Journal of researches during H. M. S. Beagle's Voyage round the world.
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
This Abstract, which I now publish, must necessarily be imperfect.
I cannot here give references and authorities for my 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,
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
co-adaptation 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 The author of the 'Vestiges of Creation' would, I presume, say
that, after a certain unknown number of
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 and safest clue. I
may venture to express my conviction of the high value 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 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 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
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
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 treatment, 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.
It has been disputed at what period of time the causes of Sterility has been said to be the bane of horticulture; but on this
view we owe variability to the same cause which produces sterility;
and variability is the source of all the choicest productions of the
garden. I may add, that as some organisms will breed most freely under
the most unnatural conditions (for instance, the rabbit and ferret
kept in hutches), showing that their reproductive system has not been
thus affected; so will some animals and plants withstand domestication
or cultivation, and vary very slightly -- perhaps hardly more
than in a state of nature.
A long list could easily be given of 'sporting plants;' by this
term gardeners mean a single bud or offset, which suddenly assumes a
new and sometimes very different character from that of the rest of
the plant.
Seedlings from the same fruit, and the young of the same litter,
sometimes differ considerably from each other, though both the young
and the parents, as Muller has remarked, have apparently been exposed
to exactly the same conditions of life; and this shows how unimportant
the direct effects of the conditions of life are in comparison with
the laws of reproduction, and of growth, and of inheritance; for had
the action of the conditions been direct, if any of the young had
varied, all would probably have varied in the same manner. To judge
how much, in the case Habit also has a deciding influence, as in the period of flowering
with plants when transported from one climate to another. In animals
it has a more marked effect; for instance, I find in the domestic duck
that the bones of the wing weigh less and the bones of the leg more,
in proportion to the whole skeleton, than do the same bones in the
wild-duck; and I presume that this change may be safely attributed to
the domestic duck flying much less, and walking more, than its wild
parent. The great and inherited development of the udders in cows and
goats in countries where they are habitually milked, in comparison
with the state of these organs in other countries, is another instance
of the effect of use. Not a single domestic animal can be named which
has not in some country drooping ears; and the view suggested by some
authors, that the drooping is due to the disuse of the muscles of the
ear, from the animals not being much alarmed by danger, seems
probable.
There are many laws regulating variation, some few of which can be
dimly seen, and will be hereafter briefly mentioned. I will here only
allude to what may be called correlation of growth. Any change in the
embryo or larva will almost certainly entail changes in the mature
animal. In monstrosities, the correlations between quite distinct
parts are very curious; and many instances are given in Isidore
Geoffroy St Hilaire's great work on this subject. Breeders believe
that long limbs are almost always The result of the various, quite unknown, or dimly seen laws of
variation is infinitely complex and diversified. It is well worth
while carefully to study the several treatises published on some of
our old cultivated plants, as on the hyacinth, potato, even the
dahlia, &c.; and it is really surprising to note the endless
points in structure and constitution in which the varieties and sub
varieties differ slightly from each other. The whole organization
seems to have become plastic, and tends to depart in some small degree
from that of the parental type.
Any variation which is not inherited is unimportant for us. But the
number and diversity of inheritable deviations of structure, both
those of slight and those of considerable physiological importance, is
endless. Dr Prosper Lucas's treatise, in two large volumes, is the
fullest and the best on this subject. No breeder doubts how strong is
the tendency to inheritance: like produces like is his fundamental
belief: doubts have been thrown on this principle by theoretical
writers alone. When a
The laws governing inheritance are quite unknown; no one can say
why the same peculiarity in different individuals of the same species,
and in individuals of different species, is sometimes inherited and
sometimes not so; why the child often reverts in certain characters to
its grandfather or grandmother or other much more remote ancestor; why
a peculiarity is often transmitted from one sex to both sexes or to
one sex alone, more commonly but not exclusively to the like sex. It
is a fact of some little importance to us, that peculiarities
appearing in the males of our domestic breeds are often transmitted
either exclusively, or in a much greater degree, to males alone. A
much more important rule, which I think may be trusted, is that, at
whatever period of life a peculiarity first appears, it tends to
appear in the offspring at a corresponding age, though sometimes
earlier. In many cases this could
Having alluded to the subject of reversion, I may here refer to a
