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Edited by his son 

Honorary Fellow of Christ's College 

Cambridge : 

Printed at the University Press 

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. 









23 June, 1909. 




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 


iv. and v. On the evidence from Geology. (The reasons for 
combining the two sections are given in the Intro- 
duction) 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 

Portrait frontispiece 

Facsimile to face p. 50 



[ ] 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. 


WE know from the contents of Charles Darwin's 
Note Book of 1837 that he was at that time a con- 
vinced 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 ex- 
termination 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. 

1 See the extracts in Life and Letters of Charles Darwin, ii. p. 5. 

2 The second volume, especially important in regard to Evolution, 
reached him in the autumn of 1832, as Prof. Judd has pointed out in his 
most interesting paper in Dancin and Modern Science. Cambridge, 1909. 

3 Obituary Notice of C. Darwin, Proc. R. Soc. vol. 44. Reprinted in 
Huxley's Collected Essays. See also Life and Letters of C. Darwin, ii. 
p. 179. 

4 See the extracts in the Life and Letters, ii. p. 5, 



They die, without they change, like golden pippins ; 
it is a generation of species like generation of indi- 

"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 scienti- 
fically 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 1 he wrote: "During the voyage 
of the Beagle I had been deeply impressed by dis- 
covering 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 2 held that these facts could not have pro- 
duced their essential effect until the voyage had 

1 Life and Letters, i. p. 82. 2 Obituary Notice, loc. cit. 


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 1 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 pre- 
paration 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 2 had not yet attained any 

1 Darwin and Modern Science. 

2 Huxley, Obituary, p. xi. 


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 1 on 
character of South American fossils, and species 
on Galapagos Archipelago. These facts origin (es- 
pecially 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 2 ' by comparing the birds shot 
by himself and by others on board. The case must 
have struck him at once, without waiting for accu- 
rate 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 sug- 
gests 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 Gala- 
pagos 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 mini- 
mised the earlier beginnings of his theorisings, and 
laid more stress on the recent thoughts which were 

1 In this citation the italics are mine. 

2 Journal of Researches, Ed. 1860, p. 394. 


naturally more vivid to him. In his letter 1 to Otto 
Zacharias (1877) he wrote, "On my return home in 
the autumn of 1836, 1 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 per- 
manence 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 recollec- 
tion 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 2 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, 

1 F. Darwin's Life of Charles Darwin (in one volume), 1892, p. 166. 

2 Life and Letters, i. p. 83. 


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 ex- 
pressed the following forecast 1 of the importance 
of the survival of the fittest. "With respect to 
extinction, we can easily see that a variety of the 
ostrich (Petise 2 ), may not be well adapted, and 
thus perish out; or on the other hand, like Or- 
pheus 3 , being favourable, many might be produced. 
This requires the principle that the permanent 
variations produced by confined breeding and 
changing circumstances are continued and pro- 
duce^!) according to the adaptation of such circum- 
stances, 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 4 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 5 , which I had fairly 
copied out and still possess." It is the first of these 
Essays, the one in 35 pages, which is now printed 
under the title The Foundations of the Origin of 

1 Life and Letters, ii. p. 8. 2 Avestruz Petise, i.e. Rhea Darwini. 

3 A bird. 

4 Life and Letters, i. p. 84. 

5 It contains as a fact 231 pp. It is a strongly bound folio, interleaved 
with blank pages, as though for notes and additions. His own MS. from 
which it was copied contains 189 pp. 


It will be noted that in the above passage he 
does not mention the MS. of the Foundations 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 cup- 
board 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 
father's Diary: 

"1842. May 18th went to Maer. June loth 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 1 ." 
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 Foundations ; 
but there is evidence in favour of an earlier date 2 . 
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 3 (Jan. 25, 1859) he speaks of his own 
contributions to the Linnean paper 4 of July 1, 1858, 
as " written in 1 839, 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 Foundations, it must, 
I think, be rejected. I can only account for his 
mistake by the supposition that my father had in 

1 Life and Letters, ii. p. 116. 2 Life and Letters, ii. p. 10. 

3 Life and Letters, ii. p. 146. 4 J. Linn. Soc. Zool. iii. p. 45. 


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 evolu- 
tion, together with words intended as headings for 
discussion, such as "Affinity, unity of type,- 
foetal 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 Foundations, 
consisting of the titles of the three chapters of 
which it was to have consisted. 

" I. The Principles of Var. in domestic organ- 

" II. The possible and probable application of 
these same principles to wild animals and conse- 
quently 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 
Foundations, 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 origin- 
ally Chapter III 1 accounts for the curious error 
which occurs in pp. 18 and 22 where the second 
Part of the Foundations is called Part III. 

The division of the Essay into two parts is main- 
tained 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 Foundations. 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 2 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 

1 It is evident that Parts and Chapters were to some extent inter- 
changeable in the author's mind, for p. 1 (of the MS. we have been dis- 
cussing) is headed in ink Chapter I, and afterwards altered in pencil to 
Part 1. 

2 On p. 23 of the MS. of the Foundations is a reference to the "back of 
p. 21 bis": this suggests that additional pages had been interpolated in 
the MS. and that it may once have had 37 in place of 35 pp. 


The sketch 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 struc- 
ture. There is much erasure and correction, appa- 
rently 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 memo- 
randa 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 1 . This matter consists partly of passages 
marked for insertion in the text, and these can gener- 
ally (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 (?). 

1 Life and Letters, i. p. 153. 


Lastly, words inserted by the editor, of which 
the appropriateness is doubtful, are printed thus 
(variation ?). 

Two kinds of erasure occur in the MS. 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 hori- 
zontal 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 intro- 
duced 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 un- 
necessary 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 

The sections into which the Essay 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 vm., and n. 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 Extermi- 
nation 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 Foundations correspond to the ten chapters 
of the MS. of 1844. 

The Origin of Species differs from the Founda- 
tions 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, varia- 
tion 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 
Foundations 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 Foundations corresponds 
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 dis- 
tribution ; affinities and classification ; unity of type 


and morphology; abortive or rudimentary organs; 
recapitulation and conclusion. 

The fact that 17 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 1 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 diver- 
gence is of course also a characteristic of the 
Foundations. 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." But if the descendants of A 
have peopled the earth by beating out other forms, 
they must have diverged in occupying the innumer- 
able diverse modes of life from which they expelled 
their predecessors. What I wrote 2 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." 

I have called attention in footnotes to many 
minor points in which the Origin agrees with the 
Foundations. One of the most interesting is the 
final sentence on p. 52, which is almost identical with 
the concluding words of the Origin. I have else- 

1 Life and Letters, i. p. 84. - Life and Letters, ii. p. 15. 


where pointed out 1 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 Bridyewater Treatise is printed opposite 
the title-page of the Origin of Species. 

Among other interesting points may be men- 
tioned 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 187G. We find 
him also (p. 2) recognising the importance of germinal 
variation, where he speaks of external conditions 
acting indirectly through the "reproductive func- 
tions." He seems to have had constantly in mind the 
need of referring each variation to a cause, a point 
of view to which he returned at the close of his life. 
This subject, though by no means wanting in the 
Origin, is there overshadowed by considerations 
which then seemed to him more pressing. 

In conclusion, I desire to express my thanks to 
Mr Wallace for a footnote he was good enough to 
supply: and to Professor Judd and Professor 
Bateson for suggestions of value. I am also 
indebted to Mr Rutherford, of the University 
Library, for his careful copy of the manuscript. 

1 Life and Letters, ii. p. 9. 


April 16, 1909. 



