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Books: On the Origin of Species

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




With respect to the horse, I have collected cases in England of the
spinal stripe in horses of the most distinct breeds, and of ALL
colours; transverse bars on the legs are not rare in duns, mouse-duns,
and in one instance in a chestnut: a faint shoulder-stripe may
sometimes be seen in duns, and I have seen a trace in a bay horse. My
son made a careful examination and sketch for me of a dun Belgian
cart-horse with a double stripe on each shoulder and with leg-stripes;
and a man, whom I can implicitly trust, has examined for me a small
dun Welch pony with THREE short parallel stripes on each shoulder.

In the north-west part of India the Kattywar breed of horses is so
generally striped, that, as I hear from Colonel Poole, who examined
the breed for the Indian Government, a horse without stripes is not
considered as purely-bred. The spine is always striped; the legs are
generally barred; and the shoulder-stripe, which is sometimes double
and sometimes treble, is common; the side of the face, moreover, is
sometimes striped. The stripes are plainest in the foal; and sometimes
quite disappear in old horses. Colonel Poole has seen both gray and
bay Kattywar horses striped when first foaled. I have, also, reason to
suspect, from information given me by Mr. W. W. Edwards, that with the
English race-horse the spinal stripe is much commoner in the foal than
in the full-grown animal. Without here entering on further details, I
may state that I have collected cases of leg and shoulder stripes in
horses of very different breeds, in various countries from Britain to
Eastern China; and from Norway in the north to the Malay Archipelago
in the south. In all parts of the world these stripes occur far
oftenest in duns and mouse-duns; by the term dun a large range of
colour is included, from one between brown and black to a close
approach to cream-colour.

I am aware that Colonel Hamilton Smith, who has written on this
subject, believes that the several breeds of the horse have descended
from several aboriginal species--one of which, the dun, was striped;
and that the above-described appearances are all due to ancient
crosses with the dun stock. But I am not at all satisfied with this
theory, and should be loth to apply it to breeds so distinct as the
heavy Belgian cart-horse, Welch ponies, cobs, the lanky Kattywar race,
etc., inhabiting the most distant parts of the world.

Now let us turn to the effects of crossing the several species of the
horse-genus. Rollin asserts, that the common mule from the ass and
horse is particularly apt to have bars on its legs. I once saw a mule
with its legs so much striped that any one at first would have thought
that it must have been the product of a zebra; and Mr. W. C. Martin,
in his excellent treatise on the horse, has given a figure of a
similar mule. In four coloured drawings, which I have seen, of hybrids
between the ass and zebra, the legs were much more plainly barred than
the rest of the body; and in one of them there was a double
shoulder-stripe. In Lord Moreton's famous hybrid from a chestnut mare
and male quagga, the hybrid, and even the pure offspring subsequently
produced from the mare by a black Arabian sire, were much more plainly
barred across the legs than is even the pure quagga. Lastly, and this
is another most remarkable case, a hybrid has been figured by Dr. Gray
(and he informs me that he knows of a second case) from the ass and
the hemionus; and this hybrid, though the ass seldom has stripes on
its legs and the hemionus has none and has not even a shoulder-stripe,
nevertheless had all four legs barred, and had three short
shoulder-stripes, like those on the dun Welch pony, and even had some
zebra-like stripes on the sides of its face. With respect to this last
fact, I was so convinced that not even a stripe of colour appears from
what would commonly be called an accident, that I was led solely from
the occurrence of the face-stripes on this hybrid from the ass and
hemionus, to ask Colonel Poole whether such face-stripes ever occur in
the eminently striped Kattywar breed of horses, and was, as we have
seen, answered in the affirmative.

