On the Origin of Species By Means of Natural Selection
by Charles Darwin · English · Gutenberg #1228 · 952,432 chars · public domain (US)
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Title: On the Origin of Species By Means of Natural Selection
Author: Charles Darwin
Release date: March 1, 1998 [eBook #1228]
Most recently updated: October 6, 2026
Language: English
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1228 1859, First Edition
22764 1860, Second Edition
2009 1872, Sixth Edition, considered the definitive edition.
On
the Origin of Species
BY MEANS OF NATURAL SELECTION,
OR THE
PRESERVATION OF FAVOURED RACES IN THE STRUGGLE FOR LIFE.
By Charles Darwin, M.A.,
Fellow Of The Royal, Geological, Linnæan, Etc., Societies;
Author Of ‘Journal Of Researches During H.M.S. Beagle’s Voyage
Round The World.’
LONDON:
JOHN MURRAY, ALBEMARLE STREET.
1859.
“But with regard to the material world, we can at least go so far as
this—we can perceive that events are brought about not by insulated
interpositions of Divine power, exerted in each particular case, but by the
establishment of general laws.”
W. WHEWELL: _Bridgewater Treatise_.
“To conclude, therefore, let no man out of a weak conceit of sobriety, or
an ill-applied moderation, think or maintain, that a man can search too far or
be too well studied in the book of God’s word, or in the book of
God’s works; divinity or philosophy; but rather let men endeavour an
endless progress or proficience in both.”
BACON: _Advancement of Learning_.
_Down, Bromley, Kent,
October_, 1_st_, 1859.
Contents
INTRODUCTION.
1. VARIATION UNDER DOMESTICATION.
2. VARIATION UNDER NATURE.
3. STRUGGLE FOR EXISTENCE.
4. NATURAL SELECTION.
5. LAWS OF VARIATION.
6. DIFFICULTIES ON THEORY.
7. INSTINCT.
8. HYBRIDISM.
9. ON THE IMPERFECTION OF THE GEOLOGICAL RECORD.
10. ON THE GEOLOGICAL SUCCESSION OF ORGANIC BEINGS.
11. GEOGRAPHICAL DISTRIBUTION.
12. GEOGRAPHICAL DISTRIBUTION—_continued_.
13. MUTUAL AFFINITIES OF ORGANIC BEINGS: MORPHOLOGY:
14. RECAPITULATION AND CONCLUSION.
INDEX
DETEAILED CONTENTS. ON THE ORIGIN OF SPECIES.
INTRODUCTION.
CHAPTER I. VARIATION UNDER DOMESTICATION.
Causes of Variability.
Effects of Habit.
Correlation of Growth.
Inheritance.
Character of Domestic Varieties.
Difficulty of distinguishing between Varieties and Species.
Origin of Domestic Varieties from one or more Species.
Domestic Pigeons, their Differences and Origin.
Principle of Selection anciently followed, its Effects.
Methodical and Unconscious Selection.
Unknown Origin of our Domestic Productions.
Circumstances favourable to Man’s power of Selection.
CHAPTER 2. VARIATION UNDER NATURE.
Variability.
Individual Differences.
Doubtful species.
Wide ranging, much diffused, and common species vary most.
Species of the larger genera in any country vary more than the
species of the smaller genera.
Many of the species of the larger genera resemble varieties in being
very closely, but unequally, related to each other, and in having
restricted ranges.
CHAPTER 3. STRUGGLE FOR EXISTENCE.
Bears on natural selection.
The term used in a wide sense.
Geometrical powers of increase.
Rapid increase of naturalised animals and plants.
Nature of the checks to increase.
Competition universal.
Effects of climate.
Protection from the number of individuals.
Complex relations of all animals and plants throughout nature.
Struggle for life most severe between individuals and varieties of
the same species; often severe between species of the same genus.
The relation of organism to organism the most important of all
relations.
CHAPTER 4. NATURAL SELECTION.
Natural Selection: its power compared with man’s selection, its power
on characters of trifling importance, its power at all ages and on
both sexes.
Sexual Selection.
On the generality of intercrosses between individuals of the same
species.
Circumstances favourable and unfavourable to Natural Selection,
namely, intercrossing, isolation, number of individuals.
Slow action.
Extinction caused by Natural Selection.
Divergence of Character, related to the diversity of inhabitants of
any small area, and to naturalisation.
Action of Natural Selection, through Divergence of Character and
Extinction, on the descendants from a common parent.
Explains the Grouping of all organic beings.
CHAPTER 5. LAWS OF VARIATION.
Effects of external conditions.
Use and disuse, combined with natural selection; organs of flight and
of vision.
Acclimatisation.
Correlation of growth.
Compensation and economy of growth.
False correlations.
Multiple, rudimentary, and lowly organised structures variable.
Parts developed in an unusual manner are highly variable: specific
characters more variable than generic: secondary sexual characters
variable.
Species of the same genus vary in an analogous manner.
Reversions to long-lost characters.
Summary.
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.
CHAPTER 7. INSTINCT.
Instincts comparable with habits, but different in their origin.
Instincts graduated.
Aphides and ants.
Instincts variable.
Domestic instincts, their origin.
Natural instincts of the cuckoo, ostrich, and parasitic bees.
Slave-making ants.
Hive-bee, its cell-making instinct.
Difficulties on the theory of the Natural Selection of instincts.
Neuter or sterile insects.
Summary.
CHAPTER 8. HYBRIDISM.
Distinction between the sterility of first crosses and of hybrids.
Sterility various in degree, not universal, affected by close
interbreeding, removed by domestication.
Laws governing the sterility of hybrids.
Sterility not a special endowment, but incidental on other
differences.
Causes of the sterility of first crosses and of hybrids.
Parallelism between the effects of changed conditions of life and
crossing.
Fertility of varieties when crossed and of their mongrel offspring
not universal.
Hybrids and mongrels compared independently of their fertility.
Summary.
CHAPTER 9. ON THE IMPERFECTION OF THE GEOLOGICAL RECORD.
On the absence of intermediate varieties at the present day.
On the nature of extinct intermediate varieties; on their number.
On the vast lapse of time, as inferred from the rate of deposition
and of denudation.
On the poorness of our palæontological collections.
On the intermittence of geological formations.
On the absence of intermediate varieties in any one formation.
On the sudden appearance of groups of species.
On their sudden appearance in the lowest known fossiliferous strata.
CHAPTER 10. ON THE GEOLOGICAL SUCCESSION OF ORGANIC BEINGS.
On the slow and successive appearance of new species.
On their different rates of change.
Species once lost do not reappear.
Groups of species follow the same general rules in their appearance
and disappearance as do single species.
On Extinction.
On simultaneous changes in the forms of life throughout the world.
On the affinities of extinct species to each other and to living
species.
On the state of development of ancient forms.
On the succession of the same types within the same areas.
Summary of preceding and present chapters.
CHAPTER 11. GEOGRAPHICAL DISTRIBUTION.
Present distribution cannot be accounted for by differences in
physical conditions.
Importance of barriers.
Affinity of the productions of the same continent.
Centres of creation.
Means of dispersal, by changes of climate and of the level of the
land, and by occasional means.
Dispersal during the Glacial period co-extensive with the world.
CHAPTER 12. GEOGRAPHICAL DISTRIBUTION—_continued_.
Distribution of fresh-water productions.
On the inhabitants of oceanic islands.
Absence of Batrachians and of terrestrial Mammals.
On the relation of the inhabitants of islands to those of the nearest
mainland.
On colonisation from the nearest source with subsequent modification.
Summary of the last and present chapters.
CHAPTER 13. MUTUAL AFFINITIES OF ORGANIC BEINGS: MORPHOLOGY:
EMBRYOLOGY: RUDIMENTARY ORGANS.
CLASSIFICATION, groups subordinate to groups.
Natural system.
Rules and difficulties in classification, explained on the theory of
descent with modification.
Classification of varieties.
Descent always used in classification.
Analogical or adaptive characters.
Affinities, general, complex and radiating.
Extinction separates and defines groups.
MORPHOLOGY, between members of the same class, between parts of the
same individual.
EMBRYOLOGY, laws of, explained by variations not supervening at an
early age, and being inherited at a corresponding age.
RUDIMENTARY ORGANS; their origin explained.
Summary.
CHAPTER 14. RECAPITULATION AND CONCLUSION.
Recapitulation of the difficulties on the theory of Natural
Selection.
Recapitulation of the general and special circumstances in its
favour.
Causes of the general belief in the immutability of species.
How far the theory of natural selection may be extended.
Effects of its adoption on the study of Natural history.
Concluding remarks.
ON THE ORIGIN OF SPECIES.
INTRODUCTION.
When on board H.M.S. ‘Beagle,’ as naturalist, I was much struck with
certain facts in the distribution of the inhabitants of South America,
and in the geological relations of the present to the past inhabitants
of that continent. These facts seemed to me to throw some light on the
origin of species—that mystery of mysteries, as it has been called by
one of our greatest philosophers. On my return home, it occurred to me,
in 1837, that something might perhaps be made out on this question by
patiently accumulating and reflecting on all sorts of facts which could
possibly have any bearing on it. After five years’ work I allowed
myself to speculate on the subject, and drew up some short notes; these
I enlarged in 1844 into a sketch of the conclusions, which then seemed
to me probable: from that period to the present day I have steadily
pursued the same object. I hope that I may be excused for entering on
these personal details, as I give them to show that I have not been
hasty in coming to a decision.
My work is now nearly finished; but as it will take me two or three
more years to complete it, and as my health is far from strong, I have
been urged to publish this Abstract. I have more especially been
induced to do this, as Mr. Wallace, who is now studying the
natural history of the Malay archipelago, has arrived at almost exactly
the same general conclusions that I have on the origin of species. Last
year he sent to me a memoir on this subject, with a request that I
would forward it to Sir Charles Lyell, who sent it to the Linnean
Society, and it is published in the third volume of the Journal of that
Society. Sir C. Lyell and Dr. Hooker, who both knew of my work—the
latter having read my sketch of 1844—honoured me by thinking it
advisable to publish, with Mr. Wallace’s excellent memoir, some brief
extracts from my manuscripts.
This Abstract, which I now publish, must necessarily be imperfect. I
cannot here give references and authorities for my several statements;
and I must trust to the reader reposing some confidence in my accuracy.
No doubt errors will have crept in, though I hope I have always been
cautious in trusting to good authorities alone. I can here give only
the general conclusions at which I have arrived, with a few facts in
illustration, but which, I hope, in most cases will suffice. No one can
feel more sensible than I do of the necessity of hereafter publishing
in detail all the facts, with references, on which my conclusions have
been grounded; and I hope in a future work to do this. For I am well
aware that scarcely a single point is discussed in this volume on which
facts cannot be adduced, often apparently leading to conclusions
directly opposite to those at which I have arrived. A fair result can
be obtained only by fully stating and balancing the facts and arguments
on both sides of each question; and this cannot possibly be here done.
I much regret that want of space prevents my having the satisfaction of
acknowledging the generous assistance which I have received from very
many naturalists, some of them personally unknown to me. I cannot,
however,
let this opportunity pass without expressing my deep obligations to Dr.
Hooker, who for the last fifteen years has aided me in every possible
way by his large stores of knowledge and his excellent judgment.
In considering the Origin of Species, it is quite conceivable that a
naturalist, reflecting on the mutual affinities of organic beings, on
their embryological relations, their geographical distribution,
geological succession, and other such facts, might come to the
conclusion that each species had not been independently created, but
had descended, like varieties, from other species. Nevertheless, such a
conclusion, even if well founded, would be unsatisfactory, until it
could be shown how the innumerable species inhabiting this world have
been modified, so as to acquire that perfection of structure and
coadaptation which most justly excites our admiration. Naturalists
continually refer to external conditions, such as climate, food, etc.,
as the only possible cause of variation. In one very limited sense, as
we shall hereafter see, this may be true; but it is preposterous to
attribute to mere external conditions, the structure, for instance, of
the woodpecker, with its feet, tail, beak, and tongue, so admirably
adapted to catch insects under the bark of trees. In the case of the
misseltoe, which draws its nourishment from certain trees, which has
seeds that must be transported by certain birds, and which has flowers
with separate sexes absolutely requiring the agency of certain insects
to bring pollen from one flower to the other, it is equally
preposterous to account for the structure of this parasite, with its
relations to several distinct organic beings, by the effects of
external conditions, or of habit, or of the volition of the plant
itself.