AN individual organism placed under new con- 
ditions [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) 1 and continues 
steadily to propagate, by buds, such new kind. 

When the organism is bred for several genera- 
tions under new or varying conditions, the variation 
is greater in amount and endless in kind [especially 2 
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 

1 Evidently a memorandum that an example should be given. 

2 The importance of exposure to new conditions for several generations 
is insisted on in the Origin, Ed. i. p. 7, also p. 131. In the latter passage 
the author guards himself against the assumption that variations are " due 
to chance," and speaks of "our ignorance of the cause of each particular 
variation." These statements are not always remembered by his critics. 

D. 1 


variation (Miiller's twins) 1 seems inevitable effect 
of process of reproduction. But more important 
is that simple (?) generation, especially under new 
conditions [when no crossing] (causes) infinite varia- 
tion and not direct effect of external conditions, 
but only in as much as it affects the reproductive 
functions 2 . There seems to be no part (beau ideal 
of liver) 3 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 4 
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 5 . 
All bisexual animals must cross, hermaphrodite 
plants do cross, it seems very possible that her- 

1 Cf. Origin, Ed. i. p. 10, vi. p. 9, "Young of the same litter, sometimes 
differ considerably from each other, though both the young and the parents, 
as Miiller has remarked, have apparently been exposed to exactly the same 
conditions of life." 

2 This is paralleled by the conclusion in the Origin, Ed. i. p. 8, that "the 
most frequent cause of variability may be attributed to the male and 
female reproductive elements having been affected prior to the act of 

3 The meaning seems to be that there must be some variability in the 
liver otherwise anatomists would not speak of the ' beau ideal ' of that 

4 The position of the following passage is uncertain. " If individuals 
of two widely different varieties be allowed to cross, a third race 
will be formed a most fertile source of the variation in domesticated 
animals. (In the Origin, Ed. i. p. 20 the author says that " the possibility 
of making distinct races by crossing has been greatly exaggerated.") If 
freely allowed, the characters of pure parents will be lost, number of 
races thus (illegible) but differences (?) besides the (illegible). But if 
varieties differing in very slight respects be allowed to cross, such small 
variation mil be destroyed, at least to our senses, a variation [clearly] 
just to be distinguished by long legs will have offspring not to be so 
distinguished. Free crossing great agent in producing uniformity in any 
breed. Introduce tendency to revert to parent form." 

5 The swamping effect of intercrossing is referred to in the Origin, 
Ed. i. p. 103., vi. p. 126. 


maphrodite animals do cross, conclusion strength- 
ened : ill effects of breeding in and in, good effects 
of crossing possibly analogous to good effects of 
change in condition (?) 1 . 

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 counter- 
acted. Secondly reversion to parent form analogue 
of vis medicatrix 2 . But if man selects, then new 
races rapidly formed, of late years systematically 
followed, in most ancient times often practically 
followed 3 . 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 4 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 

1 A discussion ou the intercrossing of hermaphrodites in relation to 
Knight's views occurs in the Origin, Ed. i. p. 96, vi. p. 119. The parallelism 
between crossing and changed conditions is briefly given in the Origin, 
Ed. i. p. 267, vi. p. 391, and was finally investigated in The Effects of 
Cross and Self -Fertilisation in the Vegetable Kingdom, 1876. 

2 There is an article on the vis medicatrix in Brougham's Dissertations, 
1839, a copy of which is in the author's library. 

3 This is the classification of selection into methodical and unconscious 
given in the Origin, Ed. i. p. 33, vi. p. 38. 

4 This passage, and a similar discussion on the power of the Creator 
(p. 6), correspond to the comparison between the selective capacities of 
man and nature, in the Origin, Ed. i. p. 83, vi. p. 102. 



[knowledge] power of selecting according to in- 
ternal 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. 


Let us see how far above principles of variation 
apply to wild animals. Wild animals vary exceed- 
ingly little yet they are known as individuals 1 . 
British Plants, in many genera number quite 
uncertain of varieties and species : in shells chiefly 
external conditions 2 . 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 con- 
stant 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. 

1 i.e. they are individually distinguishable. 

2 See Origin, Ed. i. p. 133, vi. p. 165. 


Hence (?) we should expect every now and then 
a wild form to vary l ; 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 con- 
stantly 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 

1 When the author wrote this sketch he seems not to have been so 
fully convinced of the general occuirence of variation in nature as he 
afterwards became. The above passage in the text possibly suggests that 
at this time he laid more stress on sports or mutations than was afterwards 
the case. 


climate adapting woodpecker l to crawl (?) up trees, 
(illegible) miseltoe, (sentence incomplete). But if 
every part of a plant or animal was to vary (il- 
legible), and if a being infinitely more sagacious than 
man (not an omniscient creator) during thousands 
and thousands of years were to select all the varia- 
tions 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 2 . 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 3 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 

1 The author may possibly have taken the case of the woodpecker from 
Buffou, Histoire Nat. des Oiseaux, T. vii. p. 3, 1780, where however it is 
treated from a different point of view. He uses it more than once, see for 
instance Origin, Ed. i. pp. 3, 60, 184, vi. pp. 3, 76, 220. The passage in 
the text corresponds with a discussion on the woodpecker and the mistletoe 
in Origin, Ed. i. p. 3, vi. p. 3. 

- This illustration occurs in the Origin, Ed. i. pp. 90, 91, vi. pp. 110, 111. 

3 See Origin, Ed. i. p. 83, vi. p. 102, where the word Creator is replaced 
by Nature. 


forms descending from one common type 1 . 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 com- 
mon 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 2 . 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 3 . If 
proof were wanted let any singular change of 
climate (occur) here (?), how astoundingly some 
tribes (?) increase, also introduced animals 4 , the 

1 Note in the original. "Good place to introduce, saying reasons 
hereafter to be given, how far I extend theory, say to all mammalia 
reasons growing weaker and weaker." 

2 See Origin, Ed. i. pp. 62, 63, vi. p. 77, where similar reference is made 
to De Candolle ; for Malthus see Origin, p. 5. 

3 This may possibly refer to the amount of destruction going on. See 
Origin, Ed. i. p. 68, vi. p. 84, where there is an estimate of a later date as 
to death-rate of birds in winter. " Calculate robins " probably refers to a 
calculation of the rate of increase of birds under favourable conditions. 

4 In the Origin, Ed. i. pp. 64, 65, vi. p. 80, he instances cattle and horses 
and certain plants in S. America and American species of plants in India, 
and further on, as unexpected effects of changed conditions, the enclosure 
of a heath, and the relation between the fertilisation of clover and the 
presence of cats (Origin, Ed. i. p. 74, vi. p. 91). 


pressure is always ready, capacity of alpine plants 
to endure other climates, think of endless seeds 
scattered abroad, forests regaining their percen- 
tage 1 , a thousand wedges 2 are being forced into 
the oeconomy of nature. This requires much reflec- 
tion; study Mai thus 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 innnitesimally 
small differences must inevitably tell 3 ; when unusu- 
ally 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 4 ) 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 5 increase very slowly from change of 
climate affecting peculiar plants, and some other 

^j \ j. * 

(illegible) rabbit decrease in same proportion [let 
this unsettle organisation of], a canine animal, who 

1 Origin, Ed. i. p. 74, vi. p. 91. "It has been observed that the trees 
now growing on... ancient Indian mounds... display the same beautiful 
diversity and proportion of kinds as in the surrounding virgin forests." 

2 The simile of the wedge occurs in the Origin, Ed. i. p. 67 ; it is deleted 
in Darwin's copy of the first edition : it does not occur in Ed. vi. 

