Sunday, October 27, 2013

On the Origine Of Species


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.
I have hitherto sometimes spoken as if the variations--so common and multiform in organic beings under
domestication, and in a lesser degree in those in a state of nature--had been due to chance. This, of course, is a
wholly incorrect expression, but it serves to acknowledge plainly our ignorance of the cause of each particular
variation. Some authors believe it to be as much the function of the reproductive system to produce individual
differences, or very slight deviations of structure, as to make the child like its parents. But the much greater
variability, as well as the greater frequency of monstrosities, under domestication or cultivation, than under
nature, leads me to believe that deviations of structure are in some way due to the nature of the conditions of
life, to which the parents and their more remote ancestors have been exposed during several generations. I
have remarked in the first chapter--but a long catalogue of facts which cannot be here given would be
necessary to show the truth of the remark--that the reproductive system is eminently susceptible to changes in
the conditions of life; and to this system being functionally disturbed in the parents, I chiefly attribute the
varying or plastic condition of the offspring. The male and female sexual elements seem to be affected before
that union takes place which is to form a new being. In the case of "sporting" plants, the bud, which in its
earliest condition does not apparently differ essentially from an ovule, is alone affected. But why, because the
reproductive system is disturbed, this or that part should vary more or less, we are profoundly ignorant.
Nevertheless, we can here and there dimly catch a faint ray of light, and we may feel sure that there must be
some cause for each deviation of structure, however slight.
How much direct effect difference of climate, food, etc., produces on any being is extremely doubtful. My
impression is, that the effect is extremely small in the case of animals, but perhaps rather more in that of
plants. We may, at least, safely conclude that such influences cannot have produced the many striking and
complex co-adaptations of structure between one organic being and another, which we see everywhere
throughout nature. Some little influence may be attributed to climate, food, etc.: thus, E. Forbes speaks
confidently that shells at their southern limit, and when living in shallow water, are more brightly coloured
than those of the same species further north or from greater depths. Gould believes that birds of the same
species are more brightly coloured under a clear atmosphere, than when living on islands or near the coast. So
with insects, Wollaston is convinced that residence near the sea affects their colours. Moquin-Tandon gives a
list of plants which when growing near the sea-shore have their leaves in some degree fleshy, though not
elsewhere fleshy. Several other such cases could be given.
The fact of varieties of one species, when they range into the zone of habitation of other species, often
acquiring in a very slight degree some of the characters of such species, accords with our view that species of
all kinds are only well-marked and permanent varieties. Thus the species of shells which are confined to
tropical and shallow seas are generally brighter-coloured than those confined to cold and deeper seas. The
birds which are confined to continents are, according to Mr. Gould, brighter-coloured than those of islands.
The insect-species confined to sea-coasts, as every collector knows, are often brassy or lurid. Plants which
live exclusively on the sea-side are very apt to have fleshy leaves. He who believes in the creation of each
species, will have to say that this shell, for instance, was created with bright colours for a warm sea; but that
this other shell became bright-coloured by variation when it ranged into warmer or shallower waters.
When a variation is of the slightest use to a being, we cannot tell how much of it to attribute to the
accumulative action of natural selection, and how much to the conditions of life. Thus, it is well known to
furriers that animals of the same species have thicker and better fur the more severe the climate is under which
they have lived; but who can tell how much of this difference may be due to the warmest-clad individuals
having been favoured and preserved during many generations, and how much to the direct action of the severe
climate? for it would appear that climate has some direct action on the hair of our domestic quadrupeds.
Instances could be given of the same variety being produced under conditions of life as different as can well
be conceived; and, on the other hand, of different varieties being produced from the same species under the
same conditions. Such facts show how indirectly the conditions of life must act. Again, innumerable instances
are known to every naturalist of species keeping true, or not varying at all, although living under the most
opposite climates. Such considerations as these incline me to lay very little weight on the direct action of the
conditions of life. Indirectly, as already remarked, they seem to play an important part in affecting the
reproductive system, and in thus inducing variability; and natural selection will then accumulate all profitable
variations, however slight, until they become plainly developed and appreciable by us.
EFFECTS OF USE AND DISUSE.
From the facts alluded to in the first chapter, I think there can be little doubt that use in our domestic animals
strengthens and enlarges certain parts, and disuse diminishes them; and that such modifications are inherited.
Under free nature, we can have no standard of comparison, by which to judge of the effects of long-continued
use or disuse, for we know not the parent-forms; but many animals have structures which can be explained by
the effects of disuse. As Professor Owen has remarked, there is no greater anomaly in nature than a bird that
cannot fly; yet there are several in this state. The logger-headed duck of South America can only flap along
the surface of the water, and has its wings in nearly the same condition as the domestic Aylesbury duck. As
the larger ground-feeding birds seldom take flight except to escape danger, I believe that the nearly wingless
condition of several birds, which now inhabit or have lately inhabited several oceanic islands, tenanted by no
beast of prey, has been caused by disuse. The ostrich indeed inhabits continents and is exposed to danger from
which it cannot escape by flight, but by kicking it can defend itself from enemies, as well as any of the smaller
quadrupeds. We may imagine that the early progenitor of the ostrich had habits like those of a bustard, and
that as natural selection increased in successive generations the size and weight of its body, its legs were used
more, and its wings less, until they became incapable of flight.
Kirby has remarked (and I have observed the same fact) that the anterior tarsi, or feet, of many male
dung-feeding beetles are very often broken off; he examined seventeen specimens in his own collection, and
not one had even a relic left. In the Onites apelles the tarsi are so habitually lost, that the insect has been
described as not having them. In some other genera they are present, but in a rudimentary condition. In the
Ateuchus or sacred beetle of the Egyptians, they are totally deficient. There is not sufficient evidence to
induce us to believe that mutilations are ever inherited; and I should prefer explaining the entire absence of the
anterior tarsi in Ateuchus, and their rudimentary condition in some other genera, by the long-continued effects
of disuse in their progenitors; for as the tarsi are almost always lost in many dung-feeding beetles, they must
be lost early in life, and therefore cannot be much used by these insects.
In some cases we might easily put down to disuse modifications of structure which are wholly, or mainly, due
to natural selection. Mr. Wollaston has discovered the remarkable fact that 200 beetles, out of the 550 species
inhabiting Madeira, are so far deficient in wings that they cannot fly; and that of the twenty-nine endemic
genera, no less than twenty-three genera have all their species in this condition! Several facts, namely, that
beetles in many parts of the world are very frequently blown to sea and perish; that the beetles in Madeira, as
observed by Mr. Wollaston, lie much concealed, until the wind lulls and the sun shines; that the proportion of
wingless beetles is larger on the exposed Dezertas than in Madeira itself; and especially the extraordinary fact,
so strongly insisted on by Mr. Wollaston, of the almost entire absence of certain large groups of beetles,
elsewhere excessively numerous, and which groups have habits of life almost necessitating frequent
flight;--these several considerations have made me believe that the wingless condition of so many Madeira
beetles is mainly due to the action of natural selection, but combined probably with disuse. For during
thousands of successive generations each individual beetle which flew least, either from its wings having been
ever so little less perfectly developed or from indolent habit, will have had the best chance of surviving from
not being blown out to sea; and, on the other hand, those beetles which most readily took to flight will
oftenest have been blown to sea and thus have been destroyed.
The insects in Madeira which are not ground-feeders, and which, as the flower-feeding coleoptera and
lepidoptera, must habitually use their wings to gain their subsistence, have, as Mr. Wollaston suspects, their
wings not at all reduced, but even enlarged. This is quite compatible with the action of natural selection. For
when a new insect first arrived on the island, the tendency of natural selection to enlarge or to reduce the
wings, would depend on whether a greater number of individuals were saved by successfully battling with the
winds, or by giving up the attempt and rarely or never flying. As with mariners shipwrecked near a coast, it
would have been better for the good swimmers if they had been able to swim still further, whereas it would
have been better for the bad swimmers if they had not been able to swim at all and had stuck to the wreck.
The eyes of moles and of some burrowing rodents are rudimentary in size, and in some cases are quite
covered up by skin and fur. This state of the eyes is probably due to gradual reduction from disuse, but aided
perhaps by natural selection. In South America, a burrowing rodent, the tuco-tuco, or Ctenomys, is even more
subterranean in its habits than the mole; and I was assured by a Spaniard, who had often caught them, that
they were frequently blind; one which I kept alive was certainly in this condition, the cause, as appeared on
dissection, having been inflammation of the nictitating membrane. As frequent inflammation of the eyes must
be injurious to any animal, and as eyes are certainly not indispensable to animals with subterranean habits, a
reduction in their size with the adhesion of the eyelids and growth of fur over them, might in such case be an
advantage; and if so, natural selection would constantly aid the effects of disuse.
