On the poorness of our palaeontological collections -- On the intermittence
of geological formations -- On the absence of intermediate varieties in any
one formation -- On the sudden appearance of groups of species -- On their
sudden appearance in the lowest known fossiliferous strata.
Chapter X
On the Geological Succession of Organic Beings
On the slow and successive appearance of new species -- On their different
rates of change -- Species once lost do not reappear -- Groups of species
follow the same general rules in their appearance and disappearance as do
single species -- On Extinction -- On simultaneous changes in the forms of
life throughout the world -- On the affinities of extinct species to each
other and to living species -- On the state of development of ancient forms
-- On the succession of the same types within the same areas -- Summary of
preceding and present chapters.
Chapter XI
Geographical Distribution
Present distribution cannot be accounted for by differences in physical
conditions -- Importance of barriers -- Affinity of the productions of the
same continent -- Centres of creation -- Means of dispersal, by changes of
climate and of the level of the land, and by occasional means -- Dispersal
during the Glacial period co-extensive with the world.
Chapter XII
Geographical Distribution -- continued
Distribution of fresh-water productions -- On the inhabitants of oceanic
islands -- Absence of Batrachians and of terrestrial Mammals -- On the
relation of the inhabitants of islands to those of the nearest mainland --
On colonisation from the nearest source with subsequent modification --
Summary of the last and present chapters.
Chapter XIII
Mutual Affinities of Organic Beings:
Morphology: Embryology: Rudimentary Organs
Classification, groups subordinate to groups -- Natural system -- Rules and
difficulties in classification, explained on the theory of descent with
modification -- Classification of varieties -- Descent always used in
classification -- Analogical or adaptive characters -- Affinities, general,
complex and radiating -- Extinction separates and defines groups --
Morphology, between members of the same class, between parts of the same
individual -- Embryology, laws of, explained by variations not supervening
at an early age, and being inherited at a corresponding age -- Rudimentary
Organs; their origin explained -- Summary.
Chapter XIV
Recapitulation and Conclusion
Recapitulation of the difficulties on the theory of Natural Selection --
Recapitulation of the general and special circumstances in its favour --
Causes of the general belief in the immutability of species -- How far the
theory of natural selection may be extended -- Effects of its adoption on
the study of Natural history -- Concluding remarks.
On the Origin of Species.
Introduction.
When on board H.M.S. 'Beagle,' as naturalist, I was much struck with
certain facts in the distribution of the inhabitants of South America, and
in the geological relations of the present to the past inhabitants of that
continent. These facts seemed to me to throw some light on the origin of
species--that mystery of mysteries, as it has been called by one of our
greatest philosophers. On my return home, it occurred to me, in 1837, that
something might perhaps be made out on this question by patiently
accumulating and reflecting on all sorts of facts which could possibly have
any bearing on it. After five years' work I allowed myself to speculate on
the subject, and drew up some short notes; these I enlarged in 1844 into a
sketch of the conclusions, which then seemed to me probable: from that
period to the present day I have steadily pursued the same object. I hope
that I may be excused for entering on these personal details, as I give
them to show that I have not been hasty in coming to a decision.
My work is now nearly finished; but as it will take me two or three more
years to complete it, and as my health is far from strong, I have been
urged to publish this Abstract. I have more especially been induced to do
this, as Mr. Wallace, who is now studying the natural history of the Malay
archipelago, has arrived at almost exactly the same general conclusions
that I have on the origin of species. Last year he sent to me a memoir on
this subject, with a request that I would forward it to Sir Charles Lyell,
who sent it to the Linnean Society, and it is published in the third volume
of the Journal of that Society. Sir C. Lyell and Dr. Hooker, who both knew
of my work--the latter having read my sketch of 1844--honoured me by
thinking it advisable to publish, with Mr. Wallace's excellent memoir, some
brief extracts from my manuscripts.
This Abstract, which I now publish, must necessarily be imperfect. I
cannot here give references and authorities for my several statements; and
I must trust to the reader reposing some confidence in my accuracy. No
doubt errors will have crept in, though I hope I have always been cautious
in trusting to good authorities alone. I can here give only the general
conclusions at which I have arrived, with a few facts in illustration, but
which, I hope, in most cases will suffice. No one can feel more sensible
than I do of the necessity of hereafter publishing in detail all the facts,
with references, on which my conclusions have been grounded; and I hope in
a future work to do this. For I am well aware that scarcely a single point
is discussed in this volume on which facts cannot be adduced, often
apparently leading to conclusions directly opposite to those at which I
have arrived. A fair result can be obtained only by fully stating and
balancing the facts and arguments on both sides of each question; and this
cannot possibly be here done.
