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Books: On the Origin of Species

C >> Charles Darwin >> On the Origin of Species




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 reversion to ancestral characters, if all the
domestic breeds have descended from the rock-pigeon. But if we deny
this, we must make one of the two following highly improbable
suppositions. Either, firstly, that all the several imagined
aboriginal stocks were coloured and marked like the rock-pigeon,
although no other existing species is thus coloured and marked, so
that in each separate breed there might be a tendency to revert to the
very same colours and markings. Or, secondly, that each breed, even
the purest, has within a dozen or, at most, within a score of
generations, been crossed by the rock-pigeon: I say within a dozen or
twenty generations, for we know of no fact countenancing the belief
that the child ever reverts to some one ancestor, removed by a greater
number of generations. In a breed which has been crossed only once
with some distinct breed, the tendency to reversion to any character
derived from such cross will naturally become less and less, as in
each succeeding generation there will be less of the foreign blood;
but when there has been no cross with a distinct breed, and there is a
tendency in both parents to revert to a character, which has been lost
during some former generation, this tendency, for all that we can see
to the contrary, may be transmitted undiminished for an indefinite
number of generations. These two distinct cases are often confounded
in treatises on inheritance.

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 CLEARLY DISTINCT being themselves perfectly
fertile. Some authors believe that long-continued domestication
eliminates this strong tendency to sterility: from the history of the
dog I think there is some probability in this hypothesis, if applied
to species closely related together, though it is unsupported by a
single experiment. But to extend the hypothesis so far as to suppose
that species, aboriginally as distinct as carriers, tumblers, pouters,
and fantails now are, should yield offspring perfectly fertile, inter
se, seems to me rash in the extreme.

From these several reasons, namely, the improbability of man having
formerly got seven or eight supposed species of pigeons to breed
freely under domestication; these supposed species being quite unknown
in a wild state, and their becoming nowhere feral; these species
having very abnormal characters in certain respects, as compared with
all other Columbidae, though so like in most other respects to the
rock-pigeon; the blue colour and various marks occasionally appearing
in all the breeds, both when kept pure and when crossed; the mongrel
offspring being perfectly fertile;--from these several reasons, taken
together, I can feel no doubt that all our domestic breeds have
descended from the Columba livia with its geographical sub-species.

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 Aegyptian 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 of fare in the previous dynasty. In the
time of the Romans, as we hear from Pliny, immense prices were given
for pigeons; "nay, they are come to this pass, that they can reckon up
their pedigree and race." Pigeons were much valued by Akber Khan in
India, about the year 1600; never less than 20,000 pigeons were taken
with the court. "The monarchs of Iran and Turan sent him some very
rare birds;" and, continues the courtly historian, "His Majesty by
crossing the breeds, which method was never practised before, has
improved them astonishingly." About this same period the Dutch were as
eager about pigeons as were the old Romans. The paramount importance
of these considerations in explaining the immense amount of variation
which pigeons have undergone, will be obvious when we treat of
Selection. We shall then, also, see how it is that the breeds so often
have a somewhat monstrous character. It is also a most favourable
circumstance for the production of distinct breeds, that male and
female pigeons can be easily mated for life; and thus different breeds
can be kept together in the same aviary.

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. Ask, as I have asked, a celebrated raiser of
Hereford cattle, whether his cattle might not have descended from long
horns, and he will laugh you to scorn. I have never met a pigeon, or
poultry, or duck, or rabbit fancier, who was not fully convinced that
each main breed was descended from a distinct species. Van Mons, in
his treatise on pears and apples, shows how utterly he disbelieves
that the several sorts, for instance a Ribston-pippin or Codlin-apple,
could ever have proceeded from the seeds of the same tree. Innumerable
other examples could be given. The explanation, I think, is simple:
from long-continued study they are strongly impressed with the
differences between the several races; and though they well know that
each race varies slightly, for they win their prizes by selecting such
slight differences, yet they ignore all general arguments, and refuse
to sum up in their minds slight differences accumulated during many
successive generations. May not those naturalists who, knowing far
less of the laws of inheritance than does the breeder, and knowing no
more than he does of the intermediate links in the long lines of
descent, yet admit that many of our domestic races have descended from
the same parents--may they not learn a lesson of caution, when they
deride the idea of species in a state of nature being lineal
descendants of other species?