The author of the ‘Vestiges of Creation’ would, I presume, say that,
after a certain unknown number of
generations, some bird had given birth to a woodpecker, and some plant
to the misseltoe, and that these had been produced perfect as we now
see them; but this assumption seems to me to be no explanation, for it
leaves the case of the coadaptations of organic beings to each other
and to their physical conditions of life, untouched and unexplained.
It is, therefore, of the highest importance to gain a clear insight
into the means of modification and coadaptation. At the commencement of
my observations it seemed to me probable that a careful study of
domesticated animals and of cultivated plants would offer the best
chance of making out this obscure problem. Nor have I been
disappointed; in this and in all other perplexing cases I have
invariably found that our knowledge, imperfect though it be, of
variation under domestication, afforded the best 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 to variation. In the next chapter the Struggle for
Existence amongst all organic beings throughout the world, which
inevitably follows from their high geometrical powers of
increase, will be treated of. This is the doctrine of Malthus, applied
to the whole animal and vegetable kingdoms. As many more individuals of
each species are born than can possibly survive; and as, consequently,
there is a frequently recurring struggle for existence, it follows that
any being, if it vary however slightly in any manner profitable to
itself, under the complex and sometimes varying conditions of life,
will have a better chance of surviving, and thus be _naturally
selected_. From the strong principle of inheritance, any selected
variety will tend to propagate its new and modified form.
This fundamental subject of Natural Selection will be treated at some
length in the fourth chapter; and we shall then see how Natural
Selection almost inevitably causes much Extinction of the less improved
forms of life and induces what I have called Divergence of Character.
In the next chapter I shall discuss the complex and little known laws
of variation and of correlation of growth. In the four succeeding
chapters, the most apparent and gravest difficulties on the theory will
be given: namely, first, the difficulties of transitions, or in
understanding how a simple being or a simple organ can be changed and
perfected into a highly developed being or elaborately constructed
organ; secondly the subject of Instinct, or the mental powers of
animals, thirdly, Hybridism, or the infertility of species and the
fertility of varieties when intercrossed; and fourthly, the
imperfection of the Geological Record. In the next chapter I shall
consider the geological succession of organic beings throughout time;
in the eleventh and twelfth, their geographical distribution throughout
space; in the thirteenth, their classification or mutual affinities,
both when mature and in an embryonic condition. In the last chapter I
shall give a
brief recapitulation of the whole work, and a few concluding remarks.
No one ought to feel surprise at much remaining as yet unexplained in
regard to the origin of species and varieties, if he makes due
allowance for our profound ignorance in regard to the mutual relations
of all the beings which live around us. Who can explain why one species
ranges widely and is very numerous, and why another allied species has
a narrow range and is rare? Yet these relations are of the highest
importance, for they determine the present welfare, and, as I believe,
the future success and modification of every inhabitant of this world.
Still less do we know of the mutual relations of the innumerable
inhabitants of the world during the many past geological epochs in its
history. Although much remains obscure, and will long remain obscure, I
can entertain no doubt, after the most deliberate study and
dispassionate judgment of which I am capable, that the view which most
naturalists entertain, and which I formerly entertained—namely, that
each species has been independently created—is erroneous. I am fully
convinced that species are not immutable; but that those belonging to
what are called the same genera are lineal descendants of some other
and generally extinct species, in the same manner as the acknowledged
varieties of any one species are the descendants of that species.
Furthermore, I am convinced that Natural Selection has been the main
but not exclusive means of modification.
CHAPTER I.
VARIATION UNDER DOMESTICATION.
Causes of Variability. Effects of Habit. Correlation of Growth.
Inheritance. Character of Domestic Varieties. Difficulty of
distinguishing between Varieties and Species. Origin of Domestic
Varieties from one or more Species. Domestic Pigeons, their Differences
and Origin. Principle of Selection anciently followed, its Effects.
Methodical and Unconscious Selection. Unknown Origin of our Domestic
Productions. Circumstances favourable to Man’s power of Selection.
When we look to the individuals of the same variety or sub-variety of
our older cultivated plants and animals, one of the first points which
strikes us, is, that they generally differ much more from each other,
than do the individuals of any one species or variety in a state of
nature. When we reflect on the vast diversity of the plants and animals
which have been cultivated, and which have varied during all ages under
the most different climates and 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.
No case is on record of a variable being ceasing to be variable under
cultivation. Our oldest cultivated plants, such as wheat, still often
yield new varieties: our oldest domesticated animals are still capable
of rapid improvement or modification.
It has been disputed at what period of life the causes of variability,
whatever they may be, generally act; whether during the early or late
period of development of the embryo, or at the instant of conception.
Geoffroy St. Hilaire’s experiments show that unnatural treatment of the
embryo causes monstrosities; and monstrosities cannot be separated by
any clear line of distinction from mere variations. But I am strongly
inclined to suspect that the most frequent cause of variability may be
attributed to the male and female reproductive elements having been
affected prior to the act of conception. Several reasons make me
believe in this; but the chief one is the remarkable effect which
confinement or cultivation has on the functions of the reproductive
system; this system appearing to be far more susceptible than any other
part of the organisation, to the action of any change in the conditions
of life. Nothing is more easy than to tame an animal, and few things
more difficult than to get it to breed freely under confinement, even
in the many cases when the male and female unite. How many animals
there are which will not breed, though living long under not very close
confinement in their native country! This is generally attributed to
vitiated instincts; but how many cultivated plants display the utmost
vigour, and yet rarely or never seed! In some few such cases it has
been found out that very trifling changes, such as a little more or
less water at some particular period of growth, will determine whether
or not the plant sets a seed. I cannot here enter on the copious
details which I have collected on
this curious subject; but to show how singular the laws are which
determine the reproduction of animals under confinement, I may just
mention that carnivorous animals, even from the tropics, breed in this
country pretty freely under confinement, with the exception of the
plantigrades or bear family; whereas, carnivorous birds, with the
rarest exceptions, hardly ever lay fertile eggs. Many exotic plants
have pollen utterly worthless, in the same exact condition as in the
most sterile hybrids. When, on the one hand, we see domesticated
animals and plants, though often weak and sickly, yet breeding quite
freely under confinement; and when, on the other hand, we see
individuals, though taken young from a state of nature, perfectly
tamed, long-lived, and healthy (of which I could give numerous
instances), yet having their reproductive system so seriously affected
by unperceived causes as to fail in acting, we need not be surprised at
this system, when it does act under confinement, acting not quite
regularly, and producing offspring not perfectly like their parents or
variable.
Sterility has been said to be the bane of horticulture; but on this
view we 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.
Such buds can be propagated by grafting, etc., and sometimes by seed.
These “sports” are extremely rare under nature, but far from rare under
cultivation; and in this case we see that the treatment of the parent
has affected a bud or offset, and not the ovules or pollen. But it is
the opinion of most physiologists that there is no essential difference
between a bud and an ovule in their earliest stages of formation; so
that, in fact, “sports” support my view, that variability may be
largely attributed to the ovules or pollen, or to both, having been
affected by the treatment of the parent prior to the act of conception.
These cases anyhow show that variation is not necessarily connected, as
some authors have supposed, with the act of generation.
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 Müller 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 of any variation, we should attribute to the direct action
of heat, moisture, light, food, etc., is most difficult: my impression
is, that with animals such agencies have produced very little direct
effect, though apparently more in the case of plants. Under this point
of view, Mr. Buckman’s recent experiments on plants seem extremely
valuable. When all or nearly all the individuals exposed to certain
conditions are affected in the same way, the change at first appears to
be directly due to such conditions; but in some cases it can be shown
that quite opposite conditions produce
similar changes of structure. Nevertheless some slight amount of change
may, I think, be attributed to the direct action of the conditions of
life—as, in some cases, increased size from amount of food, colour from
particular kinds of food and from light, and perhaps the thickness of
fur from climate.
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 Canidæ—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 vertebræ 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 scutellæ 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 touch; the
oil-gland is quite aborted. Several other less distinct breeds might
have been specified.
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.
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 Columbidæ 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 Columbidæ, 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 encyclopædia.
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.
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
or modify most of our plants up to their present standard of usefulness
to man, we can understand how it is that neither Australia, the Cape of
Good Hope, nor any other region inhabited by quite uncivilised man, has
afforded us a single plant worth culture. It is not that these
countries, so rich in species, do not by a strange chance possess the
aboriginal stocks of any useful plants, but that the native plants have
not been improved by continued selection up to a standard of perfection
comparable with that given to the plants in countries anciently
civilised.
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
which are first given to him in some slight degree by nature. No man
would ever try to make a fantail, till he saw a pigeon with a tail
developed in some slight degree in an unusual manner, or a pouter till
he saw a pigeon with a crop of somewhat unusual size; and the more
abnormal or unusual any character was when it first appeared, the more
likely it would be to catch his attention. But to use such an
expression as trying to make a fantail, is, I have no doubt, in most
cases, utterly incorrect. The man who first selected a pigeon with a
slightly larger tail, never dreamed what the descendants of that pigeon
would become through long-continued, partly unconscious and partly
methodical selection. Perhaps the parent bird of all fantails had only
fourteen tail-feathers somewhat expanded, like the present Java
fantail, or like individuals of other and distinct breeds, in which as
many as seventeen tail-feathers have been counted. Perhaps the first
pouter-pigeon did not inflate its crop much more than the turbit now
does the upper part of its oesophagus,—a habit which is disregarded by
all fanciers, as it is not one of the points of the breed.
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
most fleeting of characters, have lately been exhibited as distinct at
our poultry-shows.
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 falls in fashion,—perhaps more in one
district than in another, according to the state of civilisation of the
inhabitants—slowly to add to the characteristic features of the breed,
whatever they may be. But the chance will be infinitely small of any
record having been preserved of such slow, varying, and insensible
changes.
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
sufficient, with extreme care, to allow of the accumulation of a large
amount of modification in almost any desired direction. But as
variations manifestly useful or pleasing to man appear only
occasionally, the chance of their appearance will be much increased by
a large number of individuals being kept; and hence this comes to be of
the highest importance to success. On this principle Marshall has
remarked, with respect to the sheep of parts of Yorkshire, that “as
they generally belong to poor people, and are mostly _in small lots_,
they never can be improved.” On the other hand, nurserymen, from
raising large stocks of the same plants, are generally far more
successful than amateurs in getting new and valuable varieties. The
keeping of a large number of individuals of a species in any country
requires that the species should be placed under favourable conditions
of life, so as to breed freely in that country. When the individuals of
any species are scanty, all the individuals, whatever their quality may
be, will generally be allowed to breed, and this will effectually
prevent selection. But probably the most important point of all, is,
that the animal or plant should be so highly useful to man, or so much
valued by him, that the closest attention should be paid to even the
slightest deviation in the qualities or structure of each individual.
Unless such attention be paid nothing can be effected. I have seen it
gravely remarked, that it was most fortunate that the strawberry began
to vary just when gardeners began to attend closely to this plant. No
doubt the strawberry had always varied since it was cultivated, but the
slight varieties had been neglected. As soon, however, as gardeners
picked out individual plants with slightly larger, earlier, or better
fruit, and raised seedlings from them, and again picked out the best
seedlings and bred from them, then, there appeared (aided by some
crossing with distinct species) those many admirable varieties of the
strawberry which have been raised during the last thirty or forty
years.
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, etc., 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 inherent and
necessary contingency, under all circumstances, with all organic
beings, as some authors have thought. The effects of variability are
modified by various degrees of inheritance and of reversion.
Variability is governed by many unknown laws, more especially by that
of correlation of growth. Something may be attributed to the direct
action of the conditions of life. Something must be attributed to use
and disuse. The final result is thus rendered infinitely complex. In
some cases, I do not doubt that the intercrossing of species,
aboriginally distinct, has played an important part in the origin of
our domestic productions. When in any country several domestic breeds
have once been established, their occasional intercrossing, with the
aid of selection, has, no doubt, largely aided in the formation of new
sub-breeds; but the importance of the crossing of varieties has, I
believe, been greatly exaggerated, both in regard to animals and to
those plants which are propagated by seed. In plants which are
temporarily propagated by cuttings, buds, etc., the importance of the
crossing both of distinct species and of varieties is immense; for the
cultivator here quite disregards the extreme variability both of
hybrids and mongrels, and the frequent sterility of hybrids; but the
cases of plants not propagated by seed are of little importance to us,
for their endurance is only temporary. Over all these causes of Change
I am convinced that the accumulative action of Selection, whether
applied methodically and more quickly, or unconsciously and more
slowly, but more efficiently, is by far the predominant Power.
