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Books: On the Origin of Species

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

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Habit also has a decided 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 accompanied by an elongated head.
Some instances of correlation are quite whimsical; thus cats with blue
eyes are invariably deaf; colour and constitutional peculiarities go
together, of which many remarkable cases could be given amongst
animals and plants. From the facts collected by Heusinger, it appears
that white sheep and pigs are differently affected from coloured
individuals by certain vegetable poisons. Hairless dogs have imperfect
teeth; long-haired and coarse-haired animals are apt to have, as is
asserted, long or many horns; pigeons with feathered feet have skin
between their outer toes; pigeons with short beaks have small feet,
and those with long beaks large feet. Hence, if man goes on selecting,
and thus augmenting, any peculiarity, he will almost certainly
unconsciously modify other parts of the structure, owing to the
mysterious laws of the correlation of growth.

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, etc.; 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 organisation 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 deviation appears not unfrequently, and we see
it in the father and child, we cannot tell whether it may not be due
to the same original cause acting on both; but when amongst
individuals, apparently exposed to the same conditions, any very rare
deviation, due to some extraordinary combination of circumstances,
appears in the parent--say, once amongst several million
individuals--and it reappears in the child, the mere doctrine of
chances almost compels us to attribute its reappearance to
inheritance. Every one must have heard of cases of albinism, prickly
skin, hairy bodies, etc., appearing in several members of the same
family. If strange and rare deviations of structure are truly
inherited, less strange and commoner deviations may be freely admitted
to be inheritable. Perhaps the correct way of viewing the whole
subject, would be, to look at the inheritance of every character
whatever as the rule, and non-inheritance as the anomaly.

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 not be otherwise: thus the inherited
peculiarities in the horns of cattle could appear only in the
offspring when nearly mature; peculiarities in the silkworm are known
to appear at the corresponding caterpillar or cocoon stage. But
hereditary diseases and some other facts make me believe that the rule
has a wider extension, and that when there is no apparent reason why a
peculiarity should appear at any particular age, yet that it does tend
to appear in the offspring at the same period at which it first
appeared in the parent. I believe this rule to be of the highest
importance in explaining the laws of embryology. These remarks are of
course confined to the first APPEARANCE of the peculiarity, and not to
its primary cause, which may have acted on the ovules or male element;
in nearly the same manner as in the crossed offspring from a
short-horned cow by a long-horned bull, the greater length of horn,
though appearing late in life, is clearly due to the male element.

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 single variety
should be turned loose in its new home. Nevertheless, as our varieties
certainly do occasionally revert in some of their characters to
ancestral forms, it seems to me not improbable, that if we could
succeed in naturalising, or were to cultivate, during many
generations, the several races, for instance, of the cabbage, in very
poor soil (in which case, however, some effect would have to be
attributed to the direct action of the poor soil), that they would to
a large extent, or even wholly, revert to the wild aboriginal stock.
Whether or not the experiment would succeed, is not of great
importance for our line of argument; for by the experiment itself the
conditions of life are changed. If it could be shown that our domestic
varieties manifested a strong tendency to reversion,--that is, to lose
their acquired characters, whilst kept under unchanged conditions, and
whilst kept in a considerable body, so that free intercrossing might
check, by blending together, any slight deviations of structure, in
such case, I grant that we could deduce nothing from domestic
varieties in regard to species. But there is not a shadow of evidence
in favour of this view: to assert that we could not breed our cart and
race-horses, long and short-horned cattle, and poultry of various
breeds, and esculent vegetables, for an almost infinite number of
generations, would be opposed to all experience. I may add, that when
under nature the conditions of life do change, variations and
reversions of character probably do occur; but natural selection, as
will hereafter be explained, will determine how far the new characters
thus arising shall be preserved.