3 In a rough summary at the close of the Essay, occur the words : 
" Every creature lives by a straggle, smallest grain in balance must tell." 

4 Cf. Origin, Ed. i. p. 77, vi. p. 94. 

5 This is a repetition of what is given at p. 6. 


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, care- 
fully 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 pecu- 
liarities approaching to monsters. This requires 
consideration, should be introduced in first chapter 
if it holds, I believe it does. It is hypothetical at 
best 1 . 

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 

1 Compare Origin, Ed. i. p. 41, vi. p. 47. "I have seen it gravely 
remarked, that it was most fortunate that the strawberry began to vary 
just when gardeners began to attend closely to this plant. No doubt the 
strawberry had always varied since it was cultivated, but the slight varieties 
had been neglected." 


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 1 . 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 indus- 
trious, (whose) instincts (are) best developed, will 
rear more young, probably possessing her good 
qualities, and a greater number will thus (be) pre- 
pared 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 wood- 
pecker adapted to tree 2 . 

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 

1 Here we have the two types of sexual selection discussed in the Origin, 
Ed. i. pp. 88 et seq., vi. pp. 108 et seq. 

2 It is not obvious why the author objects to " chance " or "external con- 
ditions making a woodpecker." He allows that variation is ultimately 
referable to conditions and that the nature of the connexion is unknown, i.e. 
that the result is fortuitous. It is not clear in the original to how much of 
the passage the two ? refer. 


be contaminated with a cross of another race, and 
agriculturists know how difficult they find always to 
prevent this, effect would be trueness. This char- 
acter 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 1 . 
Gladiolus, Crinwn, Calceolaria* 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 3 very general. Some nearest species will 
not cross (crocus, some heath (?)), some genera cross 
readily (fowls 4 and grouse, peacock &c.). Hybrids 
no ways monstrous quite perfect except secretions 5 
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 6 ]. Will analogy throw any light 

1 The meaning is "That sterility is not universal is admitted by all." 

2 See Far. under Dom., Ed. 2, i. p. 388, where the garden forms of 
Gladiolus and Calceolaria are said to be derived from crosses between 
distinct species. Herbert's hybrid Crinums are discussed in the Origin, 
Ed. i. p. 250, vi. p. 370. It is well known that the author believed in a 
multiple origin of domestic dogs. 

3 The argument from gradation in sterility is given in the Origin, Ed. i. 
pp. 248, 255, vi. pp. 368, 375. In the Origin, I have not come across the 
cases mentioned, viz. crocus, heath, or grouse and fowl or peacock. For 
sterility between closely allied species, see Origin, Ed. i. p. 257, vi. p. 377. 
In the present essay the author does not distinguish between fertility 
between species and the fertility of the hybrid offspring, a point on which 
he insists in the Origin, Ed. i. p. 245, vi. p. 365. 

4 Ackermann (Ber. d. Vereins f. Naturkunde zu Kassel, 1898, p. 23) 
quotes from Gloger that a cross has been effected between a domestic hen 
and a Tetrao tetrix ; the offspring died when three days old. 

5 No doubt the sexual cells are meant. I do not know on what evidence 
it is stated that the mule has bred. 

6 The sentence is all but illegible. I think that the author refers to 
forms usually ranked as varieties having been marked as species when it was 


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 circum- 
stance is constitutional differences 1 , 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 distinc- 
tion of tame and domestic, elephants, ferrets 2 . 
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 un- 
favourable 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 corn- 
found that they were sterile together. See the case of the red and blue 
Anagallis given from Gartner in the Origin, Ed. i. p. 247, vi. p. 368. 

1 In the Origin, Ed. i. p. 258, where the author speaks of constitutional 
differences in this connexion, he specifies that they are confined to the 
reproductive system. 

2 The sensitiveness of the reproductive system to changed conditions is 
insisted on in the Origin, Ed. i. p. 8, vi. p. 10. 

The ferret is mentioned, as being prolific in captivity, in Var. under 
Dom., Ed. 2, ii. p. 90. 


monest 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 pro- 
pagators, Linnseus about alpine plants. American 
bog plants, pollen in exactly same state as in 
hybrids, same in geraniums. Persian and Chinese 1 
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 2 . There is here gradation (in) sterility 
and then parts, like diseases, are transmitted here- 
ditarily. We cannot assign any cause why the 
Pontic Azalea produces plenty of pollen and not 
American 3 , 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 4 , 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 off- 
spring, an American plant its American-bog 
constitution, and (with) animals, those peculiarities, 
on which 5 when placed out of their natural conditions 
they are incapable of breeding; and moreover they 
transmit every part of their constitution, their 

1 Lindley's remark is quoted in the Origin, Ed. i. p. 9. Linnaeus' 
remark is to the effect that Alpine plants tend to be sterile under cultiva- 
tion (see Var. under Dom., Ed. 2, ii. p. 147). In the same place the author 
speaks of peat-loving plants being sterile in our gardens, no doubt the 
American bog-plants referred to above. On the following page (p. 148) the 
sterility of the lilac (Syringa persica and chinensis] is referred to. 

2 The author probably means that the increase in the petals is clue to 
a greater food supply being available for them owing to sterility. See the 
discussion in Var. under Dom., Ed. 2, ii. p. 151. It must be noted that 
doubleness of the flower may exist without noticeable sterility. 

3 I have not come across this case in the author's works. 

4 For the somewhat doubtful case of the cheetah (Felisjubata) see Var. 
under Dom. y Ed. 2, ii. p. 133. I do not know to what fact "pig in India" 

6 This sentence should run " on which depends their incapacity to breed 
in unnatural conditions." 


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 1 . 
Variation depends on change of condition and 
selection 2 , 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 rigor- 
ously adapted to another, till such can be produced, 
the question is not tried 3 . 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 4 . 

1 This sentence ends in confusion : it should clearly close with the words 
"refused to breed" in place of the bracket and the present concluding 

2 The author doubtless refers to the change produced by the summation 
of variation by means of selection. 

3 The meaning of this sentence is made clear by a passage in the MS. of 
1844: "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." That is, the attempt to produce mutually 
sterile domestic breeds. 

4 This passage is to some extent a repetition of a previous one and may 


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 1 , 
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 reproduc- 
tive faculties and exposure to changed conditions 
of life whether by crossing or exposure of the indi- 
viduals 2 . 

Difficulties on theory of selection 3 . 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 mon- 
strous and to (the) end appears difficulty. But think 
of gradation, even now manifest, (Tibia and Fibula). 
Everyone will allowif every fossil preserved, gradation 

have been intended to replace an earlier sentence. I have thought it best 
to give both. In the Origin, Ed. i. p. 141, vi. p. 176, the author gives his 
opinion that the power of resisting diverse conditions, seen in man and 
his domestic animals, is an example " of a very common flexibility of con- 

1 In the Origin, Ed. i. Chs. i. and v., the author does not admit repro- 
duction, apart from environment, as being'a cause of variation. With regard 
to the cumulative effect of new conditions there are many passages in the 
Origin, Ed. i. e.g. pp. 7, 12, vi. pp. 8, 14. 

' J As already pointed out, this is the important principle investigated 
in the author's Cross and Self -Fertilisation. Professor Bateson has 
suggested to me that the experiments should be repeated with gametically 
pure individuals. 

3 In the Origin a chapter is given up to " difficulties on theory " : the 
discussion in the pi'esent essay seems slight even when it is remembered 
how small a space is here available. For Tibia &c. see p. 48. 