CHAPTER II.
VARIATION UNDER NATURE.
Variability. Individual differences. Doubtful species. Wide ranging,
much diffused, and common species vary most. Species of the larger
genera in any country vary more than the species of the smaller genera.
Many of the species of the larger genera resemble varieties in being
very closely, but unequally, related to each other, and in having
restricted ranges.
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 plants on Alpine summits, or the thicker fur of
an animal from far northwards, would not in some cases be inherited for
at least some few generations? and in this case I presume that the form
would be called a variety.
Again, we have many slight differences which may be called individual
differences, such as are known frequently to appear in the offspring
from the same parents, or which may be presumed to have thus arisen,
from being frequently observed in the individuals of the same species
inhabiting the same confined locality. No one supposes that all the
individuals of the same species are cast in the very same mould. These
individual differences are highly important for us, as they afford
materials for natural selection to accumulate, in the same manner as
man can accumulate in any given direction individual differences in his
domesticated productions. These individual differences generally affect
what naturalists consider unimportant parts; but I could show by a long
catalogue of facts, that parts which must be called important, whether
viewed under a physiological or classificatory point of view, sometimes
vary in the individuals of the same species. I am convinced that the
most experienced naturalist would be surprised at the number of the
cases of variability, even in important parts of structure, which he
could collect on good authority, as I have collected, during a course
of years. It should be remembered that systematists are far from
pleased at finding variability in important characters, and that there
are not many men who will laboriously examine internal and important
organs, and compare them in many specimens of the same species. I
should never have expected that the branching of the main nerves close
to the great central ganglion of an insect would have been variable in
the same species; I should have expected that changes of this nature
could have been effected only by slow degrees: yet quite recently Mr.
Lubbock has shown a degree of variability in these main nerves in
Coccus, which may almost be compared to the irregular branching of the
stem of a tree. This philosophical naturalist, I may add, has also
quite recently shown that the muscles in the larvæ of certain insects
are very far from uniform. Authors sometimes argue in a circle when
they state that important organs never vary; for these same authors
practically rank that character as important (as some few naturalists
have honestly confessed) which does not vary; and, under this point of
view, no instance of an important part varying will ever be found: but
under any other point of view many instances assuredly can be given.
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 because the
intermediate links have actually been found, but because analogy leads
the observer to suppose either that they do now somewhere exist, or may
formerly have existed; and here a wide door for the entry of doubt and
conjecture is opened.
Hence, in determining whether a form should be ranked as a species or a
variety, the opinion of naturalists having sound judgment 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 islets of the little Madeira group there
are many insects which are characterized as varieties in Mr.
Wollaston’s admirable work, but which it cannot
be doubted would be ranked as distinct species by many entomologists.
Even Ireland has a few animals, now generally regarded as varieties,
but which have been ranked as species by some zoologists. Several most
experienced ornithologists consider our British red grouse as only a
strongly-marked race of a Norwegian species, whereas the greater number
rank it as an undoubted species peculiar to Great Britain. A wide
distance between the homes of two doubtful forms leads many naturalists
to rank both as distinct species; but what distance, it has been well
asked, will suffice? if that between America and Europe is ample, will
that between the Continent and the Azores, or Madeira, or the Canaries,
or Ireland, be sufficient? It must be admitted that many forms,
considered by highly-competent judges as varieties, have so perfectly
the character of species that they are ranked by other highly-competent
judges as good and true species. But to discuss whether they are
rightly called species or varieties, before any definition of these
terms has been generally accepted, is vainly to beat the air.
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, etc., 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
that most careful observer Gärtner, they can be crossed only with much
difficulty. We could hardly wish for better evidence of the two forms
being specifically distinct. On the other hand, they are united by many
intermediate links, and it is very doubtful whether these links are
hybrids; and there is, as it seems to me, an overwhelming amount of
experimental evidence, showing that they descend from common parents,
and consequently must be ranked as varieties.
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
general tendency will be to make many species, for he will become
impressed, just like the pigeon or poultry-fancier before alluded to,
with the amount of difference in the forms which he is continually
studying; and he has little general knowledge of analogical variation
in other groups and in other countries, by which to correct his first
impressions. As he extends the range of his observations, he will meet
with more cases of difficulty; for he will encounter a greater number
of closely-allied forms. But if his observations be widely extended, he
will in the end generally be enabled to make up his own mind which to
call varieties and which species; but he will succeed in this at the
expense of admitting much variation,—and the truth of this admission
will often be disputed by other naturalists. When, moreover, he comes
to study allied forms brought from countries not now continuous, in
which case he can hardly hope to find the intermediate links between
his doubtful forms, he will have to trust almost entirely to analogy,
and his difficulties will rise to a climax.
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
strongly marked and more permanent varieties; and at these latter, as
leading to sub-species, and to species. The passage from one stage of
difference to another and higher stage may be, in some cases, due
merely to the long-continued action of different physical conditions in
two different regions; but I have not much faith in this view; and I
attribute the passage of a variety, from a state in which it differs
very slightly from its parent to one in which it differs more, to the
action of natural selection in accumulating (as will hereafter be more
fully explained) differences of structure in certain definite
directions. Hence I believe a well-marked variety may be justly called
an incipient species; but whether this belief be justifiable must be
judged of by the general weight of the several facts and views given
throughout this work.
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 the parent species,
it would then rank as the species, and the species as the variety; or
it might come to supplant and exterminate the parent species; or both
might co-exist, and both rank as independent species. But we shall
hereafter have to return to this subject.
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 a
different consideration from wide range, and to a certain extent from
commonness), often give rise to varieties sufficiently well-marked to
have been recorded in botanical works. Hence it is the most
flourishing, or, as they may be called, the dominant species,—those
which range widely over the world, are the most diffused in their own
country, and are the most numerous in individuals,—which oftenest
produce well-marked varieties, or, as I consider them, incipient
species. And this, perhaps, might have been anticipated; for, as
varieties, in order to become in any degree permanent, necessarily have
to struggle with the other inhabitants of the country, the species
which are already dominant will be the most likely to yield offspring
which, though in some slight degree modified, will still inherit those
advantages that enabled their parents to become dominant over their
compatriots.
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 species
belong. Again, plants low in the scale of organisation are
generally much more widely diffused than plants higher in the scale;
and here again there is no close relation to the size of the genera.
The cause of lowly-organised plants ranging widely will be discussed in
our chapter on geographical distribution.
From looking at species as only strongly-marked and well-defined
varieties, I was led to anticipate that the species of the larger
genera in each country would oftener present varieties, than the
species of the smaller genera; for wherever many closely related
species (_i.e._ species of the same genus) have been formed, many
varieties or incipient species ought, as a general rule, to be now
forming. Where many large trees grow, we expect to find saplings. Where
many species of a genus have been formed through variation,
circumstances have been favourable for variation; and hence we might
expect that the circumstances would generally be still favourable to
variation. On the other hand, if we look at each species as a special
act of creation, there is no apparent reason why more varieties should
occur in a group having many species, than in one having few.
To test the truth of this anticipation I have arranged the plants of
twelve countries, and the coleopterous insects of two districts, into
two nearly equal masses, the species of the larger genera on one side,
and those of the smaller genera on the other side, and it has
invariably proved to be the case that a larger proportion of the
species on the side of the larger genera present varieties, than on the
side of the smaller genera. Moreover, the species of the large genera
which present any varieties, invariably present a larger average number
of varieties than do the species of the small genera. Both these
results follow when another division is made, and when all the smallest
genera, with from only one to four species, are absolutely excluded
from the tables. These
facts are of plain signification on the view that species are only
strongly marked and permanent varieties; for wherever many species of
the same genus have been formed, or where, if we may use the
expression, the manufactory of species has been active, we ought
generally to find the manufactory still in action, more especially as
we have every reason to believe the process of manufacturing new
species to be a slow one. And this certainly is the case, if varieties
be looked at as incipient species; for my tables clearly show as a
general rule that, wherever many species of a genus have been formed,
the species of that genus present a number of varieties, that is of
incipient species, beyond the average. It is not that all large genera
are now varying much, and are thus increasing in the number of their
species, or that no small genera are now varying and increasing; for if
this had been so, it would have been fatal to my theory; inasmuch as
geology plainly tells us that small genera have in the lapse of time
often increased greatly in size; and that large genera have often come
to their maxima, declined, and disappeared. All that we want to show
is, that where many species of a genus have been formed, on an average
many are still forming; and this holds good.
There are other relations between the species of large genera and their
recorded varieties which deserve notice. We have seen that there is no
infallible criterion by which to distinguish species and well-marked
varieties; and in those cases in which intermediate links have not been
found between doubtful forms, naturalists are compelled to come to a
determination by the amount of difference between them, judging by
analogy whether or not the amount suffices to raise one or both to the
rank of species. Hence the amount of difference is one very important
criterion in settling whether two forms should
be ranked as species or varieties. Now Fries has remarked in regard to
plants, and Westwood in regard to insects, that in large genera the
amount of difference between the species is often exceedingly small. I
have endeavoured to test this numerically by averages, and, as far as
my imperfect results go, they always confirm the view. I have also
consulted some sagacious and most experienced observers, and, after
deliberation, they concur in this view. In this respect, therefore, the
species of the larger genera resemble varieties, more than do the
species of the smaller genera. Or the case may be put in another way,
and it may be said, that in the larger genera, in which a number of
varieties or incipient species greater than the average are now
manufacturing, many of the species already manufactured still to a
certain extent resemble varieties, for they differ from each other by a
less than usual amount of difference.
Moreover, the species of the large genera are related to each other, in
the same manner as the varieties of any one species are related to each
other. No naturalist pretends that all the species of a genus are
equally distinct from each other; they may generally be divided into
sub-genera, or sections, or lesser groups. As Fries has well remarked,
little groups of species are generally clustered like satellites around
certain other species. And what are varieties but groups of forms,
unequally related to each other, and clustered round certain forms—that
is, round their parent-species? Undoubtedly there is one most important
point of difference between varieties and species; namely, that the
amount of difference between varieties, when compared with each other
or with their parent-species, is much less than that between the
species of the same genus. But when we come to discuss the principle,
as I call it, of Divergence of Character,
we shall see how this may be explained, and how the lesser differences
between varieties will tend to increase into the greater differences
between species.
There is one other point which seems to me worth notice. Varieties
generally have much restricted ranges: this statement is indeed
scarcely more than a truism, for if a variety were found to have a
wider range than that of its supposed parent-species, their
denominations ought to be reversed. But there is also reason to
believe, that those species which are very closely allied to other
species, and in so far resemble varieties, often have much restricted
ranges. For instance, Mr. H. C. Watson has marked for me in the
well-sifted London Catalogue of plants (4th edition) 63 plants which
are therein ranked as species, but which he considers as so closely
allied to other species as to be of doubtful value: these 63 reputed
species range on an average over 6.9 of the provinces into which Mr.
Watson has divided Great Britain. Now, in this same catalogue, 53
acknowledged varieties are recorded, and these range over 7.7
provinces; whereas, the species to which these varieties belong range
over 14.3 provinces. So that the acknowledged varieties have very
nearly the same restricted average range, as have those very closely
allied forms, marked for me by Mr. Watson as doubtful species, but
which are almost universally ranked by British botanists as good and
true species.
Finally, then, varieties have the same general characters as species,
for they cannot be distinguished from species,—except, firstly, by the
discovery of intermediate linking forms, and the occurrence of such
links cannot affect the actual characters of the forms which they
connect; and except, secondly, by a certain amount of
difference, for two forms, if differing very little, are generally
ranked as varieties, notwithstanding that intermediate linking forms
have not been discovered; but the amount of difference considered
necessary to give to two forms the rank of species is quite indefinite.
In genera having more than the average number of species in any
country, the species of these genera have more than the average number
of varieties. In large genera the species are apt to be closely, but
unequally, allied together, forming little clusters round certain
species. Species very closely allied to other species apparently have
restricted ranges. In all these several respects the species of large
genera present a strong analogy with varieties. And we can clearly
understand these analogies, if species have once existed as varieties,
and have thus originated: whereas, these analogies are utterly
inexplicable if each species has been independently created.