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 the same species, also, often have a somewhat monstrous
character; by which I mean, that, although differing from each other,
and from the other species of the same genus, in several trifling
respects, they often differ in an extreme degree in some one part,
both when compared one with another, and more especially when compared
with all the species in nature to which they are nearest allied. With
these exceptions (and with that of the perfect fertility of varieties
when crossed,--a subject hereafter to be discussed), domestic races of
the same species differ from each other in the same manner as, only in
most cases in a lesser degree than, do closely-allied species of the
same genus in a state of nature. I think this must be admitted, when
we find that there are hardly any domestic races, either amongst
animals or plants, which have not been ranked by some competent judges
as mere varieties, and by other competent judges as the descendants of
aboriginally distinct species. If any marked distinction existed
between domestic races and species, this source of doubt could not so
perpetually recur. It has often been stated that domestic races do not
differ from each other in characters of generic value. I think it
could be shown that this statement is hardly correct; but naturalists
differ most widely in determining what characters are of generic
value; all such valuations being at present empirical. Moreover, on
the view of the origin of genera which I shall presently give, we have
no right to expect often to meet with generic differences in our
domesticated productions.

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 descended from one or
several parent-species. This point, if it could be cleared up, would
be interesting; if, for instance, it could be shown that the
greyhound, bloodhound, terrier, spaniel, and bull-dog, which we all
know propagate their kind so truly, were the offspring of any single
species, then such facts would have great weight in making us doubt
about the immutability of the many very closely allied and natural
species--for instance, of the many foxes--inhabiting different
quarters of the world. I do not believe, as we shall presently see,
that all our dogs have descended from any one wild species; but, in
the case of some other domestic races, there is presumptive, or even
strong, evidence in favour of this view.

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 rein-deer, 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 of our
domestic animals is, that we find in the most ancient records, more
especially on the monuments of Egypt, much diversity in the breeds;
and that some of the breeds closely resemble, perhaps are identical
with, those still existing. Even if this latter fact were found more
strictly and generally true than seems to me to be the case, what does
it show, but that some of our breeds originated there, four or five
thousand years ago? But Mr. Horner's researches have rendered it in
some degree probable that man sufficiently civilized to have
manufactured pottery existed in the valley of the Nile thirteen or
fourteen thousand years ago; and who will pretend to say how long
before these ancient periods, savages, like those of Tierra del Fuego
or Australia, who possess a semi-domestic dog, may not have existed in
Egypt?

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, etc., 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 Indian fowl (Gallus
bankiva). In regard to ducks and rabbits, the breeds of which differ
considerably from each other in structure, I do not doubt that they
all have descended from the common wild duck and rabbit.

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 that Britain has now hardly one
peculiar mammal, and France but few distinct from those of Germany and
conversely, and so with Hungary, Spain, etc., but that each of these
kingdoms possesses several peculiar breeds of cattle, sheep, etc., we
must admit that many domestic breeds have originated in Europe; for
whence could they have been derived, as these several countries do not
possess a number of peculiar species as distinct parent-stocks? So it
is in India. Even in the case of the domestic dogs of the whole world,
which I fully admit have probably descended from several wild species,
I cannot doubt that there has been an immense amount of inherited
variation. Who can believe that animals closely resembling the Italian
greyhound, the bloodhound, the bull-dog, or Blenheim spaniel, etc.--so
unlike all wild Canidae--ever existed freely in a state of nature? It
has often been loosely said that all our races of dogs have been
produced by the crossing of a few aboriginal species; but by crossing
we can get only forms in some degree intermediate between their
parents; and if we account for our several domestic races by this
process, we must admit the former existence of the most extreme forms,
as the Italian greyhound, bloodhound, bull-dog, etc., in the wild
state. Moreover, the possibility of making distinct races by crossing
has been greatly exaggerated. There can be no doubt that a race may be
modified by occasional crosses, if aided by the careful selection of
those individual mongrels, which present any desired character; but
that a race could be obtained nearly intermediate between two
extremely different races or species, I can hardly believe. Sir J.
Sebright expressly experimentised for this object, and failed. The
offspring from the first cross between two pure breeds is tolerably
and sometimes (as I have found with pigeons) extremely uniform, and
everything seems simple enough; but when these mongrels are crossed
one with another for several generations, hardly two of them will be
alike, and then the extreme difficulty, or rather utter hopelessness,
of the task becomes apparent. Certainly, a breed intermediate between
TWO VERY DISTINCT breeds could not be got without extreme care and
long-continued selection; nor can I find a single case on record of a
permanent race having been thus formed.

ON THE BREEDS OF THE DOMESTIC PIGEON.