It is well known that several animals, belonging to the most different classes, which inhabit the caves of Styria
and of Kentucky, are blind. In some of the crabs the foot-stalk for the eye remains, though the eye is gone; the
stand for the telescope is there, though the telescope with its glasses has been lost. As it is difficult to imagine
that eyes, though useless, could be in any way injurious to animals living in darkness, I attribute their loss
wholly to disuse. In one of the blind animals, namely, the cave-rat, the eyes are of immense size; and
Professor Silliman thought that it regained, after living some days in the light, some slight power of vision. In
the same manner as in Madeira the wings of some of the insects have been enlarged, and the wings of others
have been reduced by natural selection aided by use and disuse, so in the case of the cave-rat natural selection
seems to have struggled with the loss of light and to have increased the size of the eyes; whereas with all the
other inhabitants of the caves, disuse by itself seems to have done its work.
It is difficult to imagine conditions of life more similar than deep limestone caverns under a nearly similar
climate; so that on the common view of the blind animals having been separately created for the American
and European caverns, close similarity in their organisation and affinities might have been expected; but, as
Schiodte and others have remarked, this is not the case, and the cave-insects of the two continents are not
more closely allied than might have been anticipated from the general resemblance of the other inhabitants of
North America and Europe. On my view we must suppose that American animals, having ordinary powers of
vision, slowly migrated by successive generations from the outer world into the deeper and deeper recesses of
the Kentucky caves, as did European animals into the caves of Europe. We have some evidence of this
gradation of habit; for, as Schiodte remarks, "animals not far remote from ordinary forms, prepare the
transition from light to darkness. Next follow those that are constructed for twilight; and, last of all, those
destined for total darkness." By the time that an animal had reached, after numberless generations, the deepest
recesses, disuse will on this view have more or less perfectly obliterated its eyes, and natural selection will
often have effected other changes, such as an increase in the length of the antennae or palpi, as a
compensation for blindness. Notwithstanding such modifications, we might expect still to see in the
cave-animals of America, affinities to the other inhabitants of that continent, and in those of Europe, to the
inhabitants of the European continent. And this is the case with some of the American cave-animals, as I hear
from Professor Dana; and some of the European cave-insects are very closely allied to those of the
surrounding country. It would be most difficult to give any rational explanation of the affinities of the blind
cave-animals to the other inhabitants of the two continents on the ordinary view of their independent creation.
That several of the inhabitants of the caves of the Old and New Worlds should be closely related, we might
expect from the well-known relationship of most of their other productions. Far from feeling any surprise that
some of the cave-animals should be very anomalous, as Agassiz has remarked in regard to the blind fish, the
Amblyopsis, and as is the case with the blind Proteus with reference to the reptiles of Europe, I am only
surprised that more wrecks of ancient life have not been preserved, owing to the less severe competition to
which the inhabitants of these dark abodes will probably have been exposed.
ACCLIMATISATION.
Habit is hereditary with plants, as in the period of flowering, in the amount of rain requisite for seeds to
germinate, in the time of sleep, etc., and this leads me to say a few words on acclimatisation. As it is
extremely common for species of the same genus to inhabit very hot and very cold countries, and as I believe
that all the species of the same genus have descended from a single parent, if this view be correct,
acclimatisation must be readily effected during long-continued descent. It is notorious that each species is
adapted to the climate of its own home: species from an arctic or even from a temperate region cannot endure
a tropical climate, or conversely. So again, many succulent plants cannot endure a damp climate. But the
degree of adaptation of species to the climates under which they live is often overrated. We may infer this
from our frequent inability to predict whether or not an imported plant will endure our climate, and from the
number of plants and animals brought from warmer countries which here enjoy good health. We have reason
to believe that species in a state of nature are limited in their ranges by the competition of other organic beings
quite as much as, or more than, by adaptation to particular climates. But whether or not the adaptation be
generally very close, we have evidence, in the case of some few plants, of their becoming, to a certain extent,
naturally habituated to different temperatures, or becoming acclimatised: thus the pines and rhododendrons,
raised from seed collected by Dr. Hooker from trees growing at different heights on the Himalaya, were found
in this country to possess different constitutional powers of resisting cold. Mr. Thwaites informs me that he
has observed similar facts in Ceylon, and analogous observations have been made by Mr. H. C. Watson on
European species of plants brought from the Azores to England. In regard to animals, several authentic cases
could be given of species within historical times having largely extended their range from warmer to cooler
latitudes, and conversely; but we do not positively know that these animals were strictly adapted to their
native climate, but in all ordinary cases we assume such to be the case; nor do we know that they have
subsequently become acclimatised to their new homes.
As I believe that our domestic animals were originally chosen by uncivilised man because they were useful
and bred readily under confinement, and not because they were subsequently found capable of far-extended
transportation, I think the common and extraordinary capacity in our domestic animals of not only
withstanding the most different climates but of being perfectly fertile (a far severer test) under them, may be
used as an argument that a large proportion of other animals, now in a state of nature, could easily be brought
to bear widely different climates. We must not, however, push the foregoing argument too far, on account of
the probable origin of some of our domestic animals from several wild stocks: the blood, for instance, of a
tropical and arctic wolf or wild dog may perhaps be mingled in our domestic breeds. The rat and mouse
cannot be considered as domestic animals, but they have been transported by man to many parts of the world,
and now have a far wider range than any other rodent, living free under the cold climate of Faroe in the north
and of the Falklands in the south, and on many islands in the torrid zones. Hence I am inclined to look at
adaptation to any special climate as a quality readily grafted on an innate wide flexibility of constitution,
which is common to most animals. On this view, the capacity of enduring the most different climates by man
himself and by his domestic animals, and such facts as that former species of the elephant and rhinoceros were
capable of enduring a glacial climate, whereas the living species are now all tropical or sub-tropical in their
habits, ought not to be looked at as anomalies, but merely as examples of a very common flexibility of
constitution, brought, under peculiar circumstances, into play.
How much of the acclimatisation of species to any peculiar climate is due to mere habit, and how much to the
natural selection of varieties having different innate constitutions, and how much to both means combined, is
a very obscure question. That habit or custom has some influence I must believe, both from analogy, and from
the incessant advice given in agricultural works, even in the ancient Encyclopaedias of China, to be very
cautious in transposing animals from one district to another; for it is not likely that man should have
succeeded in selecting so many breeds and sub-breeds with constitutions specially fitted for their own
districts: the result must, I think, be due to habit. On the other hand, I can see no reason to doubt that natural
selection will continually tend to preserve those individuals which are born with constitutions best adapted to
their native countries. In treatises on many kinds of cultivated plants, certain varieties are said to withstand
certain climates better than others: this is very strikingly shown in works on fruit trees published in the United
States, in which certain varieties are habitually recommended for the northern, and others for the southern
States; and as most of these varieties are of recent origin, they cannot owe their constitutional differences to
habit. The case of the Jerusalem artichoke, which is never propagated by seed, and of which consequently new
varieties have not been produced, has even been advanced--for it is now as tender as ever it was--as proving
that acclimatisation cannot be effected! The case, also, of the kidney-bean has been often cited for a similar
purpose, and with much greater weight; but until some one will sow, during a score of generations, his
kidney-beans so early that a very large proportion are destroyed by frost, and then collect seed from the few
survivors, with care to prevent accidental crosses, and then again get seed from these seedlings, with the same
precautions, the experiment cannot be said to have been even tried. Nor let it be supposed that no differences
in the constitution of seedling kidney-beans ever appear, for an account has been published how much more
hardy some seedlings appeared to be than others.
On the whole, I think we may conclude that habit, use, and disuse, have, in some cases, played a considerable
part in the modification of the constitution, and of the structure of various organs; but that the effects of use
and disuse have often been largely combined with, and sometimes overmastered by, the natural selection of
innate differences.
CORRELATION OF GROWTH.