We have, also, seen that it is the most flourishing and dominant
species of the larger genera which on an average vary most; and
varieties, as we shall hereafter see, tend to become converted into new
and distinct species. The larger genera thus tend to become larger; and
throughout nature the forms of life which are now dominant tend to
become still more dominant by leaving many modified and dominant
descendants. But by steps hereafter to be explained, the larger genera
also tend to break up into smaller genera. And thus, the forms of life
throughout the universe become divided into groups subordinate to
groups.
CHAPTER III.
STRUGGLE FOR EXISTENCE.
Bears on natural selection. The term used in a wide sense. Geometrical
powers of increase. Rapid increase of naturalised animals and plants.
Nature of the checks to increase. Competition universal. Effects of
climate. Protection from the number of individuals. Complex relations
of all animals and plants throughout nature. Struggle for life most
severe between individuals and varieties of the same species; often
severe between species of the same genus. The relation of organism to
organism the most important of all relations.
Before entering on the subject of this chapter, I must make a few
preliminary remarks, to show how the struggle for existence bears on
Natural Selection. It has been seen in the last chapter that amongst
organic beings in a state of nature there is some individual
variability; indeed I am not aware that this has ever been disputed. It
is immaterial for us whether a multitude of doubtful forms be called
species or sub-species or varieties; what rank, for instance, the two
or three hundred doubtful forms of British plants are entitled to hold,
if the existence of any well-marked varieties be admitted. But the mere
existence of individual variability and of some few well-marked
varieties, though necessary as the foundation for the work, helps us
but little in understanding how species arise in nature. How have all
those exquisite adaptations of one part of the organisation to another
part, and to the conditions of life, and of one distinct organic being
to another being, been perfected? We see these beautiful co-adaptations
most plainly in the woodpecker and missletoe; and only a little less
plainly in the humblest parasite which clings
to the hairs of a quadruped or feathers of a bird; in the structure of
the beetle which dives through the water; in the plumed seed which is
wafted by the gentlest breeze; in short, we see beautiful adaptations
everywhere and in every part of the organic world.
Again, it may be asked, how is it that varieties, which I have called
incipient species, become ultimately converted into good and distinct
species, which in most cases obviously differ from each other far more
than do the varieties of the same species? How do those groups of
species, which constitute what are called distinct genera, and which
differ from each other more than do the species of the same genus,
arise? All these results, as we shall more fully see in the next
chapter, follow inevitably from the struggle for life. Owing to this
struggle for life, any variation, however slight and from whatever
cause proceeding, if it be in any degree profitable to an individual of
any species, in its infinitely complex relations to other organic
beings and to external nature, will tend to the preservation of that
individual, and will generally be inherited by its offspring. The
offspring, also, will thus have a better chance of surviving, for, of
the many individuals of any species which are periodically born, but a
small number can survive. I have called this principle, by which each
slight variation, if useful, is preserved, by the term of Natural
Selection, in order to mark its relation to man’s power of selection.
We have seen that man by selection can certainly produce great results,
and can adapt organic beings to his own uses, through the accumulation
of slight but useful variations, given to him by the hand of Nature.
But Natural Selection, as we shall hereafter see, is a power
incessantly ready for action, and is as immeasurably superior to man’s
feeble efforts, as the works of Nature are to those of Art.
We will now discuss in a little more detail the struggle for existence.
In my future work this subject shall be treated, as it well deserves,
at much greater length. The elder De Candolle and Lyell have largely
and philosophically shown that all organic beings are exposed to severe
competition. In regard to plants, no one has treated this subject with
more spirit and ability than W. Herbert, Dean of Manchester, evidently
the result of his great horticultural knowledge. Nothing is easier than
to admit in words the truth of the universal struggle for life, or more
difficult—at least I have found it so—than constantly to bear this
conclusion in mind. Yet unless it be thoroughly engrained in the mind,
I am convinced that the whole economy of nature, with every fact on
distribution, rarity, abundance, extinction, and variation, will be
dimly seen or quite misunderstood. We behold the face of nature bright
with gladness, we often see superabundance of food; we do not see, or
we forget, that the birds which are idly singing round us mostly live
on insects or seeds, and are thus constantly destroying life; or we
forget how largely these songsters, or their eggs, or their nestlings,
are destroyed by birds and beasts of prey; we do not always bear in
mind, that though food may be now superabundant, it is not so at all
seasons of each recurring year.
I should premise that I use the term Struggle for Existence in a large
and metaphorical sense, including dependence of one being on another,
and including (which is more important) not only the life of the
individual, but success in leaving progeny. Two canine animals in a
time of dearth, may be truly said to struggle with each other which
shall get food and live. But a plant on the edge of a desert is said to
struggle for life against the drought, though more properly it should
be said to be dependent on the moisture. A
plant which annually produces a thousand seeds, of which on an average
only one comes to maturity, may be more truly said to struggle with the
plants of the same and other kinds which already clothe the ground. The
missletoe is dependent on the apple and a few other trees, but can only
in a far-fetched sense be said to struggle with these trees, for if too
many of these parasites grow on the same tree, it will languish and
die. But several seedling missletoes, growing close together on the
same branch, may more truly be said to struggle with each other. As the
missletoe is disseminated by birds, its existence depends on birds; and
it may metaphorically be said to struggle with other fruit-bearing
plants, in order to tempt birds to devour and thus disseminate its
seeds rather than those of other plants. In these several senses, which
pass into each other, I use for convenience sake the general term of
struggle for existence.
A struggle for existence inevitably follows from the high rate at which
all organic beings tend to increase. Every being, which during its
natural lifetime produces several eggs or seeds, must suffer
destruction during some period of its life, and during some season or
occasional year, otherwise, on the principle of geometrical increase,
its numbers would quickly become so inordinately great that no country
could support the product. Hence, as more individuals are produced than
can possibly survive, there must in every case be a struggle for
existence, either one individual with another of the same species, or
with the individuals of distinct species, or with the physical
conditions of life. It is the doctrine of Malthus applied with manifold
force to the whole animal and vegetable kingdoms; for in this case
there can be no artificial increase of food, and no prudential
restraint from marriage. Although some species may be
now increasing, more or less rapidly, in numbers, all cannot do so, for
the world would not hold them.
There is no exception to the rule that every organic being naturally
increases at so high a rate, that if not destroyed, the earth would
soon be covered by the progeny of a single pair. Even slow-breeding man
has doubled in twenty-five years, and at this rate, in a few thousand
years, there would literally not be standing room for his progeny.
Linnæus has calculated that if an annual plant produced only two
seeds—and there is no plant so unproductive as this—and their seedlings
next year produced two, and so on, then in twenty years there would be
a million plants. The elephant is reckoned to be the slowest breeder of
all known animals, and I have taken some pains to estimate its probable
minimum rate of natural increase: it will be under the mark to assume
that it breeds when thirty years old, and goes on breeding till ninety
years old, bringing forth three pair of young in this interval; if this
be so, at the end of the fifth century there would be alive fifteen
million elephants, descended from the first pair.
But we have better evidence on this subject than mere theoretical
calculations, namely, the numerous recorded cases of the astonishingly
rapid increase of various animals in a state of nature, when
circumstances have been favourable to them during two or three
following seasons. Still more striking is the evidence from our
domestic animals of many kinds which have run wild in several parts of
the world: if the statements of the rate of increase of slow-breeding
cattle and horses in South America, and latterly in Australia, had not
been well authenticated, they would have been quite incredible. So it
is with plants: cases could be given of introduced plants which have
become common throughout whole islands in a period of less than ten
years. Several
of the plants now most numerous over the wide plains of La Plata,
clothing square leagues of surface almost to the exclusion of all other
plants, have been introduced from Europe; and there are plants which
now range in India, as I hear from Dr. Falconer, from Cape Comorin to
the Himalaya, which have been imported from America since its
discovery. In such cases, and endless instances could be given, no one
supposes that the fertility of these animals or plants has been
suddenly and temporarily increased in any sensible degree. The obvious
explanation is that the conditions of life have been very favourable,
and that there has consequently been less destruction of the old and
young, and that nearly all the young have been enabled to breed. In
such cases the geometrical ratio of increase, the result of which never
fails to be surprising, simply explains the extraordinarily rapid
increase and wide diffusion of naturalised productions in their new
homes.
In a state of nature almost every plant produces seed, and amongst
animals there are very few which do not annually pair. Hence we may
confidently assert, that all plants and animals are tending to increase
at a geometrical ratio, that all would most rapidly stock every station
in which they could any how exist, and that the geometrical tendency to
increase must be checked by destruction at some period of life. Our
familiarity with the larger domestic animals tends, I think, to mislead
us: we see no great destruction falling on them, and we forget that
thousands are annually slaughtered for food, and that in a state of
nature an equal number would have somehow to be disposed of.
The only difference between organisms which annually produce eggs or
seeds by the thousand, and those which produce extremely few, is, that
the slow-breeders would require a few more years to people, under
favourable
conditions, a whole district, let it be ever so large. The condor lays
a couple of eggs and the ostrich a score, and yet in the same country
the condor may be the more numerous of the two: the Fulmar petrel lays
but one egg, yet it is believed to be the most numerous bird in the
world. One fly deposits hundreds of eggs, and another, like the
hippobosca, a single one; but this difference does not determine how
many individuals of the two species can be supported in a district. A
large number of eggs is of some importance to those species, which
depend on a rapidly fluctuating amount of food, for it allows them
rapidly to increase in number. But the real importance of a large
number of eggs or seeds is to make up for much destruction at some
period of life; and this period in the great majority of cases is an
early one. If an animal can in any way protect its own eggs or young, a
small number may be produced, and yet the average stock be fully kept
up; but if many eggs or young are destroyed, many must be produced, or
the species will become extinct. It would suffice to keep up the full
number of a tree, which lived on an average for a thousand years, if a
single seed were produced once in a thousand years, supposing that this
seed were never destroyed, and could be ensured to germinate in a
fitting place. So that in all cases, the average number of any animal
or plant depends only indirectly on the number of its eggs or seeds.
In looking at Nature, it is most necessary to keep the foregoing
considerations always in mind—never to forget that every single organic
being around us may be said to be striving to the utmost to increase in
numbers; that each lives by a struggle at some period of its life; that
heavy destruction inevitably falls either on the young or old, during
each generation or at recurrent intervals. Lighten any check, mitigate
the
destruction ever so little, and the number of the species will almost
instantaneously increase to any amount. The face of Nature may be
compared to a yielding surface, with ten thousand sharp wedges packed
close together and driven inwards by incessant blows, sometimes one
wedge being struck, and then another with greater force.
What checks the natural tendency of each species to increase in number
is most obscure. Look at the most vigorous species; by as much as it
swarms in numbers, by so much will its tendency to increase be still
further increased. We know not exactly what the checks are in even one
single instance. Nor will this surprise any one who reflects how
ignorant we are on this head, even in regard to mankind, so
incomparably better known than any other animal. This subject has been
ably treated by several authors, and I shall, in my future work,
discuss some of the checks at considerable length, more especially in
regard to the feral animals of South America. Here I will make only a
few remarks, just to recall to the reader’s mind some of the chief
points. Eggs or very young animals seem generally to suffer most, but
this is not invariably the case. With plants there is a vast
destruction of seeds, but, from some observations which I have made, I
believe that it is the seedlings which suffer most from germinating in
ground already thickly stocked with other plants. Seedlings, also, are
destroyed in vast numbers by various enemies; for instance, on a piece
of ground three feet long and two wide, dug and cleared, and where
there could be no choking from other plants, I marked all the seedlings
of our native weeds as they came up, and out of the 357 no less than
295 were destroyed, chiefly by slugs and insects. If turf which has
long been mown, and the case would be the same with turf closely
browsed by quadrupeds, be let to grow,
the more vigorous plants gradually kill the less vigorous, though fully
grown, plants: thus out of twenty species growing on a little plot of
turf (three feet by four) nine species perished from the other species
being allowed to grow up freely.