statement often made by naturalists -- namely, that our domestic
varieties, when run wild, gradually but certainly revert in character
to their aboriginal stocks. Hence it has been argued that no
deductions can be drawn from domestic races to species in a state of
nature. I have in vain endeavoured to discover on what decisive facts
the above statement has so often and so boldly been made. There would
be great difficulty in proving its truth: we may safely conclude that
very many of the most strongly-marked domestic varieties could not
possibly live in a wild state. In many cases we do not know what the
aboriginal stock was, and so could not tell whether or not nearly
perfect reversion had ensued. It would be quite necessary, in order to
prevent the effects of intercrossing, that only a
When we look to the hereditary varieties or races of our domestic
animals and plants, and compare them with species closely allied
together, we generally perceive in each domestic race, as already
remarked, less uniformity of character than in true species. Domestic
races of
When we attempt to estimate the amount of structural difference
between the domestic races of the same species, we are soon involved
in doubt, from not knowing whether they have It has often been assumed that man has chosen for domestication
animals and plants having an extraordinary inherent tendency to vary,
and likewise to withstand diverse climates. I do not dispute that
these capacities have added largely to the value of most of our
domesticated productions; but how could a savage possibly know, when
he first tamed an animal, whether it would vary in succeeding
generations, and whether it would endure other climates? Has the
little variability of the ass or guinea-fowl, or the small power of
endurance of warmth by the reindeer, or of cold by the common camel,
prevented their domestication? I cannot doubt that if other animals
and plants, equal in number to our domesticated productions, and
belonging to equally diverse classes and countries, were taken from a
state of nature, and could be made to breed for an equal number of
generations under domestication, they would vary on an average as
largely as the parent species of our existing domesticated productions
have varied.
In the case of most of our anciently domesticated animals and
plants, I do not think it is possible to come to any definite
conclusion, whether they have descended from one or several species.
The argument mainly relied on by those who believe in the multiple
origin
The whole subject must, I think, remain vague; nevertheless, I may,
without here entering on any details, state that, from geographical
and other considerations, I think it highly probable that our domestic
dogs have descended from several wild species. In regard to sheep and
goats I can form no opinion. I should think, from facts communicated
to me by Mr Blyth, on the habits, voice, and constitution, &c., of
the humped Indian cattle, that these had descended from a different
aboriginal stock from our European cattle; and several competent
judges believe that these latter have had more than one wild parent.
With respect to horses, from reasons which I cannot give here, I am
doubtfully inclined to believe, in opposition to several authors, that
all the races have descended from one wild stock. Mr Blyth, whose
opinion, from his large and varied stores of knowledge, I should value
more than that of almost any one, thinks that all the breeds of
poultry have proceeded from the common wild
The doctrine of the origin of our several domestic races from
several aboriginal stocks, has been carried to an absurd extreme by
some authors. They believe that every race which breeds true, let the
distinctive characters be ever so slight, has had its wild prototype.
At this rate there must have existed at least a score of species of
wild cattle, as many sheep, and several goats in Europe alone, and
several even within Great Britain. One author believes that there
formerly existed in Great Britain eleven wild species of sheep
peculiar to it! When we bear in mind Believing that it is In the skeletons of the several breeds, the development of the
bones of the face in length and breadth and curvature differs
enormously. The shape, as well as the breadth and length of the ramus
of the lower jaw, varies in a highly remarkable manner. The number of
the caudal and sacral vertebrae vary; as does the number of the ribs,
together with their relative breadth and the presence of processes.
The size and shape of the apertures in the sternum are highly
variable; so is the degree of divergence and relative size of the two
arms of the furcula. The proportional width of the gape of mouth, the
proportional length of the eyelids, of the orifice of the nostrils, of
the tongue (not always in strict correlation with the length of beak),
the size of the crop and of the upper part of the oesophagus; the
development and abortion of the oil-gland; the number of the primary
wing and caudal feathers; the relative length of wing and tail to each
other and to the body; the relative length of leg and of the feet; the
number of scutellae on the toes, the development of skin between the
toes, are all points of structure which are variable. The period at
which the perfect plumage is acquired varies, as does the state of the
down with which the nestling birds are clothed when hatched. The shape
and size of the eggs vary. The manner of flight differs remarkably; as
does in some breeds the voice and disposition. Lastly, in certain
breeds, the males and females have come to differ to a slight degree
from each other.
Altogether at least a score of pigeons might be chosen, which if
shown to an ornithologist, and he were told that they were wild birds,
would certainly, I think, be ranked by him as well-defined species.