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 meta- 
morphoses ; 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 vertebrae, 
actually vary in domestic animals, less difficult 
if growth followed. Looking to whole animals, a bat 
formed not for flight 1 . Suppose we had flying fish 2 
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 3 . 

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. 

1 This may be interpreted " The general structure of a bat is the same 
as that of non-flying mammals." 

2 That is truly winged fish. 

3 The terrestrial woodpecker of S. America formed the subject of a paper 
by Darwin, Proc. Zool. Soc., 1870. See Life and Letters, vol. iii. p. 153. 




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 1 , but merely with 
their differences in each of the great divisions of 
nature. Disposition, courage, pertinacity (?), sus- 
picion, 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 2 ). 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. and 
d. Habits, as pointing and setting on certain 
occasions d. 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 3 . Origin partly habit, but the amount 
necessarily unknown, partly selection. Young 
pointers pointing stones and sheep tumbling 
pigeons sheep 4 going back to place where born. 

! The same proviso occurs in the Origin, Ed. i. p. 207, vi. p. 319. 

2 The lameness of the birds in the Galapagos is described in the Journal 
of Researches (1860), p. 398. Dogs and rabbits are probably mentioned as 
cases in which the hereditary fear of man has been lost. In the 1844 MS. 
the author states that the Cuban feral dog shows great natural wildness, 
even when caught quite young. 

3 In the Origin, Ed. i. p. 207, vi. p. 319, he refuses to define instinct. 
For Lord Brougham's definition see his Dissertations on Subjects of 
Science etc., 1839, p. 27. 

4 See James Hogg (the Ettrick Shepherd), Works, 1865, Tales and 
Sketches, p. 403. 



Instinct aided by reason, as in the taylor-bird 1 . 
Taught by parents, cows choosing food, birds singing. 
Instincts vary in wild state (birds get wilder) often 
lost 2 ; more perfect, nest without roof. These 
facts [only clear way] show how incomprehensibly 
brain has power of transmitting intellectual opera- 

Faculties 3 distinct from true instincts, finding 
[way]. It must I think be admitted that habits 
whether congenital or acquired by practice [some- 
times] often become inherited 4 ; 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. 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 5 , 1 beg 
this may be remembered, nor do I mean to attempt 
to show exact method. I want only to show that 

1 This refers to the tailor-bird making use of manufactured thread 
supplied to it, instead of thread twisted by itself. 

2 Often lost applies to instinct : birds get wilder is printed in a paren- 
thesis because it was apparently added as an after-thought. Nest without 
roof refers to the water-ousel omitting to vault its nest when building 
in a protected situation. 

3 In the MS. of 1844 is an interesting discussion on faculty as distinct 
from instinct. 

4 At this date and for long afterwards the inheritance of acquired 
characters was assumed to occur. 

5 Part II. is here intended : see the Introduction. 


whole theory ought not at once to be rejected on this 

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 1 , 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 incompre- 
hensible to us, as animal to them, geological 
changes, fishes in river, case of sheep in Spain 2 . 
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 3 , 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, howtheymake 
a hexagon, Waterhouse's theory 4 , 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 

1 The meaning is that the attitude assumed in shamming is not 
accurately like that of death. 

2 This refers to the transandantes sheep mentioned in the MS. of 1844, 
as having acquired a migratory instinct. 

3 In the Origin, Ed. i. p. 209, vi. p. 321, Mozart's pseudo-instinctive 
skill in piano-playing is mentioned. See Phil. Trans., 1770, p. 54. 

4 In the discussion on bees' cells, Origin, Ed. i. p. 225, vi. p. 343, the 
author acknowledges that his theory originated in Waterhouse's obser- 



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 \ 

-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] extraordinari- 
ness (?) 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 selec- 
tion 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 2 , none is 

1 The hawfinch- and Sylvia-types are figured in the Journal of Researches, 
p. 379. The discussion of change of form in relation to change of instinct 
is not clear, and I find it impossible to suggest a paraphrase. 

2 I should interpret this obscure sentence as follows, "No such opposing 
law is known, but in all works on the subject a law is (in flat contradiction 
to all known facts) assumed to limit the possible amount of variation." In 
the Origin, the author never limits the power of variation, as far as I know. 

OF PART I. 21 

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 1 (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. 

1 In Var. under Dom. Ed. 2, ii. p. 263, the Dahlia is described 
as showing sensitiveness to conditions in 1841. All the varieties of the 
Dahlia are said to have arisen since 1804 (ibid. i. p. 393). 


PART II 1 . 


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 2 ; what would the As- 
tronomer 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 pro- 
position (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 

1 In the original MS. the heading is: Part III.; but Part II. is clearly 
intended ; for details see the Introduction. I have not been able to 
discover where IV. ends and v. begins. 

2 This passage corresponds roughly to the conclusion of the Origin, see 
Ed. i. p. 482, vi. p. 661. 


and showed signs of common descent, and yet were 
created distinct. As long- as it was thought im- 
possible that organisms should vary, or should any- 
how become adapted to other organisms in a com- 
plicated manner, and yet be separated from them by 
an impassable barrier of sterility 1 , 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 appear- 
ance, affinities, morphology &c., &c. 

Our theory requires a very gradual introduction 
of new forms 2 , 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^, elephant [or fowl and peacock], 

1 A similar passage occurs in the conclusion of the Origin, Ed. i. p. 481, 
vi. p. 659. 

a See Origin, Ed. i. p. 312, vi. p. 453. 

3 See Origin, Ed. i. pp. 280, 281, vi. p. 414. The author uses his 
experience of pigeons for examples for what he means by intermediate ; the 
instance of the horse and tapir also occurs. 



bat 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 ex- 
isting 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 1 . 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 be- 
tween the classes (but organisms from those coun- 
tries most separated by space also fall between the 
classes (e.g.) Ornithorhyncus ?). As far as geological 
discoveries (go) they tend towards such gradation 2 . 
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 per- 
fect as cattle, that if, as many geologists seem to 

1 The absence of intermediate forms between living organisms (and also 
as regards fossils) is discussed in the Origin, Ed. i. pp. 279, 280, vi. p. 413. 
In the above discussion there is no evidence that the author felt this difficulty 
so strongly as it is expressed in the Origin, Ed. i. p. 299, as perhaps "the 
most obvious and gravest objection that can be urged against my theory." 
But in a rough summary written on the back of the penultimate page of 
the MS. he refers to the geological evidence : " Evidence, as far as it does 
go, is favourable, exceedingly incomplete, greatest difficulty on this theory. 
I am convinced not insuperable." Buckland's remarks are given in the 
Origin, Ed. i. p. 329, vi. p. 471. 

' That the evidence of geology, as far as it goes, is favourable to the 
theory of descent is claimed in the Origin, Ed. i. pp. 343 345, vi. pp. 490 
492. For the reference to net in the following sentence, see Note 1, p. 48, 
of this Essay. 


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 ; bat 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 pre- 
servation, not sand and pebble) 1 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 favour- 
able, accords with facts of European deposits 2 , but 
subsidence apt to destroy agents which produce 
sediment 3 . 

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 

1 See Origin, Ed. i. p. 288, vi. p. 422. " The remains that do become 
embedded, if in sand and gravel, will, when the beds are upraised, generally 
be dissolved by the percolation of rain-water." 