The amount of food for each species of course gives the extreme limit
to which each can increase; but very frequently it is not the obtaining
food, but the serving as prey to other animals, which determines the
average numbers of a species. Thus, there seems to be little doubt that
the stock of partridges, grouse, and hares on any large estate depends
chiefly on the destruction of vermin. If not one head of game were shot
during the next twenty years in England, and, at the same time, if no
vermin were destroyed, there would, in all probability, be less game
than at present, although hundreds of thousands of game animals are now
annually killed. On the other hand, in some cases, as with the elephant
and rhinoceros, none are destroyed by beasts of prey: even the tiger in
India most rarely dares to attack a young elephant protected by its
dam.
Climate plays an important part in determining the average numbers of a
species, and periodical seasons of extreme cold or drought, I believe
to be the most effective of all checks. I estimated that the winter of
1854-55 destroyed four-fifths of the birds in my own grounds; and this
is a tremendous destruction, when we remember that ten per cent. is an
extraordinarily severe mortality from epidemics with man. The action of
climate seems at first sight to be quite independent of the struggle
for existence; but in so far as climate chiefly acts in reducing food,
it brings on the most severe struggle between the individuals, whether
of the same or of distinct species, which subsist on the same kind of
food. Even when climate, for instance extreme
cold, acts directly, it will be the least vigorous, or those which have
got least food through the advancing winter, which will suffer most.
When we travel from south to north, or from a damp region to a dry, we
invariably see some species gradually getting rarer and rarer, and
finally disappearing; and the change of climate being conspicuous, we
are tempted to attribute the whole effect to its direct action. But
this is a very false view: we forget that each species, even where it
most abounds, is constantly suffering enormous destruction at some
period of its life, from enemies or from competitors for the same place
and food; and if these enemies or competitors be in the least degree
favoured by any slight change of climate, they will increase in
numbers, and, as each area is already fully stocked with inhabitants,
the other species will decrease. When we travel southward and see a
species decreasing in numbers, we may feel sure that the cause lies
quite as much in other species being favoured, as in this one being
hurt. So it is when we travel northward, but in a somewhat lesser
degree, for the number of species of all kinds, and therefore of
competitors, decreases northwards; hence in going northward, or in
ascending a mountain, we far oftener meet with stunted forms, due to
the _directly_ injurious action of climate, than we do in proceeding
southwards or in descending a mountain. When we reach the Arctic
regions, or snow-capped summits, or absolute deserts, the struggle for
life is almost exclusively with the elements.
That climate acts in main part indirectly by favouring other species,
we may clearly see in the prodigious number of plants in our gardens
which can perfectly well endure our climate, but which never become
naturalised, for they cannot compete with our native plants, nor resist
destruction by our native animals.
When a species, owing to highly favourable circumstances, increases
inordinately in numbers in a small tract, epidemics—at least, this
seems generally to occur with our game animals—often ensue: and here we
have a limiting check independent of the struggle for life. But even
some of these so-called epidemics appear to be due to parasitic worms,
which have from some cause, possibly in part through facility of
diffusion amongst the crowded animals, been disproportionably favoured:
and here comes in a sort of struggle between the parasite and its prey.
On the other hand, in many cases, a large stock of individuals of the
same species, relatively to the numbers of its enemies, is absolutely
necessary for its preservation. Thus we can easily raise plenty of corn
and rape-seed, etc., in our fields, because the seeds are in great
excess compared with the number of birds which feed on them; nor can
the birds, though having a superabundance of food at this one season,
increase in number proportionally to the supply of seed, as their
numbers are checked during winter: but any one who has tried, knows how
troublesome it is to get seed from a few wheat or other such plants in
a garden; I have in this case lost every single seed. This view of the
necessity of a large stock of the same species for its preservation,
explains, I believe, some singular facts in nature, such as that of
very rare plants being sometimes extremely abundant in the few spots
where they do occur; and that of some social plants being social, that
is, abounding in individuals, even on the extreme confines of their
range. For in such cases, we may believe, that a plant could exist only
where the conditions of its life were so favourable that many could
exist together, and thus save each other from utter destruction. I
should add that the good effects of frequent intercrossing, and the ill
effects
of close interbreeding, probably come into play in some of these cases;
but on this intricate subject I will not here enlarge.
Many cases are on record showing how complex and unexpected are the
checks and relations between organic beings, which have to struggle
together in the same country. I will give only a single instance,
which, though a simple one, has interested me. In Staffordshire, on the
estate of a relation where I had ample means of investigation, there
was a large and extremely barren heath, which had never been touched by
the hand of man; but several hundred acres of exactly the same nature
had been enclosed twenty-five years previously and planted with Scotch
fir. The change in the native vegetation of the planted part of the
heath was most remarkable, more than is generally seen in passing from
one quite different soil to another: not only the proportional numbers
of the heath-plants were wholly changed, but twelve species of plants
(not counting grasses and carices) flourished in the plantations, which
could not be found on the heath. The effect on the insects must have
been still greater, for six insectivorous birds were very common in the
plantations, which were not to be seen on the heath; and the heath was
frequented by two or three distinct insectivorous birds. Here we see
how potent has been the effect of the introduction of a single tree,
nothing whatever else having been done, with the exception that the
land had been enclosed, so that cattle could not enter. But how
important an element enclosure is, I plainly saw near Farnham, in
Surrey. Here there are extensive heaths, with a few clumps of old
Scotch firs on the distant hill-tops: within the last ten years large
spaces have been enclosed, and self-sown firs are now springing up in
multitudes, so close together that all cannot live.
When I ascertained that these young trees had not been sown or planted,
I was so much surprised at their numbers that I went to several points
of view, whence I could examine hundreds of acres of the unenclosed
heath, and literally I could not see a single Scotch fir, except the
old planted clumps. But on looking closely between the stems of the
heath, I found a multitude of seedlings and little trees, which had
been perpetually browsed down by the cattle. In one square yard, at a
point some hundred yards distant from one of the old clumps, I counted
thirty-two little trees; and one of them, judging from the rings of
growth, had during twenty-six years tried to raise its head above the
stems of the heath, and had failed. No wonder that, as soon as the land
was enclosed, it became thickly clothed with vigorously growing young
firs. Yet the heath was so extremely barren and so extensive that no
one would ever have imagined that cattle would have so closely and
effectually searched it for food.
Here we see that cattle absolutely determine the existence of the
Scotch fir; but in several parts of the world insects determine the
existence of cattle. Perhaps Paraguay offers the most curious instance
of this; for here neither cattle nor horses nor dogs have ever run
wild, though they swarm southward and northward in a feral state; and
Azara and Rengger have shown that this is caused by the greater number
in Paraguay of a certain fly, which lays its eggs in the navels of
these animals when first born. The increase of these flies, numerous as
they are, must be habitually checked by some means, probably by birds.
Hence, if certain insectivorous birds (whose numbers are probably
regulated by hawks or beasts of prey) were to increase in Paraguay, the
flies would decrease—then cattle and horses would become feral, and
this would certainly greatly alter (as
indeed I have observed in parts of South America) the vegetation: this
again would largely affect the insects; and this, as we just have seen
in Staffordshire, the insectivorous birds, and so onwards in
ever-increasing circles of complexity. We began this series by
insectivorous birds, and we have ended with them. Not that in nature
the relations can ever be as simple as this. Battle within battle must
ever be recurring with varying success; and yet in the long-run the
forces are so nicely balanced, that the face of nature remains uniform
for long periods of time, though assuredly the merest trifle would
often give the victory to one organic being over another. Nevertheless
so profound is our ignorance, and so high our presumption, that we
marvel when we hear of the extinction of an organic being; and as we do
not see the cause, we invoke cataclysms to desolate the world, or
invent laws on the duration of the forms of life!
I am tempted to give one more instance showing how plants and animals,
most remote in the scale of nature, are bound together by a web of
complex relations. I shall hereafter have occasion to show that the
exotic Lobelia fulgens, in this part of England, is never visited by
insects, and consequently, from its peculiar structure, never can set a
seed. Many of our orchidaceous plants absolutely require the visits of
moths to remove their pollen-masses and thus to fertilise them. I have,
also, reason to believe that humble-bees are indispensable to the
fertilisation of the heartsease (Viola tricolor), for other bees do not
visit this flower. From experiments which I have tried, I have found
that the visits of bees, if not indispensable, are at least highly
beneficial to the fertilisation of our clovers; but humble-bees alone
visit the common red clover (Trifolium pratense), as other bees cannot
reach the nectar. Hence I have very little doubt, that if the whole
genus of humble-bees became
extinct or very rare in England, the heartsease and red clover would
become very rare, or wholly disappear. The number of humble-bees in any
district depends in a great degree on the number of field-mice, which
destroy their combs and nests; and Mr. H. Newman, who has long attended
to the habits of humble-bees, believes that “more than two thirds of
them are thus destroyed all over England.” Now the number of mice is
largely dependent, as every one knows, on the number of cats; and Mr.
Newman says, “Near villages and small towns I have found the nests of
humble-bees more numerous than elsewhere, which I attribute to the
number of cats that destroy the mice.” Hence it is quite credible that
the presence of a feline animal in large numbers in a district might
determine, through the intervention first of mice and then of bees, the
frequency of certain flowers in that district!
In the case of every species, many different checks, acting at
different periods of life, and during different seasons or years,
probably come into play; some one check or some few being generally the
most potent, but all concurring in determining the average number or
even the existence of the species. In some cases it can be shown that
widely-different checks act on the same species in different districts.
When we look at the plants and bushes clothing an entangled bank, we
are tempted to attribute their proportional numbers and kinds to what
we call chance. But how false a view is this! Every one has heard that
when an American forest is cut down, a very different vegetation
springs up; but it has been observed that the trees now growing on the
ancient Indian mounds, in the Southern United States, display the same
beautiful diversity and proportion of kinds as in the surrounding
virgin forests. What a struggle between the several kinds of trees
must here have gone on during long centuries, each annually scattering
its seeds by the thousand; what war between insect and insect—between
insects, snails, and other animals with birds and beasts of prey—all
striving to increase, and all feeding on each other or on the trees or
their seeds and seedlings, or on the other plants which first clothed
the ground and thus checked the growth of the trees! Throw up a handful
of feathers, and all must fall to the ground according to definite
laws; but how simple is this problem compared to the action and
reaction of the innumerable plants and animals which have determined,
in the course of centuries, the proportional numbers and kinds of trees
now growing on the old Indian ruins!
The dependency of one organic being on another, as of a parasite on its
prey, lies generally between beings remote in the scale of nature. This
is often the case with those which may strictly be said to struggle
with each other for existence, as in the case of locusts and
grass-feeding quadrupeds. But the struggle almost invariably will be
most severe between the individuals of the same species, for they
frequent the same districts, require the same food, and are exposed to
the same dangers. In the case of varieties of the same species, the
struggle will generally be almost equally severe, and we sometimes see
the contest soon decided: for instance, if several varieties of wheat
be sown together, and the mixed seed be resown, some of the varieties
which best suit the soil or climate, or are naturally the most fertile,
will beat the others and so yield more seed, and will consequently in a
few years quite supplant the other varieties. To keep up a mixed stock
of even such extremely close varieties as the variously coloured
sweet-peas, they must be each year harvested separately, and the seed
then mixed in due proportion,
otherwise the weaker kinds will steadily decrease in numbers and
disappear. So again with the varieties of sheep: it has been asserted
that certain mountain-varieties will starve out other
mountain-varieties, so that they cannot be kept together. The same
result has followed from keeping together different varieties of the
medicinal leech. It may even be doubted whether the varieties of any
one of our domestic plants or animals have so exactly the same
strength, habits, and constitution, that the original proportions of a
mixed stock could be kept up for half a dozen generations, if they were
allowed to struggle together, like beings in a state of nature, and if
the seed or young were not annually sorted.
As species of the same genus have usually, though by no means
invariably, some similarity in habits and constitution, and always in
structure, the struggle will generally be more severe between species
of the same genus, when they come into competition with each other,
than between species of distinct genera. We see this in the recent
extension over parts of the United States of one species of swallow
having caused the decrease of another species. The recent increase of
the missel-thrush in parts of Scotland has caused the decrease of the
song-thrush. How frequently we hear of one species of rat taking the
place of another species under the most different climates! In Russia
the small Asiatic cockroach has everywhere driven before it its great
congener. One species of charlock will supplant another, and so in
other cases. We can dimly see why the competition should be most severe
between allied forms, which fill nearly the same place in the economy
of nature; but probably in no one case could we precisely say why one
species has been victorious over another in the great battle of life.