Moreover, I do not believe that any ornithologist would place
Great as the differences are between the breeds of pigeons, I am
fully convinced that the common opinion of naturalists is correct,
namely, that all have descended from the rock-pigeon An argument, as it seems to me, of great weight, and applicable in
several other cases, is, that the above-specified breeds, though
agreeing generally in constitution, habits, voice, colouring, and in
most parts of their structure, with the wild rock-pigeon, yet are
certainly highly abnormal in other parts of their structure: we may
look in vain throughout the whole great family of Columbidae for a
beak like that of the English carrier, or that of the short-faced
tumbler, or barb; for reversed feathers like those of the jacobin; for
a crop like that of the pouter; for tail-feathers like those of the
fantail. Hence it must be assumed not only that half-civilized man
succeeded in thoroughly domesticating several species, but that he
intentionally or by chance picked out extraordinarily abnormal
species; and further, that these very species have since all become
extinct or unknown. So many strange contingencies seem to me
improbable in the highest degree.
Some facts in regard to the colouring of pigeons well deserve
consideration. The rock-pigeon is of a slaty-blue, and has a white
rump (the Indian sub-species, C. intermedia of Strickland, having it
bluish); the tail has a terminal dark bar, with the bases of the outer
feathers externally edged with white; the wings have two black bars:
some semi-domestic breeds and some apparently truly wild breeds have,
besides the two black bars, the wings chequered with black. These
several marks do not occur together in any other species of the whole
family. Now, in every one of the domestic breeds, taking thoroughly
well-bred birds, all the above marks, even to the white edging of the
outer tail-feathers, sometimes concur perfectly developed. Moreover,
when two birds belonging to two distinct breeds are crossed, neither
of which is blue or has any of the above-specified marks, the mongrel
offspring are very apt suddenly to acquire these characters; for
instance, I crossed some uniformly white fantails with some uniformly
black barbs, and they produced mottled brown and black birds; these I
again crossed together, and one grandchild of the pure white fantail
and pure black barb was of as beautiful a blue colour, with the white
rump, double black wing-bar, and barred and white-edged tail-feathers,
as any wild rock-pigeon! We can understand these facts, on the
well-known principle of Lastly, the hybrids or mongrels from between all the domestic
breeds of pigeons are perfectly fertile. I can state this from my own
observations, purposely made on the most distinct breeds. Now, it is
difficult, perhaps impossible, to bring forward one case of the hybrid
offspring of two animals From these several reasons, namely, the improbability of man In favour of this view, I may add, firstly, that C. livia, or the
rock-pigeon, has been found capable of domestication in Europe and in
India; and that it agrees in habits and in a great number of points of
structure with all the domestic breeds. Secondly, although an English
carrier or short-faced tumbler differs immensely in certain characters
from the rock-pigeon, yet by comparing the several sub-breeds of these
breeds, more especially those brought from distant countries, we can
make an almost perfect series between the extremes of structure.
Thirdly, those characters which are mainly distinctive of each breed,
for instance the wattle and length of beak of the carrier, the
shortness of that of the tumbler, and the number of tail-feathers in
the fantail, are in each breed eminently variable; and the explanation
of this fact will be obvious when we come to treat of selection.
Fourthly, pigeons have been watched, and tended with the utmost care,
and loved by many people. They have been domesticated for thousands of
years in several quarters of the world; the earliest known record of
pigeons is in the fifth &Aelig;gyptian dynasty, about 3000 B.C., as was
pointed out to me by Professor Lepsius; but Mr Birch informs me that
pigeons are given in a bill
I have discussed the probable origin of domestic pigeons at some,
yet quite insufficient, length; because when I first kept pigeons and
watched the several kinds, knowing well how true they bred, I felt
fully as much difficulty in believing that they could ever have
descended from a common parent, as any naturalist could in coming to a
similar conclusion in regard to the many species of finches, or other
large groups of birds, in nature. One circumstance has struck me much;
namely, that all the breeders of the various domestic animals and the
cultivators of plants, with whom I have ever conversed, or whose
treatises I have read, are firmly convinced that the several breeds to
which each has attended, are descended from so many aboriginally
distinct species.
Let us now briefly consider
the steps by which domestic races have been produced, either from one
or from several allied species. Some little effect may, perhaps, be
attributed to the direct action of the external conditions of life,
and some little to habit; but he would be a bold man who would account
by such agencies for the differences of a dray and race horse, a
greyhound and bloodhound, a carrier and tumbler pigeon. One of the
most remarkable features in our domesticated races
The great power of this principle of selection is not hypothetical.