What now are we to say to these several facts? We see several very
distinct species of the horse-genus becoming, by simple variation,
striped on the legs like a zebra, or striped on the shoulders like an
ass. In the horse we see this tendency strong whenever a dun tint
appears--a tint which approaches to that of the general colouring of
the other species of the genus. The appearance of the stripes is not
accompanied by any change of form or by any other new character. We
see this tendency to become striped most strongly displayed in hybrids
from between several of the most distinct species. Now observe the
case of the several breeds of pigeons: they are descended from a
pigeon (including two or three sub-species or geographical races) of a
bluish colour, with certain bars and other marks; and when any breed
assumes by simple variation a bluish tint, these bars and other marks
invariably reappear; but without any other change of form or
character. When the oldest and truest breeds of various colours are
crossed, we see a strong tendency for the blue tint and bars and marks
to reappear in the mongrels. I have stated that the most probable
hypothesis to account for the reappearance of very ancient characters,
is--that there is a TENDENCY in the young of each successive
generation to produce the long-lost character, and that this tendency,
from unknown causes, sometimes prevails. And we have just seen that in
several species of the horse-genus the stripes are either plainer or
appear more commonly in the young than in the old. Call the breeds of
pigeons, some of which have bred true for centuries, species; and how
exactly parallel is the case with that of the species of the
horse-genus! For myself, I venture confidently to look back thousands
on thousands of generations, and I see an animal striped like a zebra,
but perhaps otherwise very differently constructed, the common parent
of our domestic horse, whether or not it be descended from one or more
wild stocks, of the ass, the hemionus, quagga, and zebra.

He who believes that each equine species was independently created,
will, I presume, assert that each species has been created with a
tendency to vary, both under nature and under domestication, in this
particular manner, so as often to become striped like other species of
the genus; and that each has been created with a strong tendency, when
crossed with species inhabiting distant quarters of the world, to
produce hybrids resembling in their stripes, not their own parents,
but other species of the genus. To admit this view is, as it seems to
me, to reject a real for an unreal, or at least for an unknown, cause.
It makes the works of God a mere mockery and deception; I would almost
as soon believe with the old and ignorant cosmogonists, that fossil
shells had never lived, but had been created in stone so as to mock
the shells now living on the sea-shore.

SUMMARY.

Our ignorance of the laws of variation is profound. Not in one case
out of a hundred can we pretend to assign any reason why this or that
part differs, more or less, from the same part in the parents. But
whenever we have the means of instituting a comparison, the same laws
appear to have acted in producing the lesser differences between
varieties of the same species, and the greater differences between
species of the same genus. The external conditions of life, as climate
and food, etc., seem to have induced some slight modifications. Habit
in producing constitutional differences, and use in strengthening, and
disuse in weakening and diminishing organs, seem to have been more
potent in their effects. Homologous parts tend to vary in the same
way, and homologous parts tend to cohere. Modifications in hard parts
and in external parts sometimes affect softer and internal parts. When
one part is largely developed, perhaps it tends to draw nourishment
from the adjoining parts; and every part of the structure which can be
saved without detriment to the individual, will be saved. Changes of
structure at an early age will generally affect parts subsequently
developed; and there are very many other correlations of growth, the
nature of which we are utterly unable to understand. Multiple parts
are variable in number and in structure, perhaps arising from such
parts not having been closely specialised to any particular function,
so that their modifications have not been closely checked by natural
selection. It is probably from this same cause that organic beings low
in the scale of nature are more variable than those which have their
whole organisation more specialised, and are higher in the scale.
Rudimentary organs, from being useless, will be disregarded by natural
selection, and hence probably are variable. Specific characters--that
is, the characters which have come to differ since the several species
of the same genus branched off from a common parent--are more variable
than generic characters, or those which have long been inherited, and
have not differed within this same period. In these remarks we have
referred to special parts or organs being still variable, because they
have recently varied and thus come to differ; but we have also seen in
the second Chapter that the same principle applies to the whole
individual; for in a district where many species of any genus are
found--that is, where there has been much former variation and
differentiation, or where the manufactory of new specific forms has
been actively at work--there, on an average, we now find most
varieties or incipient species. Secondary sexual characters are highly
variable, and such characters differ much in the species of the same
group. Variability in the same parts of the organisation has generally
been taken advantage of in giving secondary sexual differences to the
sexes of the same species, and specific differences to the several
species of the same genus. Any part or organ developed to an
extraordinary size or in an extraordinary manner, in comparison with
the same part or organ in the allied species, must have gone through
an extraordinary amount of modification since the genus arose; and
thus we can understand why it should often still be variable in a much
higher degree than other parts; for variation is a long-continued and
slow process, and natural selection will in such cases not as yet have
had time to overcome the tendency to further variability and to
reversion to a less modified state. But when a species with any
extraordinarily-developed organ has become the parent of many modified
descendants--which on my view must be a very slow process, requiring a
long lapse of time--in this case, natural selection may readily have
succeeded in giving a fixed character to the organ, in however
extraordinary a manner it may be developed. Species inheriting nearly
the same constitution from a common parent and exposed to similar
influences will naturally tend to present analogous variations, and
these same species may occasionally revert to some of the characters
of their ancient progenitors. Although new and important modifications
may not arise from reversion and analogous variation, such
modifications will add to the beautiful and harmonious diversity of
nature.