A corollary of the highest importance may be deduced from the foregoing
remarks, namely, that the structure of every organic being is related,
in the most essential yet often hidden manner, to that of all other
organic beings, with which it comes into competition for food or
residence, or from which it has to escape, or on which it preys. This
is obvious in the structure of the teeth and talons of the tiger; and
in that of the legs and claws of the parasite which clings to the hair
on the tiger’s body. But in the beautifully plumed seed of the
dandelion, and in the flattened and fringed legs of the water-beetle,
the relation seems at first confined to the elements of air and water.
Yet the advantage of plumed seeds no doubt stands in the closest
relation to the land being already thickly clothed by other plants; so
that the seeds may be widely distributed and fall on unoccupied ground.
In the water-beetle, the structure of its legs, so well adapted for
diving, allows it to compete with other aquatic insects, to hunt for
its own prey, and to escape serving as prey to other animals.
The store of nutriment laid up within the seeds of many plants seems at
first sight to have no sort of relation to other plants. But from the
strong growth of young plants produced from such seeds (as peas and
beans), when sown in the midst of long grass, I suspect that the chief
use of the nutriment in the seed is to favour the growth of the young
seedling, whilst struggling with other plants growing vigorously all
around.
Look at a plant in the midst of its range, why does it not double or
quadruple its numbers? We know that it can perfectly well withstand a
little more heat or cold, dampness or dryness, for elsewhere it ranges
into slightly hotter or colder, damper or drier districts. In this case
we can clearly see that if we wished in imagination to give the plant
the power of increasing in number, we should have to give it some
advantage over its competitors, or over the animals which preyed on it.
On the confines of its geographical range, a change of constitution
with respect to climate would clearly be an advantage to our plant; but
we have reason to believe that only a few plants or animals range so
far, that they are destroyed by the rigour of the climate alone. Not
until we reach the extreme confines of life, in the arctic regions or
on the borders of an utter desert, will competition cease. The land may
be extremely cold or dry, yet there will be competition between some
few species, or between the individuals of the same species, for the
warmest or dampest spots.
Hence, also, we can see that when a plant or animal is placed in a new
country amongst new competitors, though the climate may be exactly the
same as in its former home, yet the conditions of its life will
generally be changed in an essential manner. If we wished to increase
its average numbers in its new home, we should have to modify it in a
different way to what we should have done in its native country; for we
should have to give it some advantage over a different set of
competitors or enemies.
It is good thus to try in our imagination to give any form some
advantage over another. Probably in no single instance should we know
what to do, so as to succeed. It will convince us of our ignorance on
the mutual relations of all organic beings; a conviction as necessary,
as it seems to be difficult to acquire. All that we can do, is to keep
steadily in mind that each organic being is striving to increase at a
geometrical
ratio; that each at some period of its life, during some season of the
year, during each generation or at intervals, has to struggle for life,
and to suffer great destruction. When we reflect on this struggle, we
may console ourselves with the full belief, that the war of nature is
not incessant, that no fear is felt, that death is generally prompt,
and that the vigorous, the healthy, and the happy survive and multiply.
CHAPTER IV.
NATURAL SELECTION.
Natural Selection: its power compared with man’s selection, its power
on characters of trifling importance, its power at all ages and on both
sexes. Sexual Selection. On the generality of intercrosses between
individuals of the same species. Circumstances favourable and
unfavourable to Natural Selection, namely, intercrossing, isolation,
number of individuals. Slow action. Extinction caused by Natural
Selection. Divergence of Character, related to the diversity of
inhabitants of any small area, and to naturalisation. Action of Natural
Selection, through Divergence of Character and Extinction, on the
descendants from a common parent. Explains the Grouping of all organic
beings.
How will the struggle for existence, discussed too briefly in the last
chapter, act in regard to variation? Can the principle of selection,
which we have seen is so potent in the hands of man, apply in nature? I
think we shall see that it can act most effectually. Let it be borne in
mind in what an endless number of strange peculiarities our domestic
productions, and, in a lesser degree, those under nature, vary; and how
strong the hereditary tendency is. Under domestication, it may be truly
said that the whole organisation becomes in some degree plastic. Let it
be borne in mind how infinitely complex and close-fitting are the
mutual relations of all organic beings to each other and to their
physical conditions of life. Can it, then, be thought improbable,
seeing that variations useful to man have undoubtedly occurred, that
other variations useful in some way to each being in the great and
complex battle of life, should sometimes occur in the course of
thousands of generations? If such do occur, can we doubt (remembering
that many more individuals are born than can possibly survive) that
individuals having any advantage, however slight, over others, would
have the best chance of surviving and of procreating their kind? On the
other hand, we may feel sure that any variation in the least degree
injurious would be rigidly destroyed. This preservation of favourable
variations and the rejection of injurious variations, I call Natural
Selection. Variations neither useful nor injurious would not be
affected by natural selection, and would be left a fluctuating element,
as perhaps we see in the species called polymorphic.
We shall best understand the probable course of natural selection by
taking the case of a country undergoing some physical change, for
instance, of climate. The proportional numbers of its inhabitants would
almost immediately undergo a change, and some species might become
extinct. We may conclude, from what we have seen of the intimate and
complex manner in which the inhabitants of each country are bound
together, that any change in the numerical proportions of some of the
inhabitants, independently of the change of climate itself, would most
seriously affect many of the others. If the country were open on its
borders, new forms would certainly immigrate, and this also would
seriously disturb the relations of some of the former inhabitants. Let
it be remembered how powerful the influence of a single introduced tree
or mammal has been shown to be. But in the case of an island, or of a
country partly surrounded by barriers, into which new and better
adapted forms could not freely enter, we should then have places in the
economy of nature which would assuredly be better filled up, if some of
the original inhabitants were in some manner modified; for, had the
area been open to immigration, these same
places would have been seized on by intruders. In such case, every
slight modification, which in the course of ages chanced to arise, and
which in any way favoured the individuals of any of the species, by
better adapting them to their altered conditions, would tend to be
preserved; and natural selection would thus have free scope for the
work of improvement.
We have reason to believe, as stated in the first chapter, that a
change in the conditions of life, by specially acting on the
reproductive system, causes or increases variability; and in the
foregoing case the conditions of life are supposed to have undergone a
change, and this would manifestly be favourable to natural selection,
by giving a better chance of profitable variations occurring; and
unless profitable variations do occur, natural selection can do
nothing. Not that, as I believe, any extreme amount of variability is
necessary; as man can certainly produce great results by adding up in
any given direction mere individual differences, so could Nature, but
far more easily, from having incomparably longer time at her disposal.
Nor do I believe that any great physical change, as of climate, or any
unusual degree of isolation to check immigration, is actually necessary
to produce new and unoccupied places for natural selection to fill up
by modifying and improving some of the varying inhabitants. For as all
the inhabitants of each country are struggling together with nicely
balanced forces, extremely slight modifications in the structure or
habits of one inhabitant would often give it an advantage over others;
and still further modifications of the same kind would often still
further increase the advantage. No country can be named in which all
the native inhabitants are now so perfectly adapted to each other and
to the physical conditions under which they live, that none of
them could anyhow be improved; for in all countries, the natives have
been so far conquered by naturalised productions, that they have
allowed foreigners to take firm possession of the land. And as
foreigners have thus everywhere beaten some of the natives, we may
safely conclude that the natives might have been modified with
advantage, so as to have better resisted such intruders.
As man can produce and certainly has produced a great result by his
methodical and unconscious means of selection, what may not nature
effect? Man can act only on external and visible characters: nature
cares nothing for appearances, except in so far as they may be useful
to any being. She can act on every internal organ, on every shade of
constitutional difference, on the whole machinery of life. Man selects
only for his own good; Nature only for that of the being which she
tends. Every selected character is fully exercised by her; and the
being is placed under well-suited conditions of life. Man keeps the
natives of many climates in the same country; he seldom exercises each
selected character in some peculiar and fitting manner; he feeds a long
and a short beaked pigeon on the same food; he does not exercise a
long-backed or long-legged quadruped in any peculiar manner; he exposes
sheep with long and short wool to the same climate. He does not allow
the most vigorous males to struggle for the females. He does not
rigidly destroy all inferior animals, but protects during each varying
season, as far as lies in his power, all his productions. He often
begins his selection by some half-monstrous form; or at least by some
modification prominent enough to catch his eye, or to be plainly useful
to him. Under nature, the slightest difference of structure or
constitution may well turn the nicely-balanced scale in the
struggle for life, and so be preserved. How fleeting are the wishes and
efforts of man! how short his time! and consequently how poor will his
products be, compared with those accumulated by nature during whole
geological periods. Can we wonder, then, that nature’s productions
should be far “truer” in character than man’s productions; that they
should be infinitely better adapted to the most complex conditions of
life, and should plainly bear the stamp of far higher workmanship?
It may be said that natural selection is daily and hourly scrutinising,
throughout the world, every variation, even the slightest; rejecting
that which is bad, preserving and adding up all that is good; silently
and insensibly working, whenever and wherever opportunity offers, at
the improvement of each organic being in relation to its organic and
inorganic conditions of life. We see nothing of these slow changes in
progress, until the hand of time has marked the long lapse of ages, and
then so imperfect is our view into long past geological ages, that we
only see that the forms of life are now different from what they
formerly were.
Although natural selection can act only through and for the good of
each being, yet characters and structures, which we are apt to consider
as of very trifling importance, may thus be acted on. When we see
leaf-eating insects green, and bark-feeders mottled-grey; the alpine
ptarmigan white in winter, the red-grouse the colour of heather, and
the black-grouse that of peaty earth, we must believe that these tints
are of service to these birds and insects in preserving them from
danger. Grouse, if not destroyed at some period of their lives, would
increase in countless numbers; they are known to suffer largely from
birds of prey; and hawks are guided by eyesight to their prey,—so much
so, that on
parts of the Continent persons are warned not to keep white pigeons, as
being the most liable to destruction. Hence I can see no reason to
doubt that natural selection might be most effective in giving the
proper colour to each kind of grouse, and in keeping that colour, when
once acquired, true and constant. Nor ought we to think that the
occasional destruction of an animal of any particular colour would
produce little effect: we should remember how essential it is in a
flock of white sheep to destroy every lamb with the faintest trace of
black. In plants the down on the fruit and the colour of the flesh are
considered by botanists as characters of the most trifling importance:
yet we hear from an excellent horticulturist, Downing, that in the
United States smooth-skinned fruits suffer far more from a beetle, a
curculio, than those with down; that purple plums suffer far more from
a certain disease than yellow plums; whereas another disease attacks
yellow-fleshed peaches far more than those with other coloured flesh.
If, with all the aids of art, these slight differences make a great
difference in cultivating the several varieties, assuredly, in a state
of nature, where the trees would have to struggle with other trees and
with a host of enemies, such differences would effectually settle which
variety, whether a smooth or downy, a yellow or purple fleshed fruit,
should succeed.
In looking at many small points of difference between species, which,
as far as our ignorance permits us to judge, seem to be quite
unimportant, we must not forget that climate, food, etc., probably
produce some slight and direct effect. It is, however, far more
necessary to bear in mind that there are many unknown laws of
correlation of growth, which, when one part of the organisation is
modified through variation, and the modifications are accumulated by
natural selection for
the good of the being, will cause other modifications, often of the
most unexpected nature.
As we see that those variations which under domestication appear at any
particular period of life, tend to reappear in the offspring at the
same period;—for instance, in the seeds of the many varieties of our
culinary and agricultural plants; in the caterpillar and cocoon stages
of the varieties of the silkworm; in the eggs of poultry, and in the
colour of the down of their chickens; in the horns of our sheep and
cattle when nearly adult;—so in a state of nature, natural selection
will be enabled to act on and modify organic beings at any age, by the
accumulation of profitable variations at that age, and by their
inheritance at a corresponding age. If it profit a plant to have its
seeds more and more widely disseminated by the wind, I can see no
greater difficulty in this being effected through natural selection,
than in the cotton-planter increasing and improving by selection the
down in the pods on his cotton-trees. Natural selection may modify and
adapt the larva of an insect to a score of contingencies, wholly
different from those which concern the mature insect. These
modifications will no doubt affect, through the laws of correlation,
the structure of the adult; and probably in the case of those insects
which live only for a few hours, and which never feed, a large part of
their structure is merely the correlated result of successive changes
in the structure of their larvæ. So, conversely, modifications in the
adult will probably often affect the structure of the larva; but in all
cases natural selection will ensure that modifications consequent on
other modifications at a different period of life, shall not be in the
least degree injurious: for if they became so, they would cause the
extinction of the species.