Believing that it is always best to study some special group, I have,
after deliberation, taken up domestic pigeons. I have kept every breed
which I could purchase or obtain, and have been most kindly favoured
with skins from several quarters of the world, more especially by the
Honourable W. Elliot from India, and by the Honourable C. Murray from
Persia. Many treatises in different languages have been published on
pigeons, and some of them are very important, as being of considerable
antiquity. I have associated with several eminent fanciers, and have
been permitted to join two of the London Pigeon Clubs. The diversity
of the breeds is something astonishing. Compare the English carrier
and the short-faced tumbler, and see the wonderful difference in their
beaks, entailing corresponding differences in their skulls. The
carrier, more especially the male bird, is also remarkable from the
wonderful development of the carunculated skin about the head, and
this is accompanied by greatly elongated eyelids, very large external
orifices to the nostrils, and a wide gape of mouth. The short-faced
tumbler has a beak in outline almost like that of a finch; and the
common tumbler has the singular and strictly inherited habit of flying
at a great height in a compact flock, and tumbling in the air head
over heels. The runt is a bird of great size, with long, massive beak
and large feet; some of the sub-breeds of runts have very long necks,
others very long wings and tails, others singularly short tails. The
barb is allied to the carrier, but, instead of a very long beak, has a
very short and very broad one. The pouter has a much elongated body,
wings, and legs; and its enormously developed crop, which it glories
in inflating, may well excite astonishment and even laughter. The
turbit has a very short and conical beak, with a line of reversed
feathers down the breast; and it has the habit of continually
expanding slightly the upper part of the oesophagus. The Jacobin has
the feathers so much reversed along the back of the neck that they
form a hood, and it has, proportionally to its size, much elongated
wing and tail feathers. The trumpeter and laugher, as their names
express, utter a very different coo from the other breeds. The fantail
has thirty or even forty tail-feathers, instead of twelve or fourteen,
the normal number in all members of the great pigeon family; and these
feathers are kept expanded, and are carried so erect that in good
birds the head and tail touch; the oil-gland is quite aborted. Several
other less distinct breeds might have been specified.

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 the
English carrier, the short-faced tumbler, the runt, the barb, pouter,
and fantail in the same genus; more especially as in each of these
breeds several truly-inherited sub-breeds, or species as he might have
called them, could be shown him.

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 (Columba livia),
including under this term several geographical races or sub-species,
which differ from each other in the most trifling respects. As several
of the reasons which have led me to this belief are in some degree
applicable in other cases, I will here briefly give them. If the
several breeds are not varieties, and have not proceeded from the
rock-pigeon, they must have descended from at least seven or eight
aboriginal stocks; for it is impossible to make the present domestic
breeds by the crossing of any lesser number: how, for instance, could
a pouter be produced by crossing two breeds unless one of the
parent-stocks possessed the characteristic enormous crop? The supposed
aboriginal stocks must all have been rock-pigeons, that is, not
breeding or willingly perching on trees. But besides C. livia, with
its geographical sub-species, only two or three other species of
rock-pigeons are known; and these have not any of the characters of
the domestic breeds. Hence the supposed aboriginal stocks must either
still exist in the countries where they were originally domesticated,
and yet be unknown to ornithologists; and this, considering their
size, habits, and remarkable characters, seems very improbable; or
they must have become extinct in the wild state. But birds breeding on
precipices, and good fliers, are unlikely to be exterminated; and the
common rock-pigeon, which has the same habits with the domestic
breeds, has not been exterminated even on several of the smaller
British islets, or on the shores of the Mediterranean. Hence the
supposed extermination of so many species having similar habits with
the rock-pigeon seems to me a very rash assumption. Moreover, the
several above-named domesticated breeds have been transported to all
parts of the world, and, therefore, some of them must have been
carried back again into their native country; but not one has ever
become wild or feral, though the dovecot-pigeon, which is the
rock-pigeon in a very slightly altered state, has become feral in
several places. Again, all recent experience shows that it is most
difficult to get any wild animal to breed freely under domestication;
yet on the hypothesis of the multiple origin of our pigeons, it must
be assumed that at least seven or eight species were so thoroughly
domesticated in ancient times by half-civilized man, as to be quite
prolific under confinement.

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