Whatever the cause may be of each slight difference in the offspring
from their parents--and a cause for each must exist--it is the steady
accumulation, through natural selection, of such differences, when
beneficial to the individual, that gives rise to all the more
important modifications of structure, by which the innumerable beings
on the face of this earth are enabled to struggle with each other, and
the best adapted to survive.


CHAPTER 6. DIFFICULTIES ON THEORY.

Difficulties on the theory of descent with modification.
Transitions.
Absence or rarity of transitional varieties.
Transitions in habits of life.
Diversified habits in the same species.
Species with habits widely different from those of their allies.
Organs of extreme perfection.
Means of transition.
Cases of difficulty.
Natura non facit saltum.
Organs of small importance.
Organs not in all cases absolutely perfect.
The law of Unity of Type and of the Conditions of Existence embraced
by the theory of Natural Selection.

Long before having arrived at this part of my work, a crowd of
difficulties will have occurred to the reader. Some of them are so
grave that to this day I can never reflect on them without being
staggered; but, to the best of my judgment, the greater number are
only apparent, and those that are real are not, I think, fatal to my
theory.

These difficulties and objections may be classed under the following
heads:--

Firstly, why, if species have descended from other species by
insensibly fine gradations, do we not everywhere see innumerable
transitional forms? Why is not all nature in confusion instead of the
species being, as we see them, well defined?

Secondly, is it possible that an animal having, for instance, the
structure and habits of a bat, could have been formed by the
modification of some animal with wholly different habits? Can we
believe that natural selection could produce, on the one hand, organs
of trifling importance, such as the tail of a giraffe, which serves as
a fly-flapper, and, on the other hand, organs of such wonderful
structure, as the eye, of which we hardly as yet fully understand the
inimitable perfection?

Thirdly, can instincts be acquired and modified through natural
selection? What shall we say to so marvellous an instinct as that
which leads the bee to make cells, which have practically anticipated
the discoveries of profound mathematicians?

Fourthly, how can we account for species, when crossed, being sterile
and producing sterile offspring, whereas, when varieties are crossed,
their fertility is unimpaired?

The two first heads shall be here discussed--Instinct and Hybridism in
separate chapters.

ON THE ABSENCE OR RARITY OF TRANSITIONAL VARIETIES.

As natural selection acts solely by the preservation of profitable
modifications, each new form will tend in a fully-stocked country to
take the place of, and finally to exterminate, its own less improved
parent or other less-favoured forms with which it comes into
competition. Thus extinction and natural selection will, as we have
seen, go hand in hand. Hence, if we look at each species as descended
from some other unknown form, both the parent and all the transitional
varieties will generally have been exterminated by the very process of
formation and perfection of the new form.