I much regret that want of space prevents my having the satisfaction of
acknowledging the generous assistance which I have received from very many
naturalists, some of them personally unknown to me. I cannot, however, let
this opportunity pass without expressing my deep obligations to Dr. Hooker,
who for the last fifteen years has aided me in every possible way by his
large stores of knowledge and his excellent judgment.
In considering the Origin of Species, it is quite conceivable that a
naturalist, reflecting on the mutual affinities of organic beings, on their
embryological relations, their geographical distribution, geological
succession, and other such facts, might come to the conclusion that each
species had not been independently created, but had descended, like
varieties, from other species. Nevertheless, such a conclusion, even if
well founded, would be unsatisfactory, until it could be shown how the
innumerable species inhabiting this world have been modified, so as to
acquire that perfection of structure and coadaptation which most justly
excites our admiration. Naturalists continually refer to external
conditions, such as climate, food, &c., as the only possible cause of
variation. In
one very limited sense, as we shall hereafter see, this may
be true; but it is preposterous to attribute to mere external conditions,
the structure, for instance, of the woodpecker, with its feet, tail, beak,
and tongue, so admirably adapted to catch insects under the bark of trees.
In the case of the misseltoe, which draws its nourishment from certain
trees, which has seeds that must be transported by certain birds, and which
has flowers with separate sexes absolutely requiring the agency of certain
insects to bring pollen from one flower to the other, it is equally
preposterous to account for the structure of this parasite, with its
relations to several distinct organic beings, by the effects of external
conditions, or of habit, or of the volition of the plant itself.
The author of the 'Vestiges of Creation' would, I presume, say that, after
a certain unknown number of generations, some bird had given birth to a
woodpecker, and some plant to the misseltoe, and that these had been
produced perfect as we now see them; but this assumption seems to me to be
no explanation, for it leaves the case of the coadaptations of organic
beings to each other and to their physical conditions of life, untouched
and unexplained.
It is, therefore, of the highest importance to gain a clear insight into
the means of modification and coadaptation. At the commencement of my
observations it seemed to me probable that a careful study of domesticated
animals and of cultivated plants would offer the best chance of making out
this obscure problem. Nor have I been disappointed; in this and in all
other perplexing cases I have invariably found that our knowledge,
imperfect though it be, of variation under domestication, afforded the best
of such studies, although they have been very commonly neglected by
naturalists.
From these considerations, I shall devote the first chapter of this
Abstract to Variation under Domestication. We shall thus see that a large
amount of hereditary modification is at least possible, and, what is
equally or more important, we shall see how great is the power of man in
accumulating by his Selection successive slight variations. I will then
pass on to the variability of species in a state of nature; but I shall,
unfortunately, be compelled to treat this subject far too briefly, as it
can be treated properly only by giving long catalogues of facts. We shall,
however, be enabled to discuss what circumstances are most favourable to
variation. In the next chapter the Struggle for Existence amongst all
organic beings throughout the world, which inevitably follows from their
high geometrical powers of increase, will be treated of. This is the
doctrine of Malthus, applied to the whole animal and vegetable kingdoms.
As many more individuals of each species are born than can possibly
survive; and as, consequently, there is a frequently recurring struggle for
existence, it follows that any being, if it vary however slightly in any
manner profitable to itself, under the complex and sometimes varying
conditions of life, will have a better chance of surviving, and thus be
naturally selected. From the strong principle of inheritance, any selected
variety will tend to propagate its new and modified form.
This fundamental subject of Natural Selection will be treated at some
length in the fourth chapter; and we shall then see how Natural Selection
almost inevitably causes much Extinction of the less improved forms of life
and induces what I have called Divergence of Character. In the next
chapter I shall discuss the complex and little known laws of variation and
of correlation of growth. In the four succeeding chapters, the most
apparent and gravest difficulties on the theory will be given: namely,
first, the difficulties of transitions, or in understanding how a simple
being or a simple organ can be changed and perfected into a highly
developed being or elaborately constructed organ; secondly the subject of
Instinct, or the mental powers of animals, thirdly, Hybridism, or the
infertility of species and the fertility of varieties when intercrossed;
and fourthly, the imperfection of the Geological Record. In the next
chapter I shall consider the geological succession of organic beings
throughout time; in the eleventh and twelfth, their geographical
distribution throughout space; in the thirteenth, their classification or
mutual affinities, both when mature and in an embryonic condition. In the
last chapter I shall give a brief recapitulation of the whole work, and a
few concluding remarks.