Natural selection will modify the structure of the
young in relation to the parent, and of the parent in relation to the
young. In social animals it will adapt the structure of each individual
for the benefit of the community; if each in consequence profits by the
selected change. What natural selection cannot do, is to modify the
structure of one species, without giving it any advantage, for the good
of another species; and though statements to this effect may be found
in works of natural history, I cannot find one case which will bear
investigation. A structure used only once in an animal’s whole life, if
of high importance to it, might be modified to any extent by natural
selection; for instance, the great jaws possessed by certain insects,
and used exclusively for opening the cocoon—or the hard tip to the beak
of nestling birds, used for breaking the egg. It has been asserted,
that of the best short-beaked tumbler-pigeons more perish in the egg
than are able to get out of it; so that fanciers assist in the act of
hatching. Now, if nature had to make the beak of a full-grown pigeon
very short for the bird’s own advantage, the process of modification
would be very slow, and there would be simultaneously the most rigorous
selection of the young birds within the egg, which had the most
powerful and hardest beaks, for all with weak beaks would inevitably
perish: or, more delicate and more easily broken shells might be
selected, the thickness of the shell being known to vary like every
other structure.
_Sexual Selection_.—Inasmuch as peculiarities often appear under
domestication in one sex and become hereditarily attached to that sex,
the same fact probably occurs under nature, and if so, natural
selection will be able to modify one sex in its functional relations to
the other sex, or in relation to wholly different habits of life in the
two sexes, as is sometimes the case
with insects. And this leads me to say a few words on what I call
Sexual Selection. This depends, not on a struggle for existence, but on
a struggle between the males for possession of the females; the result
is not death to the unsuccessful competitor, but few or no offspring.
Sexual selection is, therefore, less rigorous than natural selection.
Generally, the most vigorous males, those which are best fitted for
their places in nature, will leave most progeny. But in many cases,
victory will depend not on general vigour, but on having special
weapons, confined to the male sex. A hornless stag or spurless cock
would have a poor chance of leaving offspring. Sexual selection by
always allowing the victor to breed might surely give indomitable
courage, length to the spur, and strength to the wing to strike in the
spurred leg, as well as the brutal cock-fighter, who knows well that he
can improve his breed by careful selection of the best cocks. How low
in the scale of nature this law of battle descends, I know not; male
alligators have been described as fighting, bellowing, and whirling
round, like Indians in a war-dance, for the possession of the females;
male salmons have been seen fighting all day long; male stag-beetles
often bear wounds from the huge mandibles of other males. The war is,
perhaps, severest between the males of polygamous animals, and these
seem oftenest provided with special weapons. The males of carnivorous
animals are already well armed; though to them and to others, special
means of defence may be given through means of sexual selection, as the
mane to the lion, the shoulder-pad to the boar, and the hooked jaw to
the male salmon; for the shield may be as important for victory, as the
sword or spear.
Amongst birds, the contest is often of a more peaceful character. All
those who have attended to the subject,
believe that there is the severest rivalry between the males of many
species to attract by singing the females. The rock-thrush of Guiana,
birds of Paradise, and some others, congregate; and successive males
display their gorgeous plumage and perform strange antics before the
females, which standing by as spectators, at last choose the most
attractive partner. Those who have closely attended to birds in
confinement well know that they often take individual preferences and
dislikes: thus Sir R. Heron has described how one pied peacock was
eminently attractive to all his hen birds. It may appear childish to
attribute any effect to such apparently weak means: I cannot here enter
on the details necessary to support this view; but if man can in a
short time give elegant carriage and beauty to his bantams, according
to his standard of beauty, I can see no good reason to doubt that
female birds, by selecting, during thousands of generations, the most
melodious or beautiful males, according to their standard of beauty,
might produce a marked effect. I strongly suspect that some well-known
laws with respect to the plumage of male and female birds, in
comparison with the plumage of the young, can be explained on the view
of plumage having been chiefly modified by sexual selection, acting
when the birds have come to the breeding age or during the breeding
season; the modifications thus produced being inherited at
corresponding ages or seasons, either by the males alone, or by the
males and females; but I have not space here to enter on this subject.
Thus it is, as I believe, that when the males and females of any animal
have the same general habits of life, but differ in structure, colour,
or ornament, such differences have been mainly caused by sexual
selection; that is, individual males have had, in successive
generations, some slight advantage over other
males, in their weapons, means of defence, or charms; and have
transmitted these advantages to their male offspring. Yet, I would not
wish to attribute all such sexual differences to this agency: for we
see peculiarities arising and becoming attached to the male sex in our
domestic animals (as the wattle in male carriers, horn-like
protuberances in the cocks of certain fowls, etc.), which we cannot
believe to be either useful to the males in battle, or attractive to
the females. We see analogous cases under nature, for instance, the
tuft of hair on the breast of the turkey-cock, which can hardly be
either useful or ornamental to this bird;—indeed, had the tuft appeared
under domestication, it would have been called a monstrosity.
_Illustrations of the action of Natural Selection_.—In order to make it
clear how, as I believe, natural selection acts, I must beg permission
to give one or two imaginary illustrations. Let us take the case of a
wolf, which preys on various animals, securing some by craft, some by
strength, and some by fleetness; and let us suppose that the fleetest
prey, a deer for instance, had from any change in the country increased
in numbers, or that other prey had decreased in numbers, during that
season of the year when the wolf is hardest pressed for food. I can
under such circumstances see no reason to doubt that the swiftest and
slimmest wolves would have the best chance of surviving, and so be
preserved or selected,—provided always that they retained strength to
master their prey at this or at some other period of the year, when
they might be compelled to prey on other animals. I can see no more
reason to doubt this, than that man can improve the fleetness of his
greyhounds by careful and methodical selection, or by that unconscious
selection which results from each man trying
to keep the best dogs without any thought of modifying the breed.
Even without any change in the proportional numbers of the animals on
which our wolf preyed, a cub might be born with an innate tendency to
pursue certain kinds of prey. Nor can this be thought very improbable;
for we often observe great differences in the natural tendencies of our
domestic animals; one cat, for instance, taking to catch rats, another
mice; one cat, according to Mr. St. John, bringing home winged game,
another hares or rabbits, and another hunting on marshy ground and
almost nightly catching woodcocks or snipes. The tendency to catch rats
rather than mice is known to be inherited. Now, if any slight innate
change of habit or of structure benefited an individual wolf, it would
have the best chance of surviving and of leaving offspring. Some of its
young would probably inherit the same habits or structure, and by the
repetition of this process, a new variety might be formed which would
either supplant or coexist with the parent-form of wolf. Or, again, the
wolves inhabiting a mountainous district, and those frequenting the
lowlands, would naturally be forced to hunt different prey; and from
the continued preservation of the individuals best fitted for the two
sites, two varieties might slowly be formed. These varieties would
cross and blend where they met; but to this subject of intercrossing we
shall soon have to return. I may add, that, according to Mr. Pierce,
there are two varieties of the wolf inhabiting the Catskill Mountains
in the United States, one with a light greyhound-like form, which
pursues deer, and the other more bulky, with shorter legs, which more
frequently attacks the shepherd’s flocks.
Let us now take a more complex case. Certain plants excrete a sweet
juice, apparently for the sake of eliminating something injurious from
their sap: this is
effected by glands at the base of the stipules in some Leguminosæ, and
at the back of the leaf of the common laurel. This juice, though small
in quantity, is greedily sought by insects. Let us now suppose a little
sweet juice or nectar to be excreted by the inner bases of the petals
of a flower. In this case insects in seeking the nectar would get
dusted with pollen, and would certainly often transport the pollen from
one flower to the stigma of another flower. The flowers of two distinct
individuals of the same species would thus get crossed; and the act of
crossing, we have good reason to believe (as will hereafter be more
fully alluded to), would produce very vigorous seedlings, which
consequently would have the best chance of flourishing and surviving.
Some of these seedlings would probably inherit the nectar-excreting
power. Those individual flowers which had the largest glands or
nectaries, and which excreted most nectar, would be oftenest visited by
insects, and would be oftenest crossed; and so in the long-run would
gain the upper hand. Those flowers, also, which had their stamens and
pistils placed, in relation to the size and habits of the particular
insects which visited them, so as to favour in any degree the
transportal of their pollen from flower to flower, would likewise be
favoured or selected. We might have taken the case of insects visiting
flowers for the sake of collecting pollen instead of nectar; and as
pollen is formed for the sole object of fertilisation, its destruction
appears a simple loss to the plant; yet if a little pollen were
carried, at first occasionally and then habitually, by the
pollen-devouring insects from flower to flower, and a cross thus
effected, although nine-tenths of the pollen were destroyed, it might
still be a great gain to the plant; and those individuals which
produced more and more pollen, and had larger and larger anthers, would
be selected.
When our plant, by this process of the continued preservation or
natural selection of more and more attractive flowers, had been
rendered highly attractive to insects, they would, unintentionally on
their part, regularly carry pollen from flower to flower; and that they
can most effectually do this, I could easily show by many striking
instances. I will give only one—not as a very striking case, but as
likewise illustrating one step in the separation of the sexes of
plants, presently to be alluded to. Some holly-trees bear only male
flowers, which have four stamens producing rather a small quantity of
pollen, and a rudimentary pistil; other holly-trees bear only female
flowers; these have a full-sized pistil, and four stamens with
shrivelled anthers, in which not a grain of pollen can be detected.
Having found a female tree exactly sixty yards from a male tree, I put
the stigmas of twenty flowers, taken from different branches, under the
microscope, and on all, without exception, there were pollen-grains,
and on some a profusion of pollen. As the wind had set for several days
from the female to the male tree, the pollen could not thus have been
carried. The weather had been cold and boisterous, and therefore not
favourable to bees, nevertheless every female flower which I examined
had been effectually fertilised by the bees, accidentally dusted with
pollen, having flown from tree to tree in search of nectar. But to
return to our imaginary case: as soon as the plant had been rendered so
highly attractive to insects that pollen was regularly carried from
flower to flower, another process might commence. No naturalist doubts
the advantage of what has been called the “physiological division of
labour;” hence we may believe that it would be advantageous to a plant
to produce stamens alone in one flower or on one whole plant, and
pistils alone in
another flower or on another plant. In plants under culture and placed
under new conditions of life, sometimes the male organs and sometimes
the female organs become more or less impotent; now if we suppose this
to occur in ever so slight a degree under nature, then as pollen is
already carried regularly from flower to flower, and as a more complete
separation of the sexes of our plant would be advantageous on the
principle of the division of labour, individuals with this tendency
more and more increased, would be continually favoured or selected,
until at last a complete separation of the sexes would be effected.
Let us now turn to the nectar-feeding insects in our imaginary case: we
may suppose the plant of which we have been slowly increasing the
nectar by continued selection, to be a common plant; and that certain
insects depended in main part on its nectar for food. I could give many
facts, showing how anxious bees are to save time; for instance, their
habit of cutting holes and sucking the nectar at the bases of certain
flowers, which they can, with a very little more trouble, enter by the
mouth. Bearing such facts in mind, I can see no reason to doubt that an
accidental deviation in the size and form of the body, or in the
curvature and length of the proboscis, etc., far too slight to be
appreciated by us, might profit a bee or other insect, so that an
individual so characterised would be able to obtain its food more
quickly, and so have a better chance of living and leaving descendants.
Its descendants would probably inherit a tendency to a similar slight
deviation of structure. The tubes of the corollas of the common red and
incarnate clovers (Trifolium pratense and incarnatum) do not on a hasty
glance appear to differ in length; yet the hive-bee can easily suck the
nectar out of the incarnate clover, but not out of the common red
clover, which is visited by humble-bees alone; so that whole fields of
the red clover offer in vain an abundant supply of precious nectar to
the hive-bee. Thus it might be a great advantage to the hive-bee to
have a slightly longer or differently constructed proboscis. On the
other hand, I have found by experiment that the fertility of clover
greatly depends on bees visiting and moving parts of the corolla, so as
to push the pollen on to the stigmatic surface. Hence, again, if
humble-bees were to become rare in any country, it might be a great
advantage to the red clover to have a shorter or more deeply divided
tube to its corolla, so that the hive-bee could visit its flowers. Thus
I can understand how a flower and a bee might slowly become, either
simultaneously or one after the other, modified and adapted in the most
perfect manner to each other, by the continued preservation of
individuals presenting mutual and slightly favourable deviations of
structure.