2 The position of the following is not clear : " Think of immense 
differences in nature of European deposits, without interposing new 
causes, think of time required by present slow changes, to cause, on very 
same area, such diverse deposits, iron-sand, chalk, sand, coral, clay ! " 

3 The paragraph which ends here is difficult to interpret. In spite of 
obscurity it is easy to recognize the general resemblance to the discussion 
on the importance of subsidence given in the Origin, Ed. i. pp. 290 et seq., 
vi. pp. 422 et seq. 


past ages mere [gaps] pages preserved 1 . Ly ell'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 2 ? We know then cattle 
existed. Compare number of living, immense dura- 
tion 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 for- 
mation 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 sud- 
denly, but in [later] tertiary not quite suddenly 3 , 
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 im- 
mensely different. Most geologists believe Silurian 4 
fossils are those which first existed in the whole world, 

1 See Note 3, p. 27. 

2 Compare Origin, Ed. i. p. 298, vi. p. 437. "We shall, perhaps, best 
perceive the improbability of our being enabled to connect species by 
numerous, fine, intermediate, fossil links, by asking ourselves whether,' for 
instance, geologists at some future period will be able to prove that our 
different breeds of cattle, sheep, horses, and dogs have descended from a 
single stock or from several aboriginal stocks." 

! The sudden appearance of groups of allied species in the lowest known 
fossiliferous strata is discussed in the Origin, Ed. i. p. 306, vi. p. 446. The 
gradual appearance in the later strata occurs in the Origin, Ed. i. 
p. 312, vi. p. 453. 

4 Compare Origin, Ed. i. p. 307, vi. p. 448. 


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 Button 
or Lyell : if first reptile 1 of Red Sandstone (?) really 
was first which existed : if Pachyderm 2 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 intro- 
duction (?) 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 3 . 

1 I have interpreted as Sandstone a scrawl which I first read as Sea ; 
I have done so at the suggestion of Professor Judd, who points out that 
" footprints in the red sandstone were known at that time, and geologists 
were not then particular to distinguish between Amphibians and Reptiles." 

2 This refers to Cuvier's discovery of Palceotherium &c. at Montmartre. 

3 This simile is more fully given in the Origin, Ed. i. p. 310, vi. p. 452. 
" For my part, following out Lyell's metaphor, I look at the natural 
geological record, as a history of the world imperfectly kept, and 
written in a changing dialect ; of this history we possess the last volume 
alone, relating only to two or three countries. Of this volume, only here 
and there a short chapter has been preserved ; and of each page, only here 
and there a few lines. Each word of the slowly-changing language, in 
which the history is supposed to be written, being more or less different 
in the interrupted succession of chapters, may repi'esent the apparently 
abruptly changed forms of life, entombed in our consecutive, but widely 
separated formations." Professor Judd has been good enough to point out 
to me, that Darwin's metaphor is founded on the comparison of geology to 
history in Ch. i. of the Principles of Geology, Ed. i. 1830, vol. i. pp. 1 4. 
Professor Judd has also called my attention to another passage, Principles, 
Ed. i. 1833, vol. iii. p. 33, when Lyell imagines an historian examining "two 
buried cities at the foot of Vesuvius, immediately superimposed upon each 


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 cir- 
cumstance 1 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 in- 
crease : fossils become rare : possibly sudden exter- 
mination 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. n. how species would 
probably be introduced and how such results accord 
with what is known. 

other." The historian would discover that the inhabitants of the lower 
town were Greeks while those of the upper one were Italians. But he 
would be wrong in supposing that there had been a sudden change from 
the Greek to the Italian language in Campania. I think it is clear that 
Darwin's metaphor is partly taken from this passage. See for instance (in 
the above passage from the Origin) such phrases as "history... written in a 
changing dialect" "apparently abruptly changed forms of life." The 
passage within [ ] in the above paragraph : "Lyell's views as far as 
they go &c.," no doubt refers, as Professor Judd points out, to Lyell not 
going so far as Darwin on the question of the imperfection of the geological 

1 On rarity and extinction see Origin, Ed. i. pp. 109, 319, vi. pp. 133, 461. 


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 sup- 
plied 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 


vi. Let us consider the absolute state of dis- 
tribution 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] divi- 
sions; 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 accord- 
ing to changes in condition 1 of different parts. But 
besides this, barriers of every kind seem to separate 

1 In the Origin, Ed. i. p. 346, vi. p. 493, the author begins his discussion 
on geographical distribution by minimising the effect of physical conditions. 
He lays great stress on the effect of barriers, as in the present Essay. 


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 1 . 

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 
(?) 2 : 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 3 .] 

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 4 of 
climate on a continent, the propagation of species 
would vary and adapt themselves to small changes 

1 Note in the original, "Would it be more striking if we took animals, 
take Rhinoceros, and study their habitats ?" 

2 Note by Mr A. R. Wallace. "The want of similarity referred to, is, 
between the mountains of Brazil and Guiana and those of the Andes. Also 
those of the Indian peninsula as compared with the Himalayas. In both 
cases there is continuous intervening land. 

"The islands referred to were, no doubt, the Galapagos for dissimilarity 
from S. America ; our own Islands as compared with Europe, and perhaps 
Java, for similarity with continental Asia." 

3 The arguments against multiple centres of creation are given in the 
Origin, Ed. i. p. 352, vi. p. 499. 

4 In the Orii/in, Ed. i. p. 366, vi. p. 516, the author does not give his 
views on the distribution of alpine plants as original but refers to Edward 
Forbes' work (GeoJog. Survey Memoirs, 1846). In his autobiography, 
Darwin refers to this. " I was forestalled" he says, "in only one important 
point, which my vanity has always made me regret." (Life and Letters, i. 
p. 88.) 


causing much extermination 1 . 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 crea- 
tionists. 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 accord- 
ance with past and extinct forms, but no persistent 
relation between areas and distribution, Geologico- 

1 (The following is written on the back of a page of the MS.) Discuss 
one or more centres of creation : allude strongly to facilities of dispersal and 
amount of geological change : allude to mountain-summits afterwards to 
be referred to. The distribution varies, as everyone knows, according to 
adaptation, explain going from N. to S. how we come to fresh groups of 
species in the same general region, but besides this we find difference, 
according to greatness of barriers, in greater proportion than can be well 
accounted for by adaptation. (On representive species see Origin, Ed. i. 
p. 349, vi. p. 496.) This very striking when we think of cattle of Pampas, 
plants (?) &c. &c. Then go into discussion ; this holds with 3 or 4 main 
divisions as well as the endless minor ones in each of these 4 great ones : in 
these I chiefly refer to mammalia &c. &c. The similarity of type, but not 
in species, in same continent has been much less insisted on than the 
dissimilarity of different great regions generically: it is more striking. 

(I have here omitted an incomprehensible sentence.) Galapagos Islands, 
Tristan d'Acunha, volcanic islands covered with craters we know lately did not 
support any organisms. How unlike these islands in nature to neighbouring 
lands. These facts perhaps more striking than almost any others. 
[Geology apt to affect geography therefore we ought to expect to find 
the above.] Geological-geographical distribution. In looking to past times 
we find Australia equally distinct. S. America was distinct, though with 
more forms in common. N. America its nearest neighbour more in common, 
in some respects more, in some less allied to Europe. Europe we find (?) 
equally European. For Europe is now part of Asia though not (illegible). 
Africa unknown, examples, Elephant, Rhinoceros, Hippopotamus, Hyaena. 
As geology destroys geography we cannot be surprised in going far back we 
find Marsupials and Edentata in Europe : but geology destroys geography. 