It is certain that several of our eminent breeders have, even within a
single lifetime, modified to
What English breeders have actually effected is proved by the
enormous prices given for animals with a good pedigree; and these have
now been exported to almost every quarter of the world. The
improvement is by no means generally due to crossing different breeds;
The same principles are followed by horticulturists; but the
variations are here often more abrupt. No one supposes that our
choicest productions have been produced by a single variation from the
aboriginal stock. We have proofs that this is not so in some cases, in
which exact records have been kept; thus, to give a very trifling
instance, the steadily-increasing size of the common gooseberry may be
quoted. We see an astonishing improvement in many florists' flowers,
when the flowers of the present day are compared with drawings made
only twenty or thirty years ago. When a race of plants is once pretty
well established, the seed-raisers do not pick out the best plants,
but merely go over their seed-beds, and pull up the 'rogues,' as they
call the plants that deviate from the proper standard. With animals
this
In regard to plants, there is another means of observing the
accumulated effects of selection -- namely, by comparing the
diversity of flowers in the different varieties of the same species in
the flower-garden; the diversity of leaves, pods, or tubers, or
whatever part is valued, in the kitchen-garden, in comparison with the
flowers of the same varieties; and the diversity of fruit of the same
species in the orchard, in comparison with the leaves It may be objected that the principle of selection has been reduced
to methodical practice for scarcely more than three-quarters of a
century; it has certainly been more attended to of late years, and
many treatises have been published on the subject; and the result, I
may add, has been, in a corresponding degree, rapid and important. But
it is very far from true that the principle is a modern discovery. I
could give several references to the full acknowledgement of the
importance of the principle in works of high antiquity. In rude and
At the present time, eminent breeders try by methodical selection,
with a distinct object in view, to make a new strain or sub-breed,
superior to anything existing in the country. But, for our purpose, a
kind of Selection, which may be called Unconscious, and which results
from every one trying to possess and breed from the best individual
animals, is more important. Thus, a man who intends keeping pointers
naturally tries to get as good dogs as he can, and afterwards breeds
from his own best dogs, but he has no wish or expectation of
permanently altering the breed. Nevertheless I cannot
By a similar process of selection, and by careful training, the
whole body of English racehorses have come to surpass in Youatt gives an excellent illustration of the effects of a course
of selection, which may be considered as unconsciously followed, in so
far that the breeders could never have expected or even have wished to
have produced the result which ensued -- namely, the production
of two distinct strains. The two flocks of Leicester sheep kept by Mr
Buckley and Mr Burgess, as Mr Youatt remarks, 'have been purely bred
from the original stock of Mr Bakewell for upwards of fifty years.
There is not a suspicion existing in the mind of any one at all
acquainted with the subject that the owner of either of them has
deviated in any one instance from the pure blood of Mr Bakewell's
flock, and yet the difference between the sheep possessed by these two
gentlemen is so great that they have the appearance of being quite
different varieties.'
If there exist savages so barbarous as never to think of the
inherited character of the offspring of their domestic animals, yet
any one animal particularly useful to them, for any special purpose,
would be carefully preserved during famines and other accidents, to
which savages are so liable, and such choice animals would thus
generally leave more offspring than the inferior ones; so that in this
case there would be a kind of unconscious selection going on. We see
the value set on animals even by the barbarians of Tierra del Fuego,
by their killing and devouring their old women, in times of dearth, as
of less value than their dogs.
In plants the same gradual process of improvement, through the
occasional preservation of the best individuals, whether or not
sufficiently distinct to be ranked
A large amount of change in our cultivated plants, thus slowly and
unconsciously accumulated, explains, as I believe, the well-known
fact, that in a vast number of cases we cannot recognise, and
therefore do not know, the wild parent-stocks of the plants which have
been longest cultivated in our flower and kitchen gardens. If it has
taken centuries or thousands of years to improve
In regard to the domestic animals kept by uncivilised man, it
should not be overlooked that they almost always have to struggle for
their own food, at least during certain seasons. And in two countries
very differently circumstanced, individuals of the same species,
having slightly different constitutions or structure, would often
succeed better in the one country than in the other, and thus by a
process of 'natural selection,' as will hereafter be more fully
explained, two sub-breeds might be formed. This, perhaps, partly
explains what has been remarked by some authors, namely, that the
varieties kept by savages have more of the character of species than
the varieties kept in civilised countries.