I mean by this expression that the whole organisation is so tied together during its growth and development,
that when slight variations in any one part occur, and are accumulated through natural selection, other parts
become modified. This is a very important subject, most imperfectly understood. The most obvious case is,
that modifications accumulated solely for the good of the young or larva, will, it may safely be concluded,
affect the structure of the adult; in the same manner as any malconformation affecting the early embryo,
seriously affects the whole organisation of the adult. The several parts of the body which are homologous, and
which, at an early embryonic period, are alike, seem liable to vary in an allied manner: we see this in the right
and left sides of the body varying in the same manner; in the front and hind legs, and even in the jaws and
limbs, varying together, for the lower jaw is believed to be homologous with the limbs. These tendencies, I do
not doubt, may be mastered more or less completely by natural selection: thus a family of stags once existed
with an antler only on one side; and if this had been of any great use to the breed it might probably have been
rendered permanent by natural selection.
Homologous parts, as has been remarked by some authors, tend to cohere; this is often seen in monstrous
plants; and nothing is more common than the union of homologous parts in normal structures, as the union of
the petals of the corolla into a tube. Hard parts seem to affect the form of adjoining soft parts; it is believed by
some authors that the diversity in the shape of the pelvis in birds causes the remarkable diversity in the shape
of their kidneys. Others believe that the shape of the pelvis in the human mother influences by pressure the
shape of the head of the child. In snakes, according to Schlegel, the shape of the body and the manner of
swallowing determine the position of several of the most important viscera.
The nature of the bond of correlation is very frequently quite obscure. M. Is. Geoffroy St. Hilaire has forcibly
remarked, that certain malconformations very frequently, and that others rarely coexist, without our being able
to assign any reason. What can be more singular than the relation between blue eyes and deafness in cats, and
the tortoise-shell colour with the female sex; the feathered feet and skin between the outer toes in pigeons, and
the presence of more or less down on the young birds when first hatched, with the future colour of their
plumage; or, again, the relation between the hair and teeth in the naked Turkish dog, though here probably
homology comes into play? With respect to this latter case of correlation, I think it can hardly be accidental,
that if we pick out the two orders of mammalia which are most abnormal in their dermal coverings, viz.
Cetacea (whales) and Edentata (armadilloes, scaly ant-eaters, etc.), that these are likewise the most abnormal
in their teeth.
I know of no case better adapted to show the importance of the laws of correlation in modifying important
structures, independently of utility and, therefore, of natural selection, than that of the difference between the
outer and inner flowers in some Compositous and Umbelliferous plants. Every one knows the difference in
the ray and central florets of, for instance, the daisy, and this difference is often accompanied with the
abortion of parts of the flower. But, in some Compositous plants, the seeds also differ in shape and sculpture;
and even the ovary itself, with its accessory parts, differs, as has been described by Cassini. These differences
have been attributed by some authors to pressure, and the shape of the seeds in the ray-florets in some
Compositae countenances this idea; but, in the case of the corolla of the Umbelliferae, it is by no means, as
Dr. Hooker informs me, in species with the densest heads that the inner and outer flowers most frequently
differ. It might have been thought that the development of the ray-petals by drawing nourishment from certain
other parts of the flower had caused their abortion; but in some Compositae there is a difference in the seeds
of the outer and inner florets without any difference in the corolla. Possibly, these several differences may be
connected with some difference in the flow of nutriment towards the central and external flowers: we know, at
least, that in irregular flowers, those nearest to the axis are oftenest subject to peloria, and become regular. I
may add, as an instance of this, and of a striking case of correlation, that I have recently observed in some
garden pelargoniums, that the central flower of the truss often loses the patches of darker colour in the two
upper petals; and that when this occurs, the adherent nectary is quite aborted; when the colour is absent from
only one of the two upper petals, the nectary is only much shortened.
With respect to the difference in the corolla of the central and exterior flowers of a head or umbel, I do not
feel at all sure that C. C. Sprengel's idea that the ray-florets serve to attract insects, whose agency is highly
advantageous in the fertilisation of plants of these two orders, is so far-fetched, as it may at first appear: and if
it be advantageous, natural selection may have come into play. But in regard to the differences both in the
internal and external structure of the seeds, which are not always correlated with any differences in the
flowers, it seems impossible that they can be in any way advantageous to the plant: yet in the Umbelliferae
these differences are of such apparent importance--the seeds being in some cases, according to Tausch,
orthospermous in the exterior flowers and coelospermous in the central flowers,--that the elder De Candolle
founded his main divisions of the order on analogous differences. Hence we see that modifications of
structure, viewed by systematists as of high value, may be wholly due to unknown laws of correlated growth,
and without being, as far as we can see, of the slightest service to the species.
We may often falsely attribute to correlation of growth, structures which are common to whole groups of
species, and which in truth are simply due to inheritance; for an ancient progenitor may have acquired through
natural selection some one modification in structure, and, after thousands of generations, some other and
independent modification; and these two modifications, having been transmitted to a whole group of
descendants with diverse habits, would naturally be thought to be correlated in some necessary manner. So,
again, I do not doubt that some apparent correlations, occurring throughout whole orders, are entirely due to
the manner alone in which natural selection can act. For instance, Alph. De Candolle has remarked that
winged seeds are never found in fruits which do not open: I should explain the rule by the fact that seeds
could not gradually become winged through natural selection, except in fruits which opened; so that the
individual plants producing seeds which were a little better fitted to be wafted further, might get an advantage
over those producing seed less fitted for dispersal; and this process could not possibly go on in fruit which did
not open.
The elder Geoffroy and Goethe propounded, at about the same period, their law of compensation or
balancement of growth; or, as Goethe expressed it, "in order to spend on one side, nature is forced to
economise on the other side." I think this holds true to a certain extent with our domestic productions: if
nourishment flows to one part or organ in excess, it rarely flows, at least in excess, to another part; thus it is
difficult to get a cow to give much milk and to fatten readily. The same varieties of the cabbage do not yield
abundant and nutritious foliage and a copious supply of oil-bearing seeds. When the seeds in our fruits
become atrophied, the fruit itself gains largely in size and quality. In our poultry, a large tuft of feathers on the
head is generally accompanied by a diminished comb, and a large beard by diminished wattles. With species
in a state of nature it can hardly be maintained that the law is of universal application; but many good
observers, more especially botanists, believe in its truth. I will not, however, here give any instances, for I see
hardly any way of distinguishing between the effects, on the one hand, of a part being largely developed
through natural selection and another and adjoining part being reduced by this same process or by disuse, and,
on the other hand, the actual withdrawal of nutriment from one part owing to the excess of growth in another
and adjoining part.
I suspect, also, that some of the cases of compensation which have been advanced, and likewise some other
facts, may be merged under a more general principle, namely, that natural selection is continually trying to
economise in every part of the organisation. If under changed conditions of life a structure before useful
becomes less useful, any diminution, however slight, in its development, will be seized on by natural
selection, for it will profit the individual not to have its nutriment wasted in building up an useless structure. I
can thus only understand a fact with which I was much struck when examining cirripedes, and of which many
other instances could be given: namely, that when a cirripede is parasitic within another and is thus protected,
it loses more or less completely its own shell or carapace. This is the case with the male Ibla, and in a truly
extraordinary manner with the Proteolepas: for the carapace in all other cirripedes consists of the three
highly-important anterior segments of the head enormously developed, and furnished with great nerves and
muscles; but in the parasitic and protected Proteolepas, the whole anterior part of the head is reduced to the
merest rudiment attached to the bases of the prehensile antennae. Now the saving of a large and complex
structure, when rendered superfluous by the parasitic habits of the Proteolepas, though effected by slow steps,
would be a decided advantage to each successive individual of the species; for in the struggle for life to which
every animal is exposed, each individual Proteolepas would have a better chance of supporting itself, by less
nutriment being wasted in developing a structure now become useless.
Thus, as I believe, natural selection will always succeed in the long run in reducing and saving every part of
the organisation, as soon as it is rendered superfluous, without by any means causing some other part to be
largely developed in a corresponding degree. And, conversely, that natural selection may perfectly well
succeed in largely developing any organ, without requiring as a necessary compensation the reduction of some
adjoining part.
It seems to be a rule, as remarked by Is. Geoffroy St. Hilaire, both in varieties and in species, that when any
part or organ is repeated many times in the structure of the same individual (as the vertebrae in snakes, and the
stamens in polyandrous flowers) the number is variable; whereas the number of the same part or organ, when
it occurs in lesser numbers, is constant. The same author and some botanists have further remarked that
multiple parts are also very liable to variation in structure. Inasmuch as this "vegetative repetition," to use
Professor Owen's expression, seems to be a sign of low organisation; the foregoing remark seems connected
with the very general opinion of naturalists, that beings low in the scale of nature are more variable than those
which are higher. I presume that lowness in this case means that the several parts of the organisation have
been but little specialised for particular functions; and as long as the same part has to perform diversified
work, we can perhaps see why it should remain variable, that is, why natural selection should have preserved
or rejected each little deviation of form less carefully than when the part has to serve for one special purpose
alone. In the same way that a knife which has to cut all sorts of things may be of almost any shape; whilst a
tool for some particular object had better be of some particular shape. Natural selection, it should never be
forgotten, can act on each part of each being, solely through and for its advantage.