But, as by this theory innumerable transitional forms must have
existed, why do we not find them embedded in countless numbers in the
crust of the earth? It will be much more convenient to discuss this
question in the chapter on the Imperfection of the geological record;
and I will here only state that I believe the answer mainly lies in
the record being incomparably less perfect than is generally supposed;
the imperfection of the record being chiefly due to organic beings not
inhabiting profound depths of the sea, and to their remains being
embedded and preserved to a future age only in masses of sediment
sufficiently thick and extensive to withstand an enormous amount of
future degradation; and such fossiliferous masses can be accumulated
only where much sediment is deposited on the shallow bed of the sea,
whilst it slowly subsides. These contingencies will concur only
rarely, and after enormously long intervals. Whilst the bed of the sea
is stationary or is rising, or when very little sediment is being
deposited, there will be blanks in our geological history. The crust
of the earth is a vast museum; but the natural collections have been
made only at intervals of time immensely remote.

But it may be urged that when several closely-allied species inhabit
the same territory we surely ought to find at the present time many
transitional forms. Let us take a simple case: in travelling from
north to south over a continent, we generally meet at successive
intervals with closely allied or representative species, evidently
filling nearly the same place in the natural economy of the land.
These representative species often meet and interlock; and as the one
becomes rarer and rarer, the other becomes more and more frequent,
till the one replaces the other. But if we compare these species where
they intermingle, they are generally as absolutely distinct from each
other in every detail of structure as are specimens taken from the
metropolis inhabited by each. By my theory these allied species have
descended from a common parent; and during the process of
modification, each has become adapted to the conditions of life of its
own region, and has supplanted and exterminated its original parent
and all the transitional varieties between its past and present
states. Hence we ought not to expect at the present time to meet with
numerous transitional varieties in each region, though they must have
existed there, and may be embedded there in a fossil condition. But in
the intermediate region, having intermediate conditions of life, why
do we not now find closely-linking intermediate varieties? This
difficulty for a long time quite confounded me. But I think it can be
in large part explained.

In the first place we should be extremely cautious in inferring,
because an area is now continuous, that it has been continuous during
a long period. Geology would lead us to believe that almost every
continent has been broken up into islands even during the later
tertiary periods; and in such islands distinct species might have been
separately formed without the possibility of intermediate varieties
existing in the intermediate zones. By changes in the form of the land
and of climate, marine areas now continuous must often have existed
within recent times in a far less continuous and uniform condition
than at present. But I will pass over this way of escaping from the
difficulty; for I believe that many perfectly defined species have
been formed on strictly continuous areas; though I do not doubt that
the formerly broken condition of areas now continuous has played an
important part in the formation of new species, more especially with
freely-crossing and wandering animals.

In looking at species as they are now distributed over a wide area, we
generally find them tolerably numerous over a large territory, then
becoming somewhat abruptly rarer and rarer on the confines, and
finally disappearing. Hence the neutral territory between two
representative species is generally narrow in comparison with the
territory proper to each. We see the same fact in ascending mountains,
and sometimes it is quite remarkable how abruptly, as Alph. De
Candolle has observed, a common alpine species disappears. The same
fact has been noticed by Forbes in sounding the depths of the sea with
the dredge. To those who look at climate and the physical conditions
of life as the all-important elements of distribution, these facts
ought to cause surprise, as climate and height or depth graduate away
insensibly. But when we bear in mind that almost every species, even
in its metropolis, would increase immensely in numbers, were it not
for other competing species; that nearly all either prey on or serve
as prey for others; in short, that each organic being is either
directly or indirectly related in the most important manner to other
organic beings, we must see that the range of the inhabitants of any
country by no means exclusively depends on insensibly changing
physical conditions, but in large part on the presence of other
species, on which it depends, or by which it is destroyed, or with
which it comes into competition; and as these species are already
defined objects (however they may have become so), not blending one
into another by insensible gradations, the range of any one species,
depending as it does on the range of others, will tend to be sharply
defined. Moreover, each species on the confines of its range, where it
exists in lessened numbers, will, during fluctuations in the number of
its enemies or of its prey, or in the seasons, be extremely liable to
utter extermination; and thus its geographical range will come to be
still more sharply defined.