SELECTION.

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
is that we see in them adaptation, not indeed to the animal's or
plant's own good, but to man's use or fancy. Some variations useful to
him have probably arisen suddenly, or by one step; many botanists, for
instance, believe that the fuller's teazle, with its hooks, which
cannot be rivalled by any mechanical contrivance, is only a variety of
the wild Dipsacus; and this amount of change may have suddenly arisen
in a seedling. So it has probably been with the turnspit dog; and this
is known to have been the case with the ancon sheep. But when we
compare the dray-horse and race-horse, the dromedary and camel, the
various breeds of sheep fitted either for cultivated land or mountain
pasture, with the wool of one breed good for one purpose, and that of
another breed for another purpose; when we compare the many breeds of
dogs, each good for man in very different ways; when we compare the
game-cock, so pertinacious in battle, with other breeds so little
quarrelsome, with "everlasting layers" which never desire to sit, and
with the bantam so small and elegant; when we compare the host of
agricultural, culinary, orchard, and flower-garden races of plants,
most useful to man at different seasons and for different purposes, or
so beautiful in his eyes, we must, I think, look further than to mere
variability. We cannot suppose that all the breeds were suddenly
produced as perfect and as useful as we now see them; indeed, in
several cases, we know that this has not been their history. The key
is man's power of accumulative selection: nature gives successive
variations; man adds them up in certain directions useful to him. In
this sense he may be said to make for himself useful breeds.

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 a large extent some breeds of cattle and
sheep. In order fully to realise what they have done, it is almost
necessary to read several of the many treatises devoted to this
subject, and to inspect the animals. Breeders habitually speak of an
animal's organisation as something quite plastic, which they can model
almost as they please. If I had space I could quote numerous passages
to this effect from highly competent authorities. Youatt, who was
probably better acquainted with the works of agriculturalists than
almost any other individual, and who was himself a very good judge of
an animal, speaks of the principle of selection as "that which enables
the agriculturist, not only to modify the character of his flock, but
to change it altogether. It is the magician's wand, by means of which
he may summon into life whatever form and mould he pleases." Lord
Somerville, speaking of what breeders have done for sheep, says:--"It
would seem as if they had chalked out upon a wall a form perfect in
itself, and then had given it existence." That most skilful breeder,
Sir John Sebright, used to say, with respect to pigeons, that "he
would produce any given feather in three years, but it would take him
six years to obtain head and beak." In Saxony the importance of the
principle of selection in regard to merino sheep is so fully
recognised, that men follow it as a trade: the sheep are placed on a
table and are studied, like a picture by a connoisseur; this is done
three times at intervals of months, and the sheep are each time marked
and classed, so that the very best may ultimately be selected for
breeding.

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; all the best
breeders are strongly opposed to this practice, except sometimes
amongst closely allied sub-breeds. And when a cross has been made, the
closest selection is far more indispensable even than in ordinary
cases. If selection consisted merely in separating some very distinct
variety, and breeding from it, the principle would be so obvious as
hardly to be worth notice; but its importance consists in the great
effect produced by the accumulation in one direction, during
successive generations, of differences absolutely inappreciable by an
uneducated eye--differences which I for one have vainly attempted to
appreciate. Not one man in a thousand has accuracy of eye and judgment
sufficient to become an eminent breeder. If gifted with these
qualities, and he studies his subject for years, and devotes his
lifetime to it with indomitable perseverance, he will succeed, and may
make great improvements; if he wants any of these qualities, he will
assuredly fail. Few would readily believe in the natural capacity and
years of practice requisite to become even a skilful pigeon-fancier.

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 kind of selection is, in fact, also followed; for hardly any one
is so careless as to allow his worst animals to breed.

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 and flowers of the same
set of varieties. See how different the leaves of the cabbage are, and
how extremely alike the flowers; how unlike the flowers of the
heartsease are, and how alike the leaves; how much the fruit of the
different kinds of gooseberries differ in size, colour, shape, and
hairiness, and yet the flowers present very slight differences. It is
not that the varieties which differ largely in some one point do not
differ at all in other points; this is hardly ever, perhaps never, the
case. The laws of correlation of growth, the importance of which
should never be overlooked, will ensure some differences; but, as a
general rule, I cannot doubt that the continued selection of slight
variations, either in the leaves, the flowers, or the fruit, will
produce races differing from each other chiefly in these characters.