Rudimentary parts, it has been stated by some authors, and I believe with truth, are apt to be highly variable.
We shall have to recur to the general subject of rudimentary and aborted organs; and I will here only add that
their variability seems to be owing to their uselessness, and therefore to natural selection having no power to
check deviations in their structure. Thus rudimentary parts are left to the free play of the various laws of
growth, to the effects of long-continued disuse, and to the tendency to reversion.
A PART DEVELOPED IN ANY SPECIES IN AN EXTRAORDINARY DEGREE OR MANNER, IN
COMPARISON WITH THE SAME PART IN ALLIED SPECIES, TENDS TO BE HIGHLY VARIABLE.
Several years ago I was much struck with a remark, nearly to the above effect, published by Mr. Waterhouse.
I infer also from an observation made by Professor Owen, with respect to the length of the arms of the
ourang-outang, that he has come to a nearly similar conclusion. It is hopeless to attempt to convince any one
of the truth of this proposition without giving the long array of facts which I have collected, and which cannot
possibly be here introduced. I can only state my conviction that it is a rule of high generality. I am aware of
several causes of error, but I hope that I have made due allowance for them. It should be understood that the
rule by no means applies to any part, however unusually developed, unless it be unusually developed in
comparison with the same part in closely allied species. Thus, the bat's wing is a most abnormal structure in
the class mammalia; but the rule would not here apply, because there is a whole group of bats having wings; it
would apply only if some one species of bat had its wings developed in some remarkable manner in
comparison with the other species of the same genus. The rule applies very strongly in the case of secondary
sexual characters, when displayed in any unusual manner. The term, secondary sexual characters, used by
Hunter, applies to characters which are attached to one sex, but are not directly connected with the act of
reproduction. The rule applies to males and females; but as females more rarely offer remarkable secondary
sexual characters, it applies more rarely to them. The rule being so plainly applicable in the case of secondary
sexual characters, may be due to the great variability of these characters, whether or not displayed in any
unusual manner--of which fact I think there can be little doubt. But that our rule is not confined to secondary
sexual characters is clearly shown in the case of hermaphrodite cirripedes; and I may here add, that I
particularly attended to Mr. Waterhouse's remark, whilst investigating this Order, and I am fully convinced
that the rule almost invariably holds good with cirripedes. I shall, in my future work, give a list of the more
remarkable cases; I will here only briefly give one, as it illustrates the rule in its largest application. The
opercular valves of sessile cirripedes (rock barnacles) are, in every sense of the word, very important
structures, and they differ extremely little even in different genera; but in the several species of one genus,
Pyrgoma, these valves present a marvellous amount of diversification: the homologous valves in the different
species being sometimes wholly unlike in shape; and the amount of variation in the individuals of several of
the species is so great, that it is no exaggeration to state that the varieties differ more from each other in the
characters of these important valves than do other species of distinct genera.
As birds within the same country vary in a remarkably small degree, I have particularly attended to them, and
the rule seems to me certainly to hold good in this class. I cannot make out that it applies to plants, and this
would seriously have shaken my belief in its truth, had not the great variability in plants made it particularly
difficult to compare their relative degrees of variability.
When we see any part or organ developed in a remarkable degree or manner in any species, the fair
presumption is that it is of high importance to that species; nevertheless the part in this case is eminently liable
to variation. Why should this be so? On the view that each species has been independently created, with all its
parts as we now see them, I can see no explanation. But on the view that groups of species have descended
from other species, and have been modified through natural selection, I think we can obtain some light. In our
domestic animals, if any part, or the whole animal, be neglected and no selection be applied, that part (for
instance, the comb in the Dorking fowl) or the whole breed will cease to have a nearly uniform character. The
breed will then be said to have degenerated. In rudimentary organs, and in those which have been but little
specialised for any particular purpose, and perhaps in polymorphic groups, we see a nearly parallel natural
case; for in such cases natural selection either has not or cannot come into full play, and thus the organisation
is left in a fluctuating condition. But what here more especially concerns us is, that in our domestic animals
those points, which at the present time are undergoing rapid change by continued selection, are also eminently
liable to variation. Look at the breeds of the pigeon; see what a prodigious amount of difference there is in the
beak of the different tumblers, in the beak and wattle of the different carriers, in the carriage and tail of our
fantails, etc., these being the points now mainly attended to by English fanciers. Even in the sub-breeds, as in
the short-faced tumbler, it is notoriously difficult to breed them nearly to perfection, and frequently
individuals are born which depart widely from the standard. There may be truly said to be a constant struggle
going on between, on the one hand, the tendency to reversion to a less modified state, as well as an innate
tendency to further variability of all kinds, and, on the other hand, the power of steady selection to keep the
breed true. In the long run selection gains the day, and we do not expect to fail so far as to breed a bird as
coarse as a common tumbler from a good short-faced strain. But as long as selection is rapidly going on, there
may always be expected to be much variability in the structure undergoing modification. It further deserves
notice that these variable characters, produced by man's selection, sometimes become attached, from causes
quite unknown to us, more to one sex than to the other, generally to the male sex, as with the wattle of carriers
and the enlarged crop of pouters.
Now let us turn to nature. When a part has been developed in an extraordinary manner in any one species,
compared with the other species of the same genus, we may conclude that this part has undergone an
extraordinary amount of modification, since the period when the species branched off from the common
progenitor of the genus. This period will seldom be remote in any extreme degree, as species very rarely
endure for more than one geological period. An extraordinary amount of modification implies an unusually
large and long-continued amount of variability, which has continually been accumulated by natural selection
for the benefit of the species. But as the variability of the extraordinarily-developed part or organ has been so
great and long-continued within a period not excessively remote, we might, as a general rule, expect still to
find more variability in such parts than in other parts of the organisation, which have remained for a much
longer period nearly constant. And this, I am convinced, is the case. That the struggle between natural
selection on the one hand, and the tendency to reversion and variability on the other hand, will in the course of
time cease; and that the most abnormally developed organs may be made constant, I can see no reason to
doubt. Hence when an organ, however abnormal it may be, has been transmitted in approximately the same
condition to many modified descendants, as in the case of the wing of the bat, it must have existed, according
to my theory, for an immense period in nearly the same state; and thus it comes to be no more variable than
any other structure. It is only in those cases in which the modification has been comparatively recent and
extraordinarily great that we ought to find the GENERATIVE VARIABILITY, as it may be called, still
present in a high degree. For in this case the variability will seldom as yet have been fixed by the continued
selection of the individuals varying in the required manner and degree, and by the continued rejection of those
tending to revert to a former and less modified condition.
The principle included in these remarks may be extended. It is notorious that specific characters are more
variable than generic. To explain by a simple example what is meant. If some species in a large genus of
plants had blue flowers and some had red, the colour would be only a specific character, and no one would be
surprised at one of the blue species varying into red, or conversely; but if all the species had blue flowers, the
colour would become a generic character, and its variation would be a more unusual circumstance. I have
chosen this example because an explanation is not in this case applicable, which most naturalists would
advance, namely, that specific characters are more variable than generic, because they are taken from parts of
less physiological importance than those commonly used for classing genera. I believe this explanation is
partly, yet only indirectly, true; I shall, however, have to return to this subject in our chapter on Classification.
It would be almost superfluous to adduce evidence in support of the above statement, that specific characters
are more variable than generic; but I have repeatedly noticed in works on natural history, that when an author
has remarked with surprise that some IMPORTANT organ or part, which is generally very constant
throughout large groups of species, has DIFFERED considerably in closely-allied species, that it has, also,
been VARIABLE in the individuals of some of the species. And this fact shows that a character, which is
generally of generic value, when it sinks in value and becomes only of specific value, often becomes variable,
though its physiological importance may remain the same. Something of the same kind applies to
monstrosities: at least Is. Geoffroy St. Hilaire seems to entertain no doubt, that the more an organ normally
differs in the different species of the same group, the more subject it is to individual anomalies.