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 1 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 2 increase in propor- 
tion 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 3 . 
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 

1 Rincon in Spanish means a nook or corner, it is here probably used to 
mean a small farm. 

2 The following is written across the page : " No one would expect a set 
of similar varieties to be produced in the different countries, so species 

3 (The following passage seems to have been meant to follow here.) The 
parent of an organism, we may generally suppose to be in less favourable 
condition than the selected offspring and therefore generally in fewer 
numbers. (This is not borne out by horticulture, mere hypothesis ; as an 
organism in favourable conditions might by selection be adapted to still 
more favourable conditions.) 

Barrier would further act in preventing species formed in one part 
migrating to another part. 


occupy an island 1 . 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 2 , 
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 dis- 
tant island might ultimately meet and a continent 
formed between them. No one doubts continents 
formed by repeated elevations and depressions 3 . 
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 enumer- 

1 (The following notes occur on the back of the page.) Number of species 
not related to capabilities of the country : furthermore not always those best 
adapted, perhaps explained by creationists by changes and progress. (See 
p. 34, note 1.) 

Although creationists can, by help of geology, explain much, how 
can he explain the marked relation of past and present in same area, 
the varying relation in other cases, between past and present, the relation 
of different parts of same great area. If island, to adjoining continent, if 
quite different, on mountain summits, the number of individuals not 
being related to capabilities, or how &c. our theory, I believe, can throw 
much light and all facts accord. 

2 See Origin, Ed. i. p. 390, vi. p. 543. 

3 On oscillation see Origin, Ed. i. p. 291, vi. p. 426. 



ated 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 1 . 

1 (From the back of MS.) Effect of climate on stationary island and 
on continent, but continent once island. Moreover repeated oscillations 
fresh diffusion when non -united, then isolation, when rising again immigra- 
tion prevented, new habitats formed, new species, when united free immi- 
gration, hence uniform characters. Hence more forms (on ?) the island. 
Mountain summits. Why not true species. First let us recall in Part I, 
conditions of variation : change of conditions during several generations, 
and if frequently altered so much better [perhaps excess of food]. Secondly, 
continued selection [while in wild state]. Thirdly, isolation in all or nearly 
all, as well to recall advantages of. 

.n continent, if we look to terrestrial animal, long continued change 
t go on, which would only cause change in numerical number 
(? proportions) : if continued long enough might ultimately affect all, though 
to most continents (there is) chance of immigration. Some few of whole 
body of species must be long affected and entire selection working same 
way. But here isolation absent, without barrier, cut off such (illegible). We 
can see advantage of isolation. But let us take case of island thrown up 
by volcanic agency at some distances, here we should have occasional 
visitants, only in few numbers and exposed to new conditions and (illegible) 
more important, a quite new grouping of organic beings, which would 
open out new sources of subsistence, or (would) control (?) old ones. The 
number would be few, can old have the very best opportunity. (The con- 
quest of the indigenes by introduced organisms shows that the indigenes 
were not perfectly adapted, see Origin, Ed. i. p. 390.) Moreover as the 
island continued changing, continued slow changes, river, marshes, lakes, 
mountains &c. &c., new races as successively formed and a fresh occasional 

If island formed continent, some species would emerge and immigrate. 
Everyone admits continents. We can see why Galapagos and C. Verde 
differ (see Origin, Ed. i. p. 398)], depressed and raised. We can see from 
this repeated action and the time required for a continent, why many more 
forms than in New Zealand (see Origin, Ed. i. p. 389 for a comparison be- 
tween New Zealand and the Cape) no mammals or other classes (see however, 
Origin, Ed. i. p. 393 for the case of the frog). We can at once see how it 
comes when there has been an old channel of migration, Cordilleras ; we 
can see why Indian Asiatic Flora, [why species] having a wide range gives 
better chance of some arriving at new points and being selected, and 
adapted to new ends. I need hardly remark no necessity for change. 



Looking now to the affinities of organisms, with- 
out 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 1 , we have no means in lower 

Finally, as continent (most extinction (?) during formation of continent) 
is formed after repeated elevation and depression, and interchange of species 
we might foretell much extinction, and that the survivor would belong to 
same type, as the extinct, in same manner as different part of same continent, 
which were once separated by space as they are by time (see Origin, Ed. i. 
pp. 339 and 349). 

As all mammals have descended from one stock, we ought to expect that 
every continent has been at some time connected, hence obliteration of 
present ranges. I do not mean that the fossil mammifers found in S. 
America are the lineal successors (ancestors) of the present forms of S. 
America : for it is highly improbable that more than one or two cases (who 
will say how many races after Plata bones) should be found. I believe 
this from numbers, who have lived, mere (?) chance of fewness. Moreover 
in every case from very existence of genera and species only few at one time 
will leave progeny, under form of new species, to distant ages ; and the more 
distant the ages the fewer the progenitors. An observation may be here 
appended, bad chance of preservation on rising island, the nurseries of new 
species, appeal to experience (see Origin, Ed. i. p. 292). This observation 
may be extended, that in all cases, subsiding land must be, in early stages, 
less favourable to formation of new species ; but it will isolate them, and 
then if land recommences rising how favourable. As preoccupation is bar 
to diffusion to species, so would it be to a selected variety. But it would 
not be if that variety was better fitted to some not fully occupied station ; 
so during elevation or the formation of new stations, is scene for new 
species. But during elevation not favourable to preservation of fossil 
(except in caverns (?)) ; when subsidence highly favourable in early stages to 
preservation of fossils ; when subsidence, less sediment. So that our strata, 
as general rule will be the tomb of old species (not undergoing any change) 
when rising land the nursery. But if there be vestige will generally be 
preserved to future ages, the new ones will not be entombed till fresh subsi- 
dence supervenes. In this long gap we shall have no record : so that 
wonderful if we should get transitional forms. I do not mean every stage, 
for we cannot expect that, as before shown, until geologists will be prepared 
to say that although under unnaturally favourable condition we can trace 
in future ages short-horn and Herefordshire (see note 2, p. 26). 

1 After " organs " is inserted, apparently as an afterthought : " no, and 
instance metamorphosis, afterwards explicable." 


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 ivhole organization very strong, 
yet no relation 1 . 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 2 . In classifying varieties the 
same language is used and the same kind of 
division : here also (in pine-apple) 3 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 4 
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 

1 For analogical resemblances see Origin, Ed. i. p. 427, vi. p. 582. 

2 " Practically when naturalists are at work, they do not trouble them- 
selves about the physiological value of the characters.... If they find a 
character nearly uniform,... they use it as one of high value," Origin, Ed. i. 
p. 417, vi. p. 573. 

3 "We are cautioned... not to class two varieties of the pine-apple 
together, merely because their fruit, though the most important part, 
happens to be nearly identical," Origin, Ed. i. p. 423, vi. p. 579. 

4 The whole of this passage is obscure, but the text is quite clear, except 
for one illegible word. 


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 be- 
come 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 ex- 
tinct and all might become extinct, and then we 
should have (an) extinct genus; a case formerly 
mentioned, of which numerous cases occur in Palae- 
ontology. 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 1 , 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 2 . External con- 
ditions air, earth, water being same 3 on globe, and 
the communication not being perfect, organisms of 
widely different descent might become adapted to 

1 (The exact position of the following passage is uncertain :) "just as it is 
not likely every present breed of fancy birds and cattle will propagate, only 
some of the best." 

2 This suggests that the author was not far from the principle of diver- 
gence on which he afterwards laid so much stress. See Origin, Ed. i. 
p. Ill, vi. p. 134, also Life and Letters, i. p. 84. 

3 That is to say the same conditions occurring in different parts of 
the globe. 


the same end and then we should have cases of 
analogy 1 , [they might even tend to become numeri- 
cally representative]. From this often happening 
each of the great divisions of nature would have 
their representative eminently adapted to earth, to 
(air) 2 , to water, and to these in (illegible) and then 
these great divisions would show numerical relations 
in their classification. 