If I am right in believing that allied or representative species, when
inhabiting a continuous area, are generally so distributed that each
has a wide range, with a comparatively narrow neutral territory
between them, in which they become rather suddenly rarer and rarer;
then, as varieties do not essentially differ from species, the same
rule will probably apply to both; and if we in imagination adapt a
varying species to a very large area, we shall have to adapt two
varieties to two large areas, and a third variety to a narrow
intermediate zone. The intermediate variety, consequently, will exist
in lesser numbers from inhabiting a narrow and lesser area; and
practically, as far as I can make out, this rule holds good with
varieties in a state of nature. I have met with striking instances of
the rule in the case of varieties intermediate between well-marked
varieties in the genus Balanus. And it would appear from information
given me by Mr. Watson, Dr. Asa Gray, and Mr. Wollaston, that
generally when varieties intermediate between two other forms occur,
they are much rarer numerically than the forms which they connect.
Now, if we may trust these facts and inferences, and therefore
conclude that varieties linking two other varieties together have
generally existed in lesser numbers than the forms which they connect,
then, I think, we can understand why intermediate varieties should not
endure for very long periods;--why as a general rule they should be
exterminated and disappear, sooner than the forms which they
originally linked together.

For any form existing in lesser numbers would, as already remarked,
run a greater chance of being exterminated than one existing in large
numbers; and in this particular case the intermediate form would be
eminently liable to the inroads of closely allied forms existing on
both sides of it. But a far more important consideration, as I
believe, is that, during the process of further modification, by which
two varieties are supposed on my theory to be converted and perfected
into two distinct species, the two which exist in larger numbers from
inhabiting larger areas, will have a great advantage over the
intermediate variety, which exists in smaller numbers in a narrow and
intermediate zone. For forms existing in larger numbers will always
have a better chance, within any given period, of presenting further
favourable variations for natural selection to seize on, than will the
rarer forms which exist in lesser numbers. Hence, the more common
forms, in the race for life, will tend to beat and supplant the less
common forms, for these will be more slowly modified and improved. It
is the same principle which, as I believe, accounts for the common
species in each country, as shown in the second chapter, presenting on
an average a greater number of well-marked varieties than do the rarer
species. I may illustrate what I mean by supposing three varieties of
sheep to be kept, one adapted to an extensive mountainous region; a
second to a comparatively narrow, hilly tract; and a third to wide
plains at the base; and that the inhabitants are all trying with equal
steadiness and skill to improve their stocks by selection; the chances
in this case will be strongly in favour of the great holders on the
mountains or on the plains improving their breeds more quickly than
the small holders on the intermediate narrow, hilly tract; and
consequently the improved mountain or plain breed will soon take the
place of the less improved hill breed; and thus the two breeds, which
originally existed in greater numbers, will come into close contact
with each other, without the interposition of the supplanted,
intermediate hill-variety.

To sum up, I believe that species come to be tolerably well-defined
objects, and do not at any one period present an inextricable chaos of
varying and intermediate links: firstly, because new varieties are
very slowly formed, for variation is a very slow process, and natural
selection can do nothing until favourable variations chance to occur,
and until a place in the natural polity of the country can be better
filled by some modification of some one or more of its inhabitants.
And such new places will depend on slow changes of climate, or on the
occasional immigration of new inhabitants, and, probably, in a still
more important degree, on some of the old inhabitants becoming slowly
modified, with the new forms thus produced and the old ones acting and
reacting on each other. So that, in any one region and at any one
time, we ought only to see a few species presenting slight
modifications of structure in some degree permanent; and this
assuredly we do see.