On the ordinary view of each species having been independently created, why should that part of the structure,
which differs from the same part in other independently-created species of the same genus, be more variable
than those parts which are closely alike in the several species? I do not see that any explanation can be given.
But on the view of species being only strongly marked and fixed varieties, we might surely expect to find
them still often continuing to vary in those parts of their structure which have varied within a moderately
recent period, and which have thus come to differ. Or to state the case in another manner:--the points in which
all the species of a genus resemble each other, and in which they differ from the species of some other genus,
are called generic characters; and these characters in common I attribute to inheritance from a common
progenitor, for it can rarely have happened that natural selection will have modified several species, fitted to
more or less widely-different habits, in exactly the same manner: and as these so-called generic characters
have been inherited from a remote period, since that period when the species first branched off from their
common progenitor, and subsequently have not varied or come to differ in any degree, or only in a slight
degree, it is not probable that they should vary at the present day. On the other hand, the points in which
species differ from other species of the same genus, are called specific characters; and as these specific
characters have varied and come to differ within the period of the branching off of the species from a common
progenitor, it is probable that they should still often be in some degree variable,--at least more variable than
those parts of the organisation which have for a very long period remained constant.
In connexion with the present subject, I will make only two other remarks. I think it will be admitted, without
my entering on details, that secondary sexual characters are very variable; I think it also will be admitted that
species of the same group differ from each other more widely in their secondary sexual characters, than in
other parts of their organisation; compare, for instance, the amount of difference between the males of
gallinaceous birds, in which secondary sexual characters are strongly displayed, with the amount of difference
between their females; and the truth of this proposition will be granted. The cause of the original variability of
secondary sexual characters is not manifest; but we can see why these characters should not have been
rendered as constant and uniform as other parts of the organisation; for secondary sexual characters have been
accumulated by sexual selection, which is less rigid in its action than ordinary selection, as it does not entail
death, but only gives fewer offspring to the less favoured males. Whatever the cause may be of the variability
of secondary sexual characters, as they are highly variable, sexual selection will have had a wide scope for
action, and may thus readily have succeeded in giving to the species of the same group a greater amount of
difference in their sexual characters, than in other parts of their structure.
It is a remarkable fact, that the secondary sexual differences between the two sexes of the same species are
generally displayed in the very same parts of the organisation in which the different species of the same genus
differ from each other. Of this fact I will give in illustration two instances, the first which happen to stand on
my list; and as the differences in these cases are of a very unusual nature, the relation can hardly be accidental.
The same number of joints in the tarsi is a character generally common to very large groups of beetles, but in
the Engidae, as Westwood has remarked, the number varies greatly; and the number likewise differs in the
two sexes of the same species: again in fossorial hymenoptera, the manner of neuration of the wings is a
character of the highest importance, because common to large groups; but in certain genera the neuration
differs in the different species, and likewise in the two sexes of the same species. This relation has a clear
meaning on my view of the subject: I look at all the species of the same genus as having as certainly
descended from the same progenitor, as have the two sexes of any one of the species. Consequently, whatever
part of the structure of the common progenitor, or of its early descendants, became variable; variations of this
part would it is highly probable, be taken advantage of by natural and sexual selection, in order to fit the
several species to their several places in the economy of nature, and likewise to fit the two sexes of the same
species to each other, or to fit the males and females to different habits of life, or the males to struggle with
other males for the possession of the females.
Finally, then, I conclude that the greater variability of specific characters, or those which distinguish species
from species, than of generic characters, or those which the species possess in common;--that the frequent
extreme variability of any part which is developed in a species in an extraordinary manner in comparison with
the same part in its congeners; and the not great degree of variability in a part, however extraordinarily it may
be developed, if it be common to a whole group of species;--that the great variability of secondary sexual
characters, and the great amount of difference in these same characters between closely allied species;--that
secondary sexual and ordinary specific differences are generally displayed in the same parts of the
organisation,--are all principles closely connected together. All being mainly due to the species of the same
group having descended from a common progenitor, from whom they have inherited much in common,--to
parts which have recently and largely varied being more likely still to go on varying than parts which have
long been inherited and have not varied,--to natural selection having more or less completely, according to the
lapse of time, overmastered the tendency to reversion and to further variability,--to sexual selection being less
rigid than ordinary selection,--and to variations in the same parts having been accumulated by natural and
sexual selection, and thus adapted for secondary sexual, and for ordinary specific purposes.
DISTINCT SPECIES PRESENT ANALOGOUS VARIATIONS; AND A VARIETY OF ONE SPECIES
OFTEN ASSUMES SOME OF THE CHARACTERS OF AN ALLIED SPECIES, OR REVERTS TO SOME
OF THE CHARACTERS OF AN EARLY PROGENITOR.
These propositions will be most readily understood by looking to our domestic races. The most distinct breeds
of pigeons, in countries most widely apart, present sub-varieties with reversed feathers on the head and
feathers on the feet,--characters not possessed by the aboriginal rock-pigeon; these then are analogous
variations in two or more distinct races. The frequent presence of fourteen or even sixteen tail-feathers in the
pouter, may be considered as a variation representing the normal structure of another race, the fantail. I
presume that no one will doubt that all such analogous variations are due to the several races of the pigeon
having inherited from a common parent the same constitution and tendency to variation, when acted on by
similar unknown influences. In the vegetable kingdom we have a case of analogous variation, in the enlarged
stems, or roots as commonly called, of the Swedish turnip and Ruta baga, plants which several botanists rank
as varieties produced by cultivation from a common parent: if this be not so, the case will then be one of
analogous variation in two so-called distinct species; and to these a third may be added, namely, the common
turnip. According to the ordinary view of each species having been independently created, we should have to
attribute this similarity in the enlarged stems of these three plants, not to the vera causa of community of
descent, and a consequent tendency to vary in a like manner, but to three separate yet closely related acts of
creation.
With pigeons, however, we have another case, namely, the occasional appearance in all the breeds, of
slaty-blue birds with two black bars on the wings, a white rump, a bar at the end of the tail, with the outer
feathers externally edged near their bases with white. As all these marks are characteristic of the parent
rock-pigeon, I presume that no one will doubt that this is a case of reversion, and not of a new yet analogous
variation appearing in the several breeds. We may I think confidently come to this conclusion, because, as we
have seen, these coloured marks are eminently liable to appear in the crossed offspring of two distinct and
differently coloured breeds; and in this case there is nothing in the external conditions of life to cause the
reappearance of the slaty-blue, with the several marks, beyond the influence of the mere act of crossing on the
laws of inheritance.
No doubt it is a very surprising fact that characters should reappear after having been lost for many, perhaps
for hundreds of generations. But when a breed has been crossed only once by some other breed, the offspring
occasionally show a tendency to revert in character to the foreign breed for many generations--some say, for a
dozen or even a score of generations. After twelve generations, the proportion of blood, to use a common
expression, of any one ancestor, is only 1 in 2048; and yet, as we see, it is generally believed that a tendency
to reversion is retained by this very small proportion of foreign blood. In a breed which has not been crossed,
but in which BOTH parents have lost some character which their progenitor possessed, the tendency, whether
strong or weak, to reproduce the lost character might be, as was formerly remarked, for all that we can see to
the contrary, transmitted for almost any number of generations. When a character which has been lost in a
breed, reappears after a great number of generations, the most probable hypothesis is, not that the offspring
suddenly takes after an ancestor some hundred generations distant, but that in each successive generation there
has been a tendency to reproduce the character in question, which at last, under unknown favourable
conditions, gains an ascendancy. For instance, it is probable that in each generation of the barb-pigeon, which
produces most rarely a blue and black-barred bird, there has been a tendency in each generation in the
plumage to assume this colour. This view is hypothetical, but could be supported by some facts; and I can see
no more abstract improbability in a tendency to produce any character being inherited for an endless number
of generations, than in quite useless or rudimentary organs being, as we all know them to be, thus inherited.
Indeed, we may sometimes observe a mere tendency to produce a rudiment inherited: for instance, in the
common snapdragon (Antirrhinum) a rudiment of a fifth stamen so often appears, that this plant must have an
inherited tendency to produce it.