I am well aware that this doctrine of natural selection, exemplified in
the above imaginary instances, is open to the same objections which
were at first urged against Sir Charles Lyell’s noble views on “the
modern changes of the earth, as illustrative of geology;” but we now
very seldom hear the action, for instance, of the coast-waves, called a
trifling and insignificant cause, when applied to the excavation of
gigantic valleys or to the formation of the longest lines of inland
cliffs. Natural selection can act only by the preservation and
accumulation of infinitesimally small inherited modifications, each
profitable to the preserved being; and as modern geology has almost
banished such views as the excavation of a great valley by a single
diluvial wave, so will natural selection, if it be a true principle,
banish the belief of the continued creation of new organic
beings, or of any great and sudden modification in their structure.
_On the Intercrossing of Individuals_.—I must here introduce a short
digression. In the case of animals and plants with separated sexes, it
is of course obvious that two individuals must always unite for each
birth; but in the case of hermaphrodites this is far from obvious.
Nevertheless I am strongly inclined to believe that with all
hermaphrodites two individuals, either occasionally or habitually,
concur for the reproduction of their kind. This view, I may add, was
first suggested by Andrew Knight. We shall presently see its
importance; but I must here treat the subject with extreme brevity,
though I have the materials prepared for an ample discussion. All
vertebrate animals, all insects, and some other large groups of
animals, pair for each birth. Modern research has much diminished the
number of supposed hermaphrodites, and of real hermaphrodites a large
number pair; that is, two individuals regularly unite for reproduction,
which is all that concerns us. But still there are many hermaphrodite
animals which certainly do not habitually pair, and a vast majority of
plants are hermaphrodites. What reason, it may be asked, is there for
supposing in these cases that two individuals ever concur in
reproduction? As it is impossible here to enter on details, I must
trust to some general considerations alone.
In the first place, I have collected so large a body of facts, showing,
in accordance with the almost universal belief of breeders, that with
animals and plants a cross between different varieties, or between
individuals of the same variety but of another strain, gives vigour and
fertility to the offspring; and on the other hand, that _close_
interbreeding diminishes vigour and fertility; that
these facts alone incline me to believe that it is a general law of
nature (utterly ignorant though we be of the meaning of the law) that
no organic being self-fertilises itself for an eternity of generations;
but that a cross with another individual is occasionally—perhaps at
very long intervals—indispensable.
On the belief that this is a law of nature, we can, I think, understand
several large classes of facts, such as the following, which on any
other view are inexplicable. Every hybridizer knows how unfavourable
exposure to wet is to the fertilisation of a flower, yet what a
multitude of flowers have their anthers and stigmas fully exposed to
the weather! but if an occasional cross be indispensable, the fullest
freedom for the entrance of pollen from another individual will explain
this state of exposure, more especially as the plant’s own anthers and
pistil generally stand so close together that self-fertilisation seems
almost inevitable. Many flowers, on the other hand, have their organs
of fructification closely enclosed, as in the great papilionaceous or
pea-family; but in several, perhaps in all, such flowers, there is a
very curious adaptation between the structure of the flower and the
manner in which bees suck the nectar; for, in doing this, they either
push the flower’s own pollen on the stigma, or bring pollen from
another flower. So necessary are the visits of bees to papilionaceous
flowers, that I have found, by experiments published elsewhere, that
their fertility is greatly diminished if these visits be prevented.
Now, it is scarcely possible that bees should fly from flower to
flower, and not carry pollen from one to the other, to the great good,
as I believe, of the plant. Bees will act like a camel-hair pencil, and
it is quite sufficient just to touch the anthers of one flower and then
the stigma of another with the same brush to ensure fertilisation; but
it must not be
supposed that bees would thus produce a multitude of hybrids between
distinct species; for if you bring on the same brush a plant’s own
pollen and pollen from another species, the former will have such a
prepotent effect, that it will invariably and completely destroy, as
has been shown by Gärtner, any influence from the foreign pollen.
When the stamens of a flower suddenly spring towards the pistil, or
slowly move one after the other towards it, the contrivance seems
adapted solely to ensure self-fertilisation; and no doubt it is useful
for this end: but, the agency of insects is often required to cause the
stamens to spring forward, as Kölreuter has shown to be the case with
the barberry; and curiously in this very genus, which seems to have a
special contrivance for self-fertilisation, it is well known that if
very closely-allied forms or varieties are planted near each other, it
is hardly possible to raise pure seedlings, so largely do they
naturally cross. In many other cases, far from there being any aids for
self-fertilisation, there are special contrivances, as I could show
from the writings of C. C. Sprengel and from my own observations, which
effectually prevent the stigma receiving pollen from its own flower:
for instance, in Lobelia fulgens, there is a really beautiful and
elaborate contrivance by which every one of the infinitely numerous
pollen-granules are swept out of the conjoined anthers of each flower,
before the stigma of that individual flower is ready to receive them;
and as this flower is never visited, at least in my garden, by insects,
it never sets a seed, though by placing pollen from one flower on the
stigma of another, I raised plenty of seedlings; and whilst another
species of Lobelia growing close by, which is visited by bees, seeds
freely. In very many other cases, though there be no special mechanical
contrivance to prevent the stigma of a flower receiving its own pollen,
yet, as
C. C. Sprengel has shown, and as I can confirm, either the anthers
burst before the stigma is ready for fertilisation, or the stigma is
ready before the pollen of that flower is ready, so that these plants
have in fact separated sexes, and must habitually be crossed. How
strange are these facts! How strange that the pollen and stigmatic
surface of the same flower, though placed so close together, as if for
the very purpose of self-fertilisation, should in so many cases be
mutually useless to each other! How simply are these facts explained on
the view of an occasional cross with a distinct individual being
advantageous or indispensable!
If several varieties of the cabbage, radish, onion, and of some other
plants, be allowed to seed near each other, a large majority, as I have
found, of the seedlings thus raised will turn out mongrels: for
instance, I raised 233 seedling cabbages from some plants of different
varieties growing near each other, and of these only 78 were true to
their kind, and some even of these were not perfectly true. Yet the
pistil of each cabbage-flower is surrounded not only by its own six
stamens, but by those of the many other flowers on the same plant. How,
then, comes it that such a vast number of the seedlings are
mongrelized? I suspect that it must arise from the pollen of a distinct
_variety_ having a prepotent effect over a flower’s own pollen; and
that this is part of the general law of good being derived from the
intercrossing of distinct individuals of the same species. When
distinct _species_ are crossed the case is directly the reverse, for a
plant’s own pollen is always prepotent over foreign pollen; but to this
subject we shall return in a future chapter.
In the case of a gigantic tree covered with innumerable flowers, it may
be objected that pollen could seldom be carried from tree to tree, and
at most only from flower
to flower on the same tree, and that flowers on the same tree can be
considered as distinct individuals only in a limited sense. I believe
this objection to be valid, but that nature has largely provided
against it by giving to trees a strong tendency to bear flowers with
separated sexes. When the sexes are separated, although the male and
female flowers may be produced on the same tree, we can see that pollen
must be regularly carried from flower to flower; and this will give a
better chance of pollen being occasionally carried from tree to tree.
That trees belonging to all Orders have their sexes more often
separated than other plants, I find to be the case in this country; and
at my request Dr. Hooker tabulated the trees of New Zealand, and Dr.
Asa Gray those of the United States, and the result was as I
anticipated. On the other hand, Dr. Hooker has recently informed me
that he finds that the rule does not hold in Australia; and I have made
these few remarks on the sexes of trees simply to call attention to the
subject.
Turning for a very brief space to animals: on the land there are some
hermaphrodites, as land-mollusca and earth-worms; but these all pair.
As yet I have not found a single case of a terrestrial animal which
fertilises itself. We can understand this remarkable fact, which offers
so strong a contrast with terrestrial plants, on the view of an
occasional cross being indispensable, by considering the medium in
which terrestrial animals live, and the nature of the fertilising
element; for we know of no means, analogous to the action of insects
and of the wind in the case of plants, by which an occasional cross
could be effected with terrestrial animals without the concurrence of
two individuals. Of aquatic animals, there are many self-fertilising
hermaphrodites; but here currents in the water offer an obvious means
for an occasional cross. And, as in the case of flowers, I have as yet
failed, after consultation with one of the highest authorities, namely,
Professor Huxley, to discover a single case of an hermaphrodite animal
with the organs of reproduction so perfectly enclosed within the body,
that access from without and the occasional influence of a distinct
individual can be shown to be physically impossible. Cirripedes long
appeared to me to present a case of very great difficulty under this
point of view; but I have been enabled, by a fortunate chance,
elsewhere to prove that two individuals, though both are
self-fertilising hermaphrodites, do sometimes cross.
It must have struck most naturalists as a strange anomaly that, in the
case of both animals and plants, species of the same family and even of
the same genus, though agreeing closely with each other in almost their
whole organisation, yet are not rarely, some of them hermaphrodites,
and some of them unisexual. But if, in fact, all hermaphrodites do
occasionally intercross with other individuals, the difference between
hermaphrodites and unisexual species, as far as function is concerned,
becomes very small.
From these several considerations and from the many special facts which
I have collected, but which I am not here able to give, I am strongly
inclined to suspect that, both in the vegetable and animal kingdoms, an
occasional intercross with a distinct individual is a law of nature. I
am well aware that there are, on this view, many cases of difficulty,
some of which I am trying to investigate. Finally then, we may conclude
that in many organic beings, a cross between two individuals is an
obvious necessity for each birth; in many others it occurs perhaps only
at long intervals; but in none, as I suspect, can self-fertilisation go
on for perpetuity.
_Circumstances favourable to Natural Selection_.—This
is an extremely intricate subject. A large amount of inheritable and
diversified variability is favourable, but I believe mere individual
differences suffice for the work. A large number of individuals, by
giving a better chance for the appearance within any given period of
profitable variations, will compensate for a lesser amount of
variability in each individual, and is, I believe, an extremely
important element of success. Though nature grants vast periods of time
for the work of natural selection, she does not grant an indefinite
period; for as all organic beings are striving, it may be said, to
seize on each place in the economy of nature, if any one species does
not become modified and improved in a corresponding degree with its
competitors, it will soon be exterminated.
In man’s methodical selection, a breeder selects for some definite
object, and free intercrossing will wholly stop his work. But when many
men, without intending to alter the breed, have a nearly common
standard of perfection, and all try to get and breed from the best
animals, much improvement and modification surely but slowly follow
from this unconscious process of selection, notwithstanding a large
amount of crossing with inferior animals. Thus it will be in nature;
for within a confined area, with some place in its polity not so
perfectly occupied as might be, natural selection will always tend to
preserve all the individuals varying in the right direction, though in
different degrees, so as better to fill up the unoccupied place. But if
the area be large, its several districts will almost certainly present
different conditions of life; and then if natural selection be
modifying and improving a species in the several districts, there will
be intercrossing with the other individuals of the same species on the
confines of each. And in this case the effects of intercrossing can
hardly be counterbalanced
by natural selection always tending to modify all the individuals in
each district in exactly the same manner to the conditions of each; for
in a continuous area, the conditions will generally graduate away
insensibly from one district to another. The intercrossing will most
affect those animals which unite for each birth, which wander much, and
which do not breed at a very quick rate. Hence in animals of this
nature, for instance in birds, varieties will generally be confined to
separated countries; and this I believe to be the case. In
hermaphrodite organisms which cross only occasionally, and likewise in
animals which unite for each birth, but which wander little and which
can increase at a very rapid rate, a new and improved variety might be
quickly formed on any one spot, and might there maintain itself in a
body, so that whatever intercrossing took place would be chiefly
between the individuals of the same new variety. A local variety when
once thus formed might subsequently slowly spread to other districts.
On the above principle, nurserymen always prefer getting seed from a
large body of plants of the same variety, as the chance of
intercrossing with other varieties is thus lessened.
Even in the case of slow-breeding animals, which unite for each birth,
we must not overrate the effects of intercrosses in retarding natural
selection; for I can bring a considerable catalogue of facts, showing
t
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