Nothing more wonderful in Nat. Hist, than look- 
ing 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 3 , having bones 4 with 
same name, we see there is some deep bond of union 
between them 5 , 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 6 . 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 

1 The position of the following is uncertain, " greyhound and racehorse 
have an analogy to each other." The same comparison occurs in the Origin, 
Ed. i. p. 427, vi. p. 583. 

2 Air is evidently intended; in the MS. water is written twice. 

3 Written between the lines occurs :" extend to birds and other 

4 Written between the lines occurs: "many bones merely represented." 

5 In the Origin, Ed. i. p. 434, vi. p. 595, the term morphology is taken 
as including unity of type. The paddle of the porpoise and the wing of 
the bat are there used as instances of morphological resemblance. 

c The sentence is difficult to decipher. 


selections adapted to various conditions. We know 
that proportion, size, shape of bones and their accom- 
panying 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 verte- 
brse 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 1 . The unity of type in past and 
present ages of certain great divisions thus un- 
doubtedly receives the simplest explanation. 

There is another class of allied and almost 
identical facts, admitted by the soberest physio- 
logists, [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 in- 
dividual, 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, germeii 
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 crusta- 
ceans 2 , and all insects are considered as metamor- 
phosed (limbs) and to see the series is to admit this 
phraseology. The skulls of the vertebrates are un- 
doubtedly composed of three metamorphosed verte- 
brae; thus we can understand the strange form of 

1 In the Origin, Ed. i. p. 436, vi. p. 598, the author speaks of the 
" general pattern " being obscured in the paddles of " extinct gigantic sea- 

a See Origin, Ed. i. p. 437, vi. p. 599. 


the separate bones which compose the casket holding 
man's brain. These 1 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 mon- 
strous 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 2 . For let us take case of 
Vertebrata, which if 3 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 vertebrae 
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 vertebrae. 

1 The following passage seems to have been meant to precede the 
sentence beginning " These facts " : " It is evident, that when in each 
individual species, organs are metamorph. a unity of type extends." 

2 This is, I believe, the first place in which the author uses the words 
" theory of descent." 

3 The sentence should probably run, "Let us take the case of the verte- 
brata : if we assume them to be descended from one parent, then by this 
theory they have been altered &c." 


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 mon- 
strosities &c. we should say, instead of there 
existing a unity of type, morphology 1 , as we do 
when we trace the head as being the vertebrse 
metamorphosed. Be it observed that Naturalists, 
as they use terms of affinity without attaching real 
meaning, here also they are obliged to use meta- 
morphosis, 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 pro- 
bable, 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 2 . 

1 That is " we should call it a morphological fact." 

2 In the Origin, Ed. i. p. 438, vi. p. 602, the author, referring to the 
expressions used by naturalists in regard to morphology and metamorphosis, 
says " On my view these terms may be used literally." 


(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 foetus 
passes \ 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 2 (illegible). It is 
not true that one passes through the form of a lower 
group, though no doubt fish more nearly related to 
foetal state 3 . 

This similarity at the earliest stage is remark- 
ably shown in the course of the arteries which 
become greatly altered, as foetus 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 4 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 5 . 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 

1 See Origin, Ed. i. p. 439, vi. p. 605. 

2 In the Origin, Ed. i. p. 440, vi. p. 606, the author argues that the 
"loop-like course of the arteries" in the vertebrate embryo has no 
direct relation to the conditions of existence. 

3 The following passages are written across the page : " They pass 
through the same phases, but some, generally called the higher groups, are 
further metamorphosed. 

? Degradation and complication ? no tendency to perfection. 
? Justly argued against Lamarck ? " 

4 An almost identical passage occurs in the Origin, Ed. i. p. 440, vi. 
p. 606. 

5 The following: "Deaths of brothers (when) old by same peculiar 
disease " which is written between the lines seems to have been a memo- 
randum which is expanded a few lines lower. I believe the case of the 
brothers came from Dr R. W. Darwin. 


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 l and we must suspect that at whatever time 
of life the alteration of fetus is effected, it tends 
to appear at same period. When we (see) a ten- 
dency 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 2 . 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 3 , 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 foetal state : and this, I think, 
throws light on above marvellous fact. In larvae, 
which have long life selection, perhaps, does 
much, in the pupa not so much 4 . There is no 

1 See the discussion to this effect in the Origin, Ed. i. pp. 443-4, vi. p. 
610. The author there makes the distinction between a cause affecting the 
germ-cell and the reaction occurring at a late period of life. 

2 Possibly the sentence was meant to end " is not visible till then." 

3 See Origin, Ed. i. pp. 444-5, vi. p. 611. The query appended to much 
less is justified, since measurement was necessary to prove that the grey- 
hound and bulldog puppies had not neai'ly acquired " their full amount of 
proportional difference." 

4 (The following discussion, from the back of the page, is in large measure 
the same as the text.) I think light can be thrown on these facts. From the 
following peculiarities being hereditary, [we know that some change in the 
germinal vesicle is effected, which will only betray itself years after] diseases 
man, goitre, gout, baldness, fatness, size, [longevity (illegible) time of 


object gained in varying form &c. of foetus (beyond 
certain adaptations to mother's womb) and there- 
fore 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 

reproduction, shape of horns, case of old brothers dying of same disease]. 
And we know that the germinal vesicle must have been affected, though no 
effect is apparent or can be apparent till years afterwards, no more 
apparent than when these peculiarities appear by the exposure of the full- 
grown individual. (That is, " the young individual is as apparently free from 
the hereditary changes which will appear later, as the young is actually free 
from the changes produced by exposure to certain conditions in adult life.") 
So that when we see a variety in cattle, even if the variety be due to act of 
reproduction, we cannot feel sure at what period this change became 
apparent. It may have been effected during early age of free life (or) fetal 
existence, as monsters show. From arguments before used, and crossing, 
we may generally suspect in germ ; but I repeat it does not follow, that the 
change should be apparent till life fully developed ; any more than fatness 
depending on heredity should be apparent during early childhood, still 
less during fetal existence. In case of horns of cattle, which when inherited 
must depend on germinal vesicle, obviously no effect till cattle full-grown. 
Practically it would appear that the [hereditary] peculiarities characterising 
our domestic races, therefore resulting from vesicle, do not appear with 
their full characters in very early states ; thus though two breeds of cows 
have calves different, they are not so different, grey-hound and bull-dog. 
And this is what is (to) be expected, for man is indifferent to characters of 
young animals and hence would select those full-grown animals which 
possessed the desirable characteristics. So that from mere chance we 
might expect that some of the characters would be such only as became 
fully apparent in mature life. Furthermore we may suspect it to be a law, 
that at whatever time a new character appears, whether from vesicle, or 
effects of external conditions, it would appear at corresponding time 
(see Origin, Ed. i. p. 444). Thus diseases appearing in old age produce 
children with d., early maturity, longevity, old men, brothers, of 
same disease young children of d. I said men do not select for 
quality of young, calf with big bullocks. Silk-worms, peculiarities which, 
appear in caterpillar state or cocoon state, are transmitted to corre- 
sponding states. The effect of this would be that if some peculiarity was 
bora in a young animal, but never exercised, it might be inherited in young 
animal ; but if exercised that part of structure would be increased and 
would be inherited in corresponding time of life after such training. 