As all the species of the same genus are supposed, on my theory, to have descended from a common parent, it
might be expected that they would occasionally vary in an analogous manner; so that a variety of one species
would resemble in some of its characters another species; this other species being on my view only a
well-marked and permanent variety. But characters thus gained would probably be of an unimportant nature,
for the presence of all important characters will be governed by natural selection, in accordance with the
diverse habits of the species, and will not be left to the mutual action of the conditions of life and of a similar
inherited constitution. It might further be expected that the species of the same genus would occasionally
exhibit reversions to lost ancestral characters. As, however, we never know the exact character of the common
ancestor of a group, we could not distinguish these two cases: if, for instance, we did not know that the
rock-pigeon was not feather-footed or turn-crowned, we could not have told, whether these characters in our
domestic breeds were reversions or only analogous variations; but we might have inferred that the blueness
was a case of reversion, from the number of the markings, which are correlated with the blue tint, and which it
does not appear probable would all appear together from simple variation. More especially we might have
inferred this, from the blue colour and marks so often appearing when distinct breeds of diverse colours are
crossed. Hence, though under nature it must generally be left doubtful, what cases are reversions to an
anciently existing character, and what are new but analogous variations, yet we ought, on my theory,
sometimes to find the varying offspring of a species assuming characters (either from reversion or from
analogous variation) which already occur in some other members of the same group. And this undoubtedly is
the case in nature.
A considerable part of the difficulty in recognising a variable species in our systematic works, is due to its
varieties mocking, as it were, some of the other species of the same genus. A considerable catalogue, also,
could be given of forms intermediate between two other forms, which themselves must be doubtfully ranked
as either varieties or species; and this shows, unless all these forms be considered as independently created
species, that the one in varying has assumed some of the characters of the other, so as to produce the
intermediate form. But the best evidence is afforded by parts or organs of an important and uniform nature
occasionally varying so as to acquire, in some degree, the character of the same part or organ in an allied
species. I have collected a long list of such cases; but here, as before, I lie under a great disadvantage in not
being able to give them. I can only repeat that such cases certainly do occur, and seem to me very remarkable.
I will, however, give one curious and complex case, not indeed as affecting any important character, but from
occurring in several species of the same genus, partly under domestication and partly under nature. It is a case
apparently of reversion. The ass not rarely has very distinct transverse bars on its legs, like those on the legs of
a zebra: it has been asserted that these are plainest in the foal, and from inquiries which I have made, I believe
this to be true. It has also been asserted that the stripe on each shoulder is sometimes double. The shoulder
stripe is certainly very variable in length and outline. A white ass, but NOT an albino, has been described
without either spinal or shoulder-stripe; and these stripes are sometimes very obscure, or actually quite lost, in
dark-coloured asses. The koulan of Pallas is said to have been seen with a double shoulder-stripe. The
hemionus has no shoulder-stripe; but traces of it, as stated by Mr. Blyth and others, occasionally appear: and I
have been informed by Colonel Poole that the foals of this species are generally striped on the legs, and faintly
on the shoulder. The quagga, though so plainly barred like a zebra over the body, is without bars on the legs;
but Dr. Gray has figured one specimen with very distinct zebra-like bars on the hocks.
With respect to the horse, I have collected cases in England of the spinal stripe in horses of the most distinct
breeds, and of ALL colours; transverse bars on the legs are not rare in duns, mouse-duns, and in one instance
in a chestnut: a faint shoulder-stripe may sometimes be seen in duns, and I have seen a trace in a bay horse.
My son made a careful examination and sketch for me of a dun Belgian cart-horse with a double stripe on
each shoulder and with leg-stripes; and a man, whom I can implicitly trust, has examined for me a small dun
Welch pony with THREE short parallel stripes on each shoulder.
In the north-west part of India the Kattywar breed of horses is so generally striped, that, as I hear from
Colonel Poole, who examined the breed for the Indian Government, a horse without stripes is not considered
as purely-bred. The spine is always striped; the legs are generally barred; and the shoulder-stripe, which is
sometimes double and sometimes treble, is common; the side of the face, moreover, is sometimes striped. The
stripes are plainest in the foal; and sometimes quite disappear in old horses. Colonel Poole has seen both gray
and bay Kattywar horses striped when first foaled. I have, also, reason to suspect, from information given me
by Mr. W. W. Edwards, that with the English race-horse the spinal stripe is much commoner in the foal than
in the full-grown animal. Without here entering on further details, I may state that I have collected cases of leg
and shoulder stripes in horses of very different breeds, in various countries from Britain to Eastern China; and
from Norway in the north to the Malay Archipelago in the south. In all parts of the world these stripes occur
far oftenest in duns and mouse-duns; by the term dun a large range of colour is included, from one between
brown and black to a close approach to cream-colour.
I am aware that Colonel Hamilton Smith, who has written on this subject, believes that the several breeds of
the horse have descended from several aboriginal species--one of which, the dun, was striped; and that the
above-described appearances are all due to ancient crosses with the dun stock. But I am not at all satisfied
with this theory, and should be loth to apply it to breeds so distinct as the heavy Belgian cart-horse, Welch
ponies, cobs, the lanky Kattywar race, etc., inhabiting the most distant parts of the world.
Now let us turn to the effects of crossing the several species of the horse-genus. Rollin asserts, that the
common mule from the ass and horse is particularly apt to have bars on its legs. I once saw a mule with its
legs so much striped that any one at first would have thought that it must have been the product of a zebra;
and Mr. W. C. Martin, in his excellent treatise on the horse, has given a figure of a similar mule. In four
coloured drawings, which I have seen, of hybrids between the ass and zebra, the legs were much more plainly
barred than the rest of the body; and in one of them there was a double shoulder-stripe. In Lord Moreton's
famous hybrid from a chestnut mare and male quagga, the hybrid, and even the pure offspring subsequently
produced from the mare by a black Arabian sire, were much more plainly barred across the legs than is even
the pure quagga. Lastly, and this is another most remarkable case, a hybrid has been figured by Dr. Gray (and
he informs me that he knows of a second case) from the ass and the hemionus; and this hybrid, though the ass
seldom has stripes on its legs and the hemionus has none and has not even a shoulder-stripe, nevertheless had
all four legs barred, and had three short shoulder-stripes, like those on the dun Welch pony, and even had
some zebra-like stripes on the sides of its face. With respect to this last fact, I was so convinced that not even
a stripe of colour appears from what would commonly be called an accident, that I was led solely from the
occurrence of the face-stripes on this hybrid from the ass and hemionus, to ask Colonel Poole whether such
face-stripes ever occur in the eminently striped Kattywar breed of horses, and was, as we have seen, answered
in the affirmative.
What now are we to say to these several facts? We see several very distinct species of the horse-genus
becoming, by simple variation, striped on the legs like a zebra, or striped on the shoulders like an ass. In the
horse we see this tendency strong whenever a dun tint appears--a tint which approaches to that of the general
colouring of the other species of the genus. The appearance of the stripes is not accompanied by any change of
form or by any other new character. We see this tendency to become striped most strongly displayed in
hybrids from between several of the most distinct species. Now observe the case of the several breeds of
pigeons: they are descended from a pigeon (including two or three sub-species or geographical races) of a
bluish colour, with certain bars and other marks; and when any breed assumes by simple variation a bluish
tint, these bars and other marks invariably reappear; but without any other change of form or character. When
the oldest and truest breeds of various colours are crossed, we see a strong tendency for the blue tint and bars
and marks to reappear in the mongrels. I have stated that the most probable hypothesis to account for the
reappearance of very ancient characters, is--that there is a TENDENCY in the young of each successive
generation to produce the long-lost character, and that this tendency, from unknown causes, sometimes
prevails. And we have just seen that in several species of the horse-genus the stripes are either plainer or
appear more commonly in the young than in the old. Call the breeds of pigeons, some of which have bred true
for centuries, species; and how exactly parallel is the case with that of the species of the horse-genus! For
myself, I venture confidently to look back thousands on thousands of generations, and I see an animal striped
like a zebra, but perhaps otherwise very differently constructed, the common parent of our domestic horse,
whether or not it be descended from one or more wild stocks, of the ass, the hemionus, quagga, and zebra.
He who believes that each equine species was independently created, will, I presume, assert that each species
has been created with a tendency to vary, both under nature and under domestication, in this particular
manner, so as often to become striped like other species of the genus; and that each has been created with a
strong tendency, when crossed with species inhabiting distant quarters of the world, to produce hybrids
resembling in their stripes, not their own parents, but other species of the genus. To admit this view is, as it
seems to me, to reject a real for an unreal, or at least for an unknown, cause. It makes the works of God a mere
mockery and deception; I would almost as soon believe with the old and ignorant cosmogonists, that fossil
shells had never lived, but had been created in stone so as to mock the shells now living on the sea-shore.