On the view here given of the all-important part which selection by
man has played, it becomes at once obvious, how it is that our
domestic races show adaptation in their structure or in their habits
to man's wants or fancies. We can, I think, further understand the
frequently abnormal character of our domestic races, and likewise
their differences being so great in external characters and relatively
so slight in internal parts or organs. Man can hardly select, or only
with much difficulty, any deviation of structure excepting such as is
externally visible; and indeed he rarely cares for what is internal.
He can never act by selection, excepting on variations
Nor let it be thought that some great deviation of structure would
be necessary to catch the fancier's eye: he perceives extremely small
differences, and it is in human nature to value any novelty, however
slight, in one's own possession. Nor must the value which would
formerly be set on any slight differences in the individuals of the
same species, be judged of by the value which would now be set on
them, after several breeds have once fairly been established. Many
slight differences might, and indeed do now, arise amongst pigeons,
which are rejected as faults or deviations from the standard of
perfection of each breed. The common goose has not given rise to any
marked varieties; hence the Thoulouse and the common breed, which
differ only in colour, that I think these views further explain what has sometimes been noticed
-- namely that we know nothing about the origin or history of any
of our domestic breeds. But, in fact, a breed, like a dialect of a
language, can hardly be said to have had a definite origin. A man
preserves and breeds from an individual with some slight deviation of
structure, or takes more care than usual in matching his best animals
and thus improves them, and the improved individuals slowly spread in
the immediate neighbourhood. But as yet they will hardly have a
distinct name, and from being only slightly valued, their history will
be disregarded. When further improved by the same slow and gradual
process, they will spread more widely, and will get recognised as
something distinct and valuable, and will then probably first receive
a provincial name. In semi-civilised countries, with little free
communication, the spreading and knowledge of any new sub-breed will
be a slow process. As soon as the points of value of the new sub-breed
are once fully acknowledged, the principle, as I have called it, of
unconscious selection will always tend, -- perhaps more at one
period than at another, as the breed rises or I must now say a few words on the circumstances, favourable, or the
reverse, to man's power of selection. A high degree of variability is
obviously favourable, as freely giving the materials for selection to
work on; not that mere individual differences are not amply
In the case of animals with separate sexes, facility in preventing
crosses is an important element of success in the formation of new
races, -- at least, in a country which is already stocked with
other races. In this respect enclosure of the land plays a part.
Wandering savages or the inhabitants of open plains rarely possess
more than one breed of the same species. Pigeons can be mated for
life, and this is a great convenience to the fancier, for thus many
races may be kept true, though mingled in the same aviary; and this
circumstance must have largely favoured the improvement and formation
of new breeds. Pigeons, I may add, can be propagated in great numbers
and at a very quick rate, and inferior birds may be freely rejected,
as when killed they serve for food. On the other hand, cats, from
their nocturnal rambling habits, cannot be matched, and, although so
much valued by women and children, we hardly ever see a distinct breed
kept up; such breeds as we do sometimes see are almost always imported
from some other country, often from islands. Although I do not doubt
that some domestic animals vary less than others, yet the rarity or
absence of distinct breeds of the cat, the donkey, peacock, goose,
&c., may be attributed in main part to selection not having been
brought into play: in cats, from the difficulty in pairing them; in
donkeys, from only a few being kept by poor people, and little
attention paid to their breeding; in peacocks, from not being very
easily reared and a large stock not kept; in geese, from being
valuable only for two purposes, food and feathers, and more especially
from no pleasure having been felt in the display of distinct breeds.
