly developed have certain molar teeth deficient; or again, in that division of the Crustaceans in which the tail is much developed, the thorax is little so, and the converse. The points of difference between different races is often strikingly analogous to that between species of the same genus: trifling spots or marks of colour[{261}] (as the bars on pigeons’ wings) are often preserved in races of plants and animals, precisely in the same manner as similar trifling characters often pervade all the species of a genus, and even of a family. Flowers in varying their colours often become veined and spotted and the leaves become divided like true species: it is known that the varieties of the same plant never have red, blue and yellow flowers, though the hyacinth makes a very near approach to an exception[{262}]; and different species of the same genus seldom, though sometimes they have flowers of these three colours. Dun-coloured horses having a dark stripe down their backs, and certain domestic asses having transverse bars on their legs, afford striking examples of a variation analogous in character to the distinctive marks of other species of the same genus.

External characters of Hybrids and Mongrels.

There is, however, as it appears to me, a more important method of comparison between species and races, namely the character of the offspring[{263}] when species are crossed and when races are crossed: I believe, in no one respect, except in sterility, is there any difference. It would, I think, be a marvellous fact, if species have been formed by distinct acts of creation, that they should act upon each other in uniting, like races descended from a common stock. In the first place, by repeated crossing one species can absorb and wholly obliterate the characters of another, or of several other species, in the same manner as one race will absorb by crossing another race. Marvellous, that one act of creation should absorb another or even several acts of creation! The offspring of species, that is hybrids, and the offspring of races, that is mongrels, resemble each other in being either intermediate in character (as is most frequent in hybrids) or in resembling sometimes closely one and sometimes the other parent; in both the offspring produced by the same act of conception sometimes differ in their degree of resemblance; both hybrids and mongrels sometimes retain a certain part or organ very like that of either parent, both, as we have seen, become in succeeding generations variable; and this tendency to vary can be transmitted by both; in both for many generations there is a strong tendency to reversion to their ancestral form. In the case of a hybrid laburnum and of a supposed mongrel vine different parts of the same plants took after each of their two parents. In the hybrids from some species, and in the mongrel of some races, the offspring differ according as which of the two species, or of the two races, is the father (as in the common mule and hinny) and which the mother. Some races will breed together, which differ so greatly in size, that the dam often perishes in labour; so it is with some species when crossed; when the dam of one species has borne offspring to the male of another species, her succeeding offspring are sometimes stained (as in Lord Morton’s mare by the quagga, wonderful as the fact[{264}] is) by this first cross; so agriculturists positively affirm is the case when a pig or sheep of one breed has produced offspring by the sire of another breed.

Summary of second chapter[{265}].

Let us sum up this second chapter. If slight variations do occur in organic beings in a state of nature; if changes of condition from geological causes do produce in the course of ages effects analogous to those of domestication on any, however few, organisms; and how can we doubt it,—from what is actually known, and from what may be presumed, since thousands of organisms taken by man for sundry uses, and placed in new conditions, have varied. If such variations tend to be hereditary; and how can we doubt it,—when we see shades of expression, peculiar manners, monstrosities of the strangest kinds, diseases, and a multitude of other peculiarities, which characterise and form, being inherited, the endless races (there are 1200 kinds of cabbages[{266}]) of our domestic plants and animals. If we admit that every organism maintains its place by an almost periodically recurrent struggle; and how can we doubt it,—when we know that all beings tend to increase in a geometrical ratio (as is instantly seen when the conditions become for a time more favourable); whereas on an average the amount of food must remain constant, if so, there will be a natural means of selection, tending to preserve those individuals with any slight deviations of structure more favourable to the then existing conditions, and tending to destroy any with deviations of an opposite nature. If the above propositions be correct, and there be no law of nature limiting the possible amount of variation, new races of beings will,—perhaps only rarely, and only in some few districts,—be formed.

Limits of Variation.

That a limit to variation does exist in nature is assumed by most authors, though I am unable to discover a single fact on which this belief is grounded[{267}]. One of the commonest statements is that plants do not become acclimatised; and I have even observed that kinds not raised by seed, but propagated by cuttings, &c., are instanced. A good instance has, however, been advanced in the case of kidney beans, which it is believed are now as tender as when first introduced. Even if we overlook the frequent introduction of seed from warmer countries, let me observe that as long as the seeds are gathered promiscuously from the bed, without continual observation and careful selection of those plants which have stood the climate best during their whole growth, the experiment of acclimatisation has hardly been begun. Are not all those plants and animals, of which we have the greatest number of races, the oldest domesticated? Considering the quite recent progress[{268}] of systematic agriculture and horticulture, is it not opposed to every fact, that we have exhausted the capacity of variation in our cattle and in our corn,—even if we have done so in some trivial points, as their fatness or kind of wool? Will any one say, that if horticulture continues to flourish during the next few centuries, that we shall not have numerous new kinds of the potato and Dahlia? But take two varieties of each of these plants, and adapt them to certain fixed conditions and prevent any cross for 5000 years, and then again vary their conditions; try many climates and situations; and who[{269}] will predict the number and degrees of difference which might arise from these stocks? I repeat that we know nothing of any limit to the possible amount of variation, and therefore to the number and differences of the races, which might be produced by the natural means of selection, so infinitely more efficient than the agency of man. Races thus produced would probably be very “true”; and if from having been adapted to different conditions of existence, they possessed different constitutions, if suddenly removed to some new station, they would perhaps be sterile and their offspring would perhaps be infertile. Such races would be undistinguishable from species. But is there any evidence that the species, which surround us on all sides, have been thus produced? This is a question which an examination of the economy of nature we might expect would answer either in the affirmative or negative[{270}].