Sunday, October 20, 2013

On the Origin of Species,

Chapter 4

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 that
within the same area, varieties of the same animal can long remain distinct, from haunting different stations,
from breeding at slightly different seasons, or from varieties of the same kind preferring to pair together.
Intercrossing plays a very important part in nature in keeping the individuals of the same species, or of the
same variety, true and uniform in character. It will obviously thus act far more efficiently with those animals
which unite for each birth; but I have already attempted to show that we have reason to believe that occasional
intercrosses take place with all animals and with all plants. Even if these take place only at long intervals, I am
convinced that the young thus produced will gain so much in vigour and fertility over the offspring from
long-continued self-fertilisation, that they will have a better chance of surviving and propagating their kind;and thus, in the long run, the influence of intercrosses, even at rare intervals, will be great. If there exist
organic beings which never intercross, uniformity of character can be retained amongst them, as long as their
conditions of life remain the same, only through the principle of inheritance, and through natural selection
destroying any which depart from the proper type; but if their conditions of life change and they undergo
modification, uniformity of character can be given to their modified offspring, solely by natural selection
preserving the same favourable variations.
Isolation, also, is an important element in the process of natural selection. In a confined or isolated area, if not
very large, the organic and inorganic conditions of life will generally be in a great degree uniform; so that
natural selection will tend to modify all the individuals of a varying species throughout the area in the same
manner in relation to the same conditions. Intercrosses, also, with the individuals of the same species, which
otherwise would have inhabited the surrounding and differently circumstanced districts, will be prevented.
But isolation probably acts more efficiently in checking the immigration of better adapted organisms, after
any physical change, such as of climate or elevation of the land, etc.; and thus new places in the natural
economy of the country are left open for the old inhabitants to struggle for, and become adapted to, through
modifications in their structure and constitution. Lastly, isolation, by checking immigration and consequently
competition, will give time for any new variety to be slowly improved; and this may sometimes be of
importance in the production of new species. If, however, an isolated area be very small, either from being
surrounded by barriers, or from having very peculiar physical conditions, the total number of the individuals
supported on it will necessarily be very small; and fewness of individuals will greatly retard the production of
new species through natural selection, by decreasing the chance of the appearance of favourable variations.
If we turn to nature to test the truth of these remarks, and look at any small isolated area, such as an oceanic
island, although the total number of the species inhabiting it, will be found to be small, as we shall see in our
chapter on geographical distribution; yet of these species a very large proportion are endemic,--that is, have
been produced there, and nowhere else. Hence an oceanic island at first sight seems to have been highly
favourable for the production of new species. But we may thus greatly deceive ourselves, for to ascertain
whether a small isolated area, or a large open area like a continent, has been most favourable for the
production of new organic forms, we ought to make the comparison within equal times; and this we are
incapable of doing.
Although I do not doubt that isolation is of considerable importance in the production of new species, on the
whole I am inclined to believe that largeness of area is of more importance, more especially in the production
of species, which will prove capable of enduring for a long period, and of spreading widely. Throughout a
great and open area, not only will there be a better chance of favourable variations arising from the large
number of individuals of the same species there supported, but the conditions of life are infinitely complex
from the large number of already existing species; and if some of these many species become modified and
improved, others will have to be improved in a corresponding degree or they will be exterminated. Each new
form, also, as soon as it has been much improved, will be able to spread over the open and continuous area,
and will thus come into competition with many others. Hence more new places will be formed, and the
competition to fill them will be more severe, on a large than on a small and isolated area. Moreover, great
areas, though now continuous, owing to oscillations of level, will often have recently existed in a broken
condition, so that the good effects of isolation will generally, to a certain extent, have concurred. Finally, I
conclude that, although small isolated areas probably have been in some respects highly favourable for the
production of new species, yet that the course of modification will generally have been more rapid on large
areas; and what is more important, that the new forms produced on large areas, which already have been
victorious over many competitors, will be those that will spread most widely, will give rise to most new
varieties and species, and will thus play an important part in the changing history of the organic world.
We can, perhaps, on these views, understand some facts which will be again alluded to in our chapter on
geographical distribution; for instance, that the productions of the smaller continent of Australia have
formerly yielded, and apparently are now yielding, before those of the larger Europaeo-Asiatic area. Thus,
 also, it is that continental productions have everywhere become so largely naturalised on islands. On a small
island, the race for life will have been less severe, and there will have been less modification and less
extermination. Hence, perhaps, it comes that the flora of Madeira, according to Oswald Heer, resembles the
extinct tertiary flora of Europe. All fresh-water basins, taken together, make a small area compared with that
of the sea or of the land; and, consequently, the competition between fresh-water productions will have been
less severe than elsewhere; new forms will have been more slowly formed, and old forms more slowly
exterminated. And it is in fresh water that we find seven genera of Ganoid fishes, remnants of a once
preponderant order: and in fresh water we find some of the most anomalous forms now known in the world, as
the Ornithorhynchus and Lepidosiren, which, like fossils, connect to a certain extent orders now widely
separated in the natural scale. These anomalous forms may almost be called living fossils; they have endured
to the present day, from having inhabited a confined area, and from having thus been exposed to less severe
competition.
To sum up the circumstances favourable and unfavourable to natural selection, as far as the extreme intricacy
of the subject permits. I conclude, looking to the future, that for terrestrial productions a large continental
area, which will probably undergo many oscillations of level, and which consequently will exist for long
periods in a broken condition, will be the most favourable for the production of many new forms of life, likely
to endure long and to spread widely. For the area will first have existed as a continent, and the inhabitants, at
this period numerous in individuals and kinds, will have been subjected to very severe competition. When
converted by subsidence into large separate islands, there will still exist many individuals of the same species
on each island: intercrossing on the confines of the range of each species will thus be checked: after physical
changes of any kind, immigration will be prevented, so that new places in the polity of each island will have
to be filled up by modifications of the old inhabitants; and time will be allowed for the varieties in each to
become well modified and perfected. When, by renewed elevation, the islands shall be re-converted into a
continental area, there will again be severe competition: the most favoured or improved varieties will be
enabled to spread: there will be much extinction of the less improved forms, and the relative proportional
numbers of the various inhabitants of the renewed continent will again be changed; and again there will be a
fair field for natural selection to improve still further the inhabitants, and thus produce new species.
That natural selection will always act with extreme slowness, I fully admit. Its action depends on there being
places in the polity of nature, which can be better occupied by some of the inhabitants of the country
undergoing modification of some kind. The existence of such places will often depend on physical changes,
which are generally very slow, and on the immigration of better adapted forms having been checked. But the
action of natural selection will probably still oftener depend on some of the inhabitants becoming slowly
modified; the mutual relations of many of the other inhabitants being thus disturbed. Nothing can be effected,
unless favourable variations occur, and variation itself is apparently always a very slow process. The process
will often be greatly retarded by free intercrossing. Many will exclaim that these several causes are amply
sufficient wholly to stop the action of natural selection. I do not believe so. On the other hand, I do believe
that natural selection will always act very slowly, often only at long intervals of time, and generally on only a
very few of the inhabitants of the same region at the same time. I further believe, that this very slow,
intermittent action of natural selection accords perfectly well with what geology tells us of the rate and
manner at which the inhabitants of this world have changed.
Slow though the process of selection may be, if feeble man can do much by his powers of artificial selection, I
can see no limit to the amount of change, to the beauty and infinite complexity of the coadaptations between
all organic beings, one with another and with their physical conditions of life, which may be effected in the
long course of time by nature's power of selection.
EXTINCTION.
This subject will be more fully discussed in our chapter on Geology; but it must be here alluded to from being
intimately connected with natural selection. Natural selection acts solely through the preservation of variations
in some way advantageous, which consequently endure. But as from the high geometrical powers of increase
of all organic beings, each area is already fully stocked with inhabitants, it follows that as each selected and
favoured form increases in number, so will the less favoured forms decrease and become rare. Rarity, as
geology tells us, is the precursor to extinction. We can, also, see that any form represented by few individuals
will, during fluctuations in the seasons or in the number of its enemies, run a good chance of utter extinction.
But we may go further than this; for as new forms are continually and slowly being produced, unless we
believe that the number of specific forms goes on perpetually and almost indefinitely increasing, numbers
inevitably must become extinct. That the number of specific forms has not indefinitely increased, geology
shows us plainly; and indeed we can see reason why they should not have thus increased, for the number of
places in the polity of nature is not indefinitely great,--not that we have any means of knowing that any one
region has as yet got its maximum of species. Probably no region is as yet fully stocked, for at the Cape of
Good Hope, where more species of plants are crowded together than in any other quarter of the world, some
foreign plants have become naturalised, without causing, as far as we know, the extinction of any natives.