To sum up on the origin of our Domestic Races of animals and
plants. I believe that the conditions of life, from their action on
the reproductive system, are so far of the highest importance as
causing variability. I do not believe that variability is an
BEFORE applying the principles arrived at in the last chapter to organic
beings in a state of nature, we must briefly discuss whether these
latter are subject to any variation. To treat this subject at all
properly, a long catalogue of dry facts should be given; but these I
shall reserve for my future work. Nor shall I here discuss the various
definitions which have been given of the term species. No one
definition has as yet satisfied all naturalists; yet every naturalist
knows vaguely what he means when he speaks of a species. Generally the
term includes the unknown element of a distinct act of creation. The
term 'variety' is almost equally difficult to define; but here
community of descent is almost universally implied, though it can
rarely be proved. We have also what are called monstrosities; but they
graduate into varieties. By a monstrosity I presume is meant some
considerable deviation of structure in one part, either injurious to
or not useful to the species, and not generally propagated. Some
authors use the term 'variation' in a technical sense, as implying a
modification directly due to the physical conditions of life; and
'variations' in this sense are supposed not to be inherited: but who
can say that the dwarfed condition of shells in the brackish waters of
the Baltic, or dwarfed
Again, we have many slight differences which may be called There is one point connected with individual differences, which
seems to me extremely perplexing: I refer to those genera which have
sometimes been called 'protean' or 'polymorphic,' in which the species
present an inordinate amount of variation; and hardly two naturalists
can agree which forms to rank as species and which as varieties. We
may instance Rubus, Rosa, and Hieracium amongst plants, several genera
of insects, and several genera of Brachiopod shells. In most
polymorphic genera some of the species have fixed and definite
characters. Genera which are polymorphic in one country seem to be,
with some few exceptions, polymorphic in other countries, and
likewise, judging from Brachiopod shells, at former periods of time.
These facts seem to be very perplexing, for they seem to show that
this kind of variability is independent of the conditions of life. I
am inclined to suspect that we see in these polymorphic genera
variations in points of structure which are of no service or
disservice to the species, and which consequently have not been seized
on and rendered definite by natural selection, as hereafter will be
explained.
Those forms which possess in some considerable degree the character
of species, but which are so closely similar to some other forms, or
are so closely linked to them by intermediate gradations, that
naturalists do not like to rank them as distinct species, are in
several respects the most important for us. We have every reason to
believe that many of these doubtful and closely-allied forms have
permanently retained their characters in their own country for a long
time; for as long, as far as we know, as have good and true species.
practically, when a naturalist can unite two forms together by others
having intermediate characters, he treats the one as a variety of the
other, ranking the most common, but sometimes the one first described,
as the species, and the other as the variety. But cases of great
difficulty, which I will not here enumerate, sometimes occur in
deciding whether or not to rank one form as a variety of another, even
when they are closely connected by intermediate links; nor will the
commonly-assumed hybrid nature of the intermediate links always remove
the difficulty. In very many cases, however, one form is ranked as a
variety of another, not Hence, in determining whether a form should be ranked as a species
or a variety, the opinion of naturalists having sound judgement and
wide experience seems the only guide to follow. We must, however, in
many cases, decide by a majority of naturalists, for few well-marked
and well-known varieties can be named which have not been ranked as
species by at least some competent judges.
That varieties of this doubtful nature are far from uncommon cannot
be disputed. Compare the several floras of Great Britain, of France or
of the United States, drawn up by different botanists, and see what a
surprising number of forms have been ranked by one botanist as good
species, and by another as mere varieties. Mr H. C. Watson, to whom I
lie under deep obligation for assistance of all kinds, has marked for
me 182 British plants, which are generally considered as varieties,
but which have all been ranked by botanists as species; and in making
this list he has omitted many trifling varieties, but which
nevertheless have been ranked by some botanists as species, and he has
entirely omitted several highly polymorphic genera. Under genera,
including the most polymorphic forms, Mr Babington gives 251 species,
whereas Mr Bentham gives only 112, -- a difference of 139
doubtful forms! Amongst animals which unite for each birth, and which
are highly locomotive, doubtful forms, ranked by one zoologist as a
species and by another as a variety, can rarely be found within the
same country, but are common in separated areas. How many of those
birds and insects in North America and Europe, which differ very
slightly from each other, have been ranked by one eminent naturalist
as undoubted species, and by another as varieties, or, as they are
often called, as geographical races! Many years ago, when comparing,
and seeing others compare, the birds from the separate islands of the
Galapagos Archipelago, both one with another, and with those from the
American mainland, I was much struck how entirely vague and arbitrary
is the distinction between species and varieties. On the Many of the cases of strongly-marked varieties or doubtful species
well deserve consideration; for several interesting lines of argument,
from geographical distribution, analogical variation, hybridism,
&c., have been brought to bear on the attempt to determine their
rank. I will here give only a single instance, -- the well-known
one of the primrose and cowslip, or Primula veris and elatior. These
plants differ considerably in appearance; they have a different
flavour and emit a different odour; they flower at slightly different
periods; they grow in somewhat different stations; they ascend
mountains to different heights; they have different geographical
ranges; and lastly, according to very numerous experiments made during
several years by
Close investigation, in most cases, will bring naturalists to an
agreement how to rank doubtful forms. Yet it must be confessed, that
it is in the best-known countries that we find the greatest number of
forms of doubtful value. I have been struck with the fact, that if
any animal or plant in a state of nature be highly useful to man, or
from any cause closely attract his attention, varieties of it will
almost universally be found recorded. These varieties, moreover, will
be often ranked by some authors as species. Look at the common oak,
how closely it has been studied; yet a German author makes more than a
dozen species out of forms, which are very generally considered as
varieties; and in this country the highest botanical authorities and
practical men can be quoted to show that the sessile and pedunculated
oaks are either good and distinct species or mere varieties.