I have said that man selects in full-life, so would it be in Nature. In 
struggle of existence, it matters nothing to a feline animal, whether kitten 
eminently feline, as long as it sucks. Therefore natural selection would act 
equally well on character which was fully (developed) only in full age. 
Selection could tend to alter no character in foetus, (except relation to 
mother) it would alter less in young state (putting on one side larva condition) 
but alter every part in full-grown condition. Look to a fetus and its parent, 
and again after ages fetus and its (i.e. the above mentioned parents) 
descendant ; the parent more variable (?) than fetus, which explains all. 


the arteries, as long as they nourish the foetus; it 
is the selection of slight changes which supervene at 
any time during (illegible) of life. 

The less differences of foetus, 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 foetus, retaining traces of the ancestral form, 
is of the highest value in classification. 


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 1 , 
whale, narwhal, bone on tibia, muscles which do not 
move, little bone of wing of Apteryx, bone repre- 
senting extremities in some snake, little wings 
within (?) soldered cover of beetles, men and 
bulls, mammae : 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 2 . 

The term abortive organ has been thus applied 

1 Some of these examples occur in Origin, Ed. i. pp. 450-51, vi. 
pp. 619-20. 

2 The two following sentences are written, one down the margin, the 
other across the page. " Abortive organs eminently useful in classification. 
Embryonic state of organs. Rudiments of organs." 


to above structure (as invariable as all other parts 1 ) 
from their absolute similarity to monstrous cases, 
where from accident, certain organs are not de- 
veloped; 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 2 . 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 Linuinflavum, stamen in 
Geranium pyrenaicwm?. Nectaries abort into petals 
in Columbine (Aquilegia], produced from some acci- 
dent 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 4 ?) muscles of dog's ears, (and 
of) rabbits, muscles wither, arteries grow up. When 
eye born defective, optic nerve (Tuco Tuco) is atro- 
phied. 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 

1 I imagine the meaning to be that abortive organs are specific characters 
in contrast to monstrosities. 

2 Minute hanging horns are mentioned in the Origin, Ed. i. p. 454, vi. p. 
625, as occurring in hornless breeds of cattle. 

3 Linum flacum is dimorphic : thyme gynodisecious. It is not clear 
what point is referred to under Geranium pyrenaicum. 

4 The author's work on duck's wings &c. is in Var. under Dom., Ed. 2, 
i. p. 299. 


hereditariness 1 . 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 anti- 
cipated, 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 2 , indeed in one point of view, 
the heads of [vertebrated] animal may be said to be 
abortive vertebrae turned into other use: legs of 
some Crustacea abortive jaws, &c., &c. De Candolle's 
analogy of table covered with dishes 3 . 

(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. 

1 The words vis medicatrix are inserted after " useless," apparently as a 

2 In the male florets of certain Composites the style functions merely as 
a piston for forcing out the pollen. 

3 (On the back of the page is the following.) If abortive organs are a 
trace preserved by hereditary tendency, of organ in ancestor of use, we can 
at once see why important in natural classification, also why more plain in 
young animal because, as in last section, the selection has altered the old 
animal most. I repeat, these wondrous facts, of parts created for no use in 
past and present time, all can by my theory receive simple explanation ; or 
they receive none and we must be content with some such empty metaphor, 
as that of De Candolle, who compares creation to a well covered table, and 
says abortive organs may be compared to the dishes (some should be empty) 
placed symmetrically ! 



Let us recapitulate the whole (?) (of) these latter 
sections by taking case of the three species of 
Rhinoceros, which inhabit Java, Sumatra, and main- 
land 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] 1 set of links in the broken chain 
representing the Pachydermata, as the chain like- 
wise 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 vertebrae, short neck, same bones as giraffe. 
In the upper jaws of all three we find small teeth 
like rabbit's. In dissecting them in foetal 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, 

1 The author doubtless meant that the complex relationships between 
organisms can be roughly represented by a net in which the knots stand 
for species. 


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 1 different (from) their aboriginal 
forms; might be selected to serve different ends. 

Now the Creationist believes these three Rhi- 
noceroses were created 2 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, differ- 
ently adapted, now inhabiting different countries, 
with different structures and instincts, are ad- 
mitted 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 im- 
perfect 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 3 . But 
there remains one other way of comparing species 

1 Between the lines occurs : " one (?) form be lost." 

2 The original sentence is here broken up by the insertion of: "out of 
the dust of Java, Sumatra, these (?) allied to past and present age and 
(illegible), with the stamp of inutility in some of their organs and conversion 
in others." 

3 Between the lines occur the words : " Species vary according to same 
general laws as varieties ; they cross according to same laws." 



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 1 . 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 Kolreuter 
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 ! 


Such are my reasons for believing that specific 
forms are not immutable. The affinity of different 
groups, the unity of types of structure, the repre- 
sentative forms through which foetus 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 2 , or other great work of 
art, as a thing wholly beyond comprehension, but we 

1 " A cross with a bull-dog has affected for many generations the courage 
and obstinacy of greyhounds," Origin, Ed. i. p. 214, vi. p. 327. 

2 The simile of the savage and the ship occurs in the Origin, Ed. i. 
p. 485, vi. p. 665. 


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feel far more interest in examining it. How in- 
teresting 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 sum- 
mary of a long history, (as the summing up) of 1 useful 
contrivances, much like a work of art. How in- 
teresting 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 2 , 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 3 . 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 

1 In the Origin, Ed. i. p. 486, vi. p. 665, the author speaks of the " sum- 
ming up of many contrivances " : I have therefore introduced the above 
words which make the passage clearer. In the Origin the comparison is 
with "a great mechanical invention," not with a work of art. 

2 See a similar passage in the Origin, Ed. i. p. 487, vi. p. 667. 

3 See the Origin, Ed. i. p. 488, vi. p. 668. 


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 con- 
ceive, the creation of the higher animals has directly 
come. Doubtless it at first transcends our humble 
powers, to conceive laws capable of creating in- 
dividual organisms, each characterised by the most 
exquisite workmanship and widely- extended adap- 
tations. 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 1 . 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 2 . 

1 The following discussion, together with some memoranda are on the 
last page of the MS. " The supposed creative spirit does not create either 
number or kind which (are) from analogy adapted to site (viz. New Zealand) : 
it does not keep them all permanently adapted to any country, it works on 
spots or areas of creation, it is not persistent for great periods, it creates 
forms of same groups in same regions, with no physical similarity, it 
creates, on islands or mountain summits, species allied to the neighbouring 
ones, and not allied to alpine nature as shown in other mountain summits 
even different on different island of similarly constituted archipelago, not 
created on two points : never mammifers created on small isolated island ; 
nor number of organisms adapted to locality : its power seems influenced or 
related to the range of other species wholly distinct of the same genus, it 
does not equally effect, in amount of difference, all the groups of the same 

2 This passage is the ancestor of the concluding words in the first edition 
of the Origin of Species which have remained substantially unchanged 
throughout subsequent editions, "There is grandeur in this view of life, 
with its several powers, having been originally breathed into a few forms 
or into one ; and that whilst this planet has gone cycling on according to the 
fixed law of gravity, from so simple a beginning endless forms most 


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 -ac- 
climatisation 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 per- 
ceiving every step, like the erosion of valleys 1 . 

beautiful and most wonderful have been, and are being, evolved." In the 
2nd edition " by the Creator " is introduced after " originally breathed." 

1 Compare the Origin, Ed. i. p. 481, vi. p. 659, "The difficulty is the 
same as that felt by so many geologists, when Lyell first insisted that long 
lines of inland cliffs had been formed, and great valleys excavated, by the 
slow action of the coast-waves." 

. a 

9 V