No one ought to feel surprise at much remaining as yet unexplained in
regard to the origin of species and varieties, if he makes due allowance
for our profound ignorance in regard to the mutual relations of all the
beings which live around us. Who can explain why one species ranges widely
and is very numerous, and why another allied species has a narrow range and
is rare? Yet these relations are of the highest importance, for they
determine the present welfare, and, as I believe, the future success and
modification of every inhabitant of this world. Still less do we know of
the mutual relations of the innumerable inhabitants of the world during the
many past geological epochs in its history. Although much remains obscure,
and will long remain obscure, I can entertain no doubt, after the most
deliberate study and dispassionate judgment of which I am capable, that the
view which most naturalists entertain, and which I formerly
entertained--namely, that each species has been independently created--is
erroneous. I am fully convinced that species are not immutable; but that
those belonging to what are called the same genera are lineal descendants
of some other and generally extinct species, in the same manner as the
acknowledged varieties of any one species are the descendants of that
species. Furthermore, I am convinced that Natural Selection has been the
main but not exclusive means of modification.
Chapter I
Variation under Domestication
Causes of Variability -- Effects of Habit -- Correlation of Growth --
Inheritance -- Character of Domestic Varieties -- Difficulty of
Varieties from one or more Species -- Domestic Pigeons, their Differences
and Origin -- Principle of Selection anciently followed, its Effects --
Methodical and Unconscious Selection -- Unknown Origin of our Domestic
Productions -- Circumstances favourable to Man's power of Selection.
When we look to the individuals of the same variety or sub-variety of our
older cultivated plants and animals, one of the first points which strikes
us, is, that they generally differ much more from each other, than do the
individuals of any one species or variety in a state of nature. When we
reflect on the vast diversity of the plants and animals which have been
cultivated, and which have varied during all ages under the most different
climates and treatme
nt, I think we are driven to conclude that this greater
variability is simply due to our domestic productions having been raised
under conditions of life not so uniform as, and somewhat different from,
those to which the parent-species have been exposed under nature. There
is, also, I think, some probability in the view propounded by Andrew
Knight, that this variability may be partly connected with excess of food.
It seems pretty clear that organic beings must be exposed during several
generations to the new conditions of life to cause any appreciable amount
of variation; and that when the organisation has once begun to vary, it
generally continues to vary for many generations. No case is on record of
a variable being ceasing to be variable under cultivation. Our oldest
cultivated plants, such as wheat, still often yield new varieties: our
oldest domesticated animals are still capable of rapid improvement or
modification.
It has been disputed at what period of life the causes of variability,
whatever they may be, generally act; whether during the early or late
period of development of the embryo, or at the instant of conception.
Geoffroy St. Hilaire's experiments show that unnatural treatment of the
embryo causes monstrosities; and monstrosities cannot be separated by any
clear line of distinction from mere variations. But I am strongly inclined
to suspect 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 conception. Several reasons make me believe in this; but the chief
one is the remarkable effect which confinement or cultivation has on the
functions of the reproductive system; this system appearing to be far more
susceptible than any other part of the organisation, to the action of any
change in the conditions of life. Nothing is more easy than to tame an
animal, and few things more difficult than to get it to breed freely under
confinement, even in the many cases when the male and female unite. How
many animals there are which will not breed, though living long under not
very close confinement in their native country! This is generally
attributed to vitiated instincts; but how many cultivated plants display
the utmost vigour, and yet rarely or never seed! In some few such cases it
has been found out that very trifling changes, such as a little more or
less water at some particular period of growth, will determine whether or
not the plant sets a seed. I cannot here enter on the copious details
which I have collected on this curious subject; but to show how singular
the laws are which determine the reproduction of animals under confinement,
I may just mention that carnivorous animals, even from the tropics, breed
in this country pretty freely under confinement, with the exception of the
plantigrades or bear family; whereas, carnivorous birds, with the rarest
exceptions, hardly ever lay fertile eggs. Many exotic plants have pollen
utterly worthless, in the same exact condition as in the most sterile
hybrids. When, on the one hand, we see domesticated animals and plants,
though often weak and sickly, yet breeding quite freely under confinement;
and when, on the other hand, we see individuals, though taken young from a
state of nature, perfectly tamed, long-lived, and healthy (of which I could
give numerous instances), yet having their reproductive system so seriously
affected by unperceived causes as to fail in acting, we need not be
surprised at this system, when it does act under confinement, acting not
quite regularly, and producing offspring not perfectly like their parents
or variable.
Sterility has been said to be the bane of horticulture; but on this view we