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 acknowledgment of the
importance of the principle in works of high antiquity. In rude and
barbarous periods of English history choice animals were often
imported, and laws were passed to prevent their exportation: the
destruction of horses under a certain size was ordered, and this may
be compared to the "roguing" of plants by nurserymen. The principle of
selection I find distinctly given in an ancient Chinese encyclopaedia.
Explicit rules are laid down by some of the Roman classical writers.
From passages in Genesis, it is clear that the colour of domestic
animals was at that early period attended to. Savages now sometimes
cross their dogs with wild canine animals, to improve the breed, and
they formerly did so, as is attested by passages in Pliny. The savages
in South Africa match their draught cattle by colour, as do some of
the Esquimaux their teams of dogs. Livingstone shows how much good
domestic breeds are valued by the negroes of the interior of Africa
who have not associated with Europeans. Some of these facts do not
show actual selection, but they show that the breeding of domestic
animals was carefully attended to in ancient times, and is now
attended to by the lowest savages. It would, indeed, have been a
strange fact, had attention not been paid to breeding, for the
inheritance of good and bad qualities is so obvious.

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 doubt that this
process, continued during centuries, would improve and modify any
breed, in the same way as Bakewell, Collins, etc., by this very same
process, only carried on more methodically, did greatly modify, even
during their own lifetimes, the forms and qualities of their cattle.
Slow and insensible changes of this kind could never be recognised
unless actual measurements or careful drawings of the breeds in
question had been made long ago, which might serve for comparison. In
some cases, however, unchanged or but little changed individuals of
the same breed may be found in less civilised districts, where the
breed has been less improved. There is reason to believe that King
Charles's spaniel has been unconsciously modified to a large extent
since the time of that monarch. Some highly competent authorities are
convinced that the setter is directly derived from the spaniel, and
has probably been slowly altered from it. It is known that the English
pointer has been greatly changed within the last century, and in this
case the change has, it is believed, been chiefly effected by crosses
with the fox-hound; but what concerns us is, that the change has been
effected unconsciously and gradually, and yet so effectually, that,
though the old Spanish pointer certainly came from Spain, Mr. Borrow
has not seen, as I am informed by him, any native dog in Spain like
our pointer.

By a similar process of selection, and by careful training, the whole
body of English racehorses have come to surpass in fleetness and size
the parent Arab stock, so that the latter, by the regulations for the
Goodwood Races, are favoured in the weights they carry. Lord Spencer
and others have shown how the cattle of England have increased in
weight and in early maturity, compared with the stock formerly kept in
this country. By comparing the accounts given in old pigeon treatises
of carriers and tumblers with these breeds as now existing in Britain,
India, and Persia, we can, I think, clearly trace the stages through
which they have insensibly passed, and come to differ so greatly from
the rock-pigeon.

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 at their first appearance as
distinct varieties, and whether or not two or more species or races
have become blended together by crossing, may plainly be recognised in
the increased size and beauty which we now see in the varieties of the
heartsease, rose, pelargonium, dahlia, and other plants, when compared
with the older varieties or with their parent-stocks. No one would
ever expect to get a first-rate heartsease or dahlia from the seed of
a wild plant. No one would expect to raise a first-rate melting pear
from the seed of a wild pear, though he might succeed from a poor
seedling growing wild, if it had come from a garden-stock. The pear,
though cultivated in classical times, appears, from Pliny's
description, to have been a fruit of very inferior quality. I have
seen great surprise expressed in horticultural works at the wonderful
skill of gardeners, in having produced such splendid results from such
poor materials; but the art, I cannot doubt, has been simple, and, as
far as the final result is concerned, has been followed almost
unconsciously. It has consisted in always cultivating the best known
variety, sowing its seeds, and, when a slightly better variety has
chanced to appear, selecting it, and so onwards. But the gardeners of
the classical period, who cultivated the best pear they could procure,
never thought what splendid fruit we should eat; though we owe our
excellent fruit, in some small degree, to their having naturally
chosen and preserved the best varieties they could anywhere find.