CHAPTER III
ON THE VARIATION OF INSTINCTS AND OTHER MENTAL ATTRIBUTES UNDER DOMESTICATION AND IN STATE OF NATURE; ON THE DIFFICULTIES IN THIS SUBJECT; AND ON ANALOGOUS DIFFICULTIES WITH RESPECT TO CORPOREAL STRUCTURES

Variation of mental attributes under domestication.

I have as yet only alluded to the mental qualities which differ greatly in different species. Let me here premise that, as will be seen in the Second Part, there is no evidence and consequently no attempt to show that all existing organisms have descended from any one common parent-stock, but that only those have so descended which, in the language of naturalists, are clearly related to each other. Hence the facts and reasoning advanced in this chapter do not apply to the first origin of the senses[{271}], or of the chief mental attributes, such as of memory, attention, reasoning, &c., &c., by which most or all of the great related groups are characterised, any more than they apply to the first origin of life, or growth, or the power of reproduction. The application of such facts as I have collected is merely to the differences of the primary mental qualities and of the instincts in the species[{272}] of the several great groups. In domestic animals every observer has remarked in how great a degree, in the individuals of the same species, the dispositions, namely courage, pertinacity, suspicion, restlessness, confidence, temper, pugnaciousness, affection, care of their young, sagacity, &c., &c., vary. It would require a most able metaphysician to explain how many primary qualities of the mind must be changed to cause these diversities of complex dispositions. From these dispositions being inherited, of which the testimony is unanimous, families and breeds arise, varying in these respects. I may instance the good and ill temper of different stocks of bees and of horses,—the pugnacity and courage of game fowls,—the pertinacity of certain dogs, as bull-dogs, and the sagacity of others,—for restlessness and suspicion compare a wild rabbit reared with the greatest care from its earliest age with the extreme tameness of the domestic breed of the same animal. The offspring of the domestic dogs which have run wild in Cuba[{273}], though caught quite young, are most difficult to tame, probably nearly as much so as the original parent-stock from which the domestic dog descended. The habitual “periods” of different families of the same species differ, for instance, in the time of year of reproduction, and the period of life when the capacity is acquired, and the hour of roosting (in Malay fowls), &c., &c. These periodical habits are perhaps essentially corporeal, and may be compared to nearly similar habits in plants, which are known to vary extremely. Consensual movements (as called by Müller) vary and are inherited,—such as the cantering and ambling paces in horses, the tumbling of pigeons, and perhaps the handwriting, which is sometimes so similar between father and sons, may be ranked in this class. Manners, and even tricks which perhaps are only peculiar manners, according to W. Hunter and my father, are distinctly inherited in cases where children have lost their parent in early infancy. The inheritance of expression, which often reveals the finest shades of character, is familiar to everyone.

Again the tastes and pleasures of different breeds vary, thus the shepherd-dog delights in chasing the sheep, but has no wish to kill them,—the terrier (see Knight) delights in killing vermin, and the spaniel in finding game. But it is impossible to separate their mental peculiarities in the way I have done: the tumbling of pigeons, which I have instanced as a consensual movement, might be called a trick and is associated with a taste for flying in a close flock at a great height. Certain breeds of fowls have a taste for roosting in trees. The different actions of pointers and setters might have been adduced in the same class, as might the peculiar manner of hunting of the spaniel. Even in the same breed of dogs, namely in fox-hounds, it is the fixed opinion of those best able to judge that the different pups are born with different tendencies; some are best to find their fox in the cover; some are apt to run straggling, some are best to make casts and to recover the lost scent, &c.; and that these peculiarities undoubtedly are transmitted to their progeny. Or again the tendency to point might be adduced as a distinct habit which has become inherited,—as might the tendency of a true sheep dog (as I have been assured is the case) to run round the flock instead of directly at them, as is the case with other young dogs when attempted to be taught. The "transandantes" sheep[{274}] in Spain, which for some centuries have been yearly taken a journey of several hundred miles from one province to another, know when the time comes, and show the greatest restlessness (like migratory birds in confinement), and are prevented with difficulty from starting by themselves, which they sometimes do, and find their own way. There is a case on good evidence[{275}] of a sheep which, when she lambed, would return across a mountainous country to her own birth-place, although at other times of year not of a rambling disposition. Her lambs inherited this same disposition, and would go to produce their young on the farm whence their parent came; and so troublesome was this habit that the whole family was destroyed.