When a young naturalist commences the study of a group of organisms
quite unknown to him, he is at first much perplexed to determine what
differences to consider as specific, and what as varieties; for he
knows nothing of the amount and kind of variation to which the group
is subject; and this shows, at least, how very generally there is some
variation. But if he confine his attention to one class within one
country, he will soon make up his mind how to rank most of the
doubtful forms. His
Certainly no clear line of demarcation has as yet been drawn
between species and sub-species -- that is, the forms which in
the opinion of some naturalists come very near to, but do not quite
arrive at the rank of species; or, again, between sub-species and
well-marked varieties, or between lesser varieties and individual
differences. These differences blend into each other in an insensible
series; and a series impresses the mind with the idea of an actual
passage.
Hence I look at individual differences, though of small interest to
the systematist, as of high importance for us, as being the first step
towards such slight varieties as are barely thought worth recording in
works on natural history. And I look at varieties which are in any
degree more distinct and permanent, as steps leading to more
It need not be supposed that all varieties or incipient species
necessarily attain the rank of species. They may whilst in this
incipient state become extinct, or they may endure as varieties for
very long periods, as has been shown to be the case by Mr Wollaston
with the varieties of certain fossil land-shells in Madeira. If a
variety were to flourish so as to exceed in numbers From these remarks it will be seen that I look at the term species,
as one arbitrarily given for the sake of convenience to a set of
individuals closely resembling each other, and that it does not
essentially differ from the term variety, which is given to less
distinct and more fluctuating forms. The term variety, again, in
comparison with mere individual differences, is also applied
arbitrarily, and for mere convenience sake.
Guided by theoretical considerations, I thought that some
interesting results might be obtained in regard to the nature and
relations of the species which vary most, by tabulating all the
varieties in several well-worked floras. At first this seemed a
simple task; but Mr H. C. Watson, to whom I am much indebted for
valuable advice and assistance on this subject, soon convinced me that
there were many difficulties, as did subsequently Dr Hooker, even in
stronger terms. I shall reserve for my future work the discussion of
these difficulties, and the tables themselves of the proportional
numbers of the varying species. Dr Hooker permits me to add, that
after having carefully read my manuscript, and examined the tables, he
thinks that the following statements are fairly well established. The
whole subject, however, treated as it necessarily here is with much
brevity, is rather perplexing, and allusions cannot be avoided to the
'struggle for existence,' 'divergence of character,' and other
questions, hereafter to be discussed.
Alph. De Candolle and others have shown that plants which have very
wide ranges generally present varieties; and this might have been
expected, as they become exposed to diverse physical conditions, and
as they come into competition (which, as we shall hereafter see, is a
far more important circumstance) with different sets of organic
beings. But my tables further show that, in any limited country, the
species which are most common, that is abound most in individuals, and
the species which are most widely diffused within their own country
(and this is If the plants inhabiting a country and described in any Flora be
divided into two equal masses, all those in the larger genera being
placed on one side, and all those in the smaller genera on the other
side, a somewhat larger number of the very common and much diffused or
dominant species will be found on the side of the larger genera. This,
again, might have been anticipated; for the mere fact of many species
of the same genus inhabiting any country, shows that there is
something in the organic or inorganic conditions of that country
favourable to the genus; and, consequently, we might have expected to
have found in the larger genera, or those including many species, a
large proportional number of dominant species. But so many causes
tend to obscure this result, that I am surprised that my tables show
even a small majority on the side of the larger genera. I will here
allude to only two causes of obscurity. Fresh-water and salt-loving
plants have generally very wide ranges and are much diffused, but this
seems to be connected with the nature of the stations inhabited by
them, and has little or no relation to the size of the genera to which
the specie