Furthermore, the species which are most numerous in individuals will have the best chance of producing
within any given period favourable variations. We have evidence of this, in the facts given in the second
chapter, showing that it is the common species which afford the greatest number of recorded varieties, or
incipient species. Hence, rare species will be less quickly modified or improved within any given period, and
they will consequently be beaten in the race for life by the modified descendants of the commoner species.
From these several considerations I think it inevitably follows, that as new species in the course of time are
formed through natural selection, others will become rarer and rarer, and finally extinct. The forms which
stand in closest competition with those undergoing modification and improvement, will naturally suffer most.
And we have seen in the chapter on the Struggle for Existence that it is the most closely-allied
forms,--varieties of the same species, and species of the same genus or of related genera,--which, from having
nearly the same structure, constitution, and habits, generally come into the severest competition with each
other. Consequently, each new variety or species, during the progress of its formation, will generally press
hardest on its nearest kindred, and tend to exterminate them. We see the same process of extermination
amongst our domesticated productions, through the selection of improved forms by man. Many curious
instances could be given showing how quickly new breeds of cattle, sheep, and other animals, and varieties of
flowers, take the place of older and inferior kinds. In Yorkshire, it is historically known that the ancient black
cattle were displaced by the long-horns, and that these "were swept away by the short-horns" (I quote the
words of an agricultural writer) "as if by some murderous pestilence."
DIVERGENCE OF CHARACTER.
The principle, which I have designated by this term, is of high importance on my theory, and explains, as I
believe, several important facts. In the first place, varieties, even strongly-marked ones, though having
somewhat of the character of species--as is shown by the hopeless doubts in many cases how to rank
them--yet certainly differ from each other far less than do good and distinct species. Nevertheless, according
to my view, varieties are species in the process of formation, or are, as I have called them, incipient species.
How, then, does the lesser difference between varieties become augmented into the greater difference between
species? That this does habitually happen, we must infer from most of the innumerable species throughout
nature presenting well-marked differences; whereas varieties, the supposed prototypes and parents of future
well-marked species, present slight and ill-defined differences. Mere chance, as we may call it, might cause
one variety to differ in some character from its parents, and the offspring of this variety again to differ from its
parent in the very same character and in a greater degree; but this alone would never account for so habitual
and large an amount of difference as that between varieties of the same species and species of the same genus.
As has always been my practice, let us seek light on this head from our domestic productions. We shall here
find something analogous. A fancier is struck by a pigeon having a slightly shorter beak; another fancier is
struck by a pigeon having a rather longer beak; and on the acknowledged principle that "fanciers do not and
will not admire a medium standard, but like extremes," they both go on (as has actually occurred with
tumbler-pigeons) choosing and breeding from birds with longer and longer beaks, or with shorter and shorter
beaks. Again, we may suppose that at an early period one man preferred swifter horses; another stronger and
more bulky horses. The early differences would be very slight; in the course of time, from the continued
selection of swifter horses by some breeders, and of stronger ones by others, the differences would become
greater, and would be noted as forming two sub-breeds; finally, after the lapse of centuries, the sub-breeds
would become converted into two well-established and distinct breeds. As the differences slowly become
greater, the inferior animals with intermediate characters, being neither very swift nor very strong, will have
been neglected, and will have tended to disappear. Here, then, we see in man's productions the action of what
may be called the principle of divergence, causing differences, at first barely appreciable, steadily to increase,
and the breeds to diverge in character both from each other and from their common parent.
But how, it may be asked, can any analogous principle apply in nature? I believe it can and does apply most
efficiently, from the simple circumstance that the more diversified the descendants from any one species
become in structure, constitution, and habits, by so much will they be better enabled to seize on many and
widely diversified places in the polity of nature, and so be enabled to increase in numbers.
We can clearly see this in the case of animals with simple habits. Take the case of a carnivorous quadruped, of
which the number that can be supported in any country has long ago arrived at its full average. If its natural
powers of increase be allowed to act, it can succeed in increasing (the country not undergoing any change in
its conditions) only by its varying descendants seizing on places at present occupied by other animals: some of
them, for instance, being enabled to feed on new kinds of prey, either dead or alive; some inhabiting new
stations, climbing trees, frequenting water, and some perhaps becoming less carnivorous. The more
diversified in habits and structure the descendants of our carnivorous animal became, the more places they
would be enabled to occupy. What applies to one animal will apply throughout all time to all animals--that is,
if they vary--for otherwise natural selection can do nothing. So it will be with plants. It has been
experimentally proved, that if a plot of ground be sown with one species of grass, and a similar plot be sown
with several distinct genera of grasses, a greater number of plants and a greater weight of dry herbage can thus
be raised. The same has been found to hold good when first one variety and then several mixed varieties of
wheat have been sown on equal spaces of ground. Hence, if any one species of grass were to go on varying,
and those varieties were continually selected which differed from each other in at all the same manner as
distinct species and genera of grasses differ from each other, a greater number of individual plants of this
species of grass, including its modified descendants, would succeed in living on the same piece of ground.
And we well know that each species and each variety of grass is annually sowing almost countless seeds; and
thus, as it may be said, is striving its utmost to increase its numbers. Consequently, I cannot doubt that in the
course of many thousands of generations, the most distinct varieties of any one species of grass would always
have the best chance of succeeding and of increasing in numbers, and thus of supplanting the less distinct
varieties; and varieties, when rendered very distinct from each other, take the rank of species.
The truth of the principle, that the greatest amount of life can be supported by great diversification of
structure, is seen under many natural circumstances. In an extremely small area, especially if freely open to
immigration, and where the contest between individual and individual must be severe, we always find great
diversity in its inhabitants. For instance, I found that a piece of turf, three feet by four in size, which had been
exposed for many years to exactly the same conditions, supported twenty species of plants, and these
belonged to eighteen genera and to eight orders, which shows how much these plants differed from each
other. So it is with the plants and insects on small and uniform islets; and so in small ponds of fresh water.
Farmers find that they can raise most food by a rotation of plants belonging to the most different orders:
nature follows what may be called a simultaneous rotation. Most of the animals and plants which live close
round any small piece of ground, could live on it (supposing it not to be in any way peculiar in its nature), and
may be said to be striving to the utmost to live there; but, it is seen, that where they come into the closest
competition with each other, the advantages of diversification of structure, with the accompanying differences
of habit and constitution, determine that the inhabitants, which thus jostle each other most closely, shall, as a
general rule, belong to what we call different genera and orders.
The same principle is seen in the naturalisation of plants through man's agency in foreign lands. It might have
been expected that the plants which have succeeded in becoming naturalised in any land would generally have
been closely allied to the indigenes; for these are commonly looked at as specially created and adapted for
their own country. It might, also, perhaps have been expected that naturalised plants would have belonged to a
few groups more especially adapted to certain stations in their new homes. But the case is very different; and
Alph. De Candolle has well remarked in his great and admirable work, that floras gain by naturalisation,
proportionally with the number of the native genera and species, far more in new genera than in new species.
To give a single instance: in the last edition of Dr. Asa Gray's 'Manual of the Flora of the Northern United
States,' 260 naturalised plants are enumerated, and these belong to 162 genera. We thus see that these
naturalised plants are of a highly diversified nature. They differ, moreover, to a large extent from the
indigenes, for out of the 162 genera, no less than 100 genera are not there indigenous, and thus a large
proportional addition is made to the genera of these States.
By considering the nature of the plants or animals which have struggled successfully with the indigenes of any
country, and have there become naturalised, we can gain some crude idea in what manner some of the natives
would have had to be modified, in order to have gained an advantage over the other natives; and we may, I
think, at least safely infer that diversification of structure, amounting to new generic differences, would have
been profitable to them.
The advantage of diversification in the inhabitants of the same region is, in fact, the same as that of the
physiological division of labour in the organs of the same individual body--a subject so well elucidated by
Milne Edwards. No physiologist doubts that a stomach by being adapted to digest vegetable matter alone, or
flesh alone, draws most nutriment from these substances. So in the general economy of any land, the more
widely and perfectly the animals and plants are diversified for different habits of life, so will a greater number
of individuals be capable of there supporting themselves. A set of animals, with their organisation but little
diversified, could hardly compete with a set more perfectly diversified in structure. It may be doubted, for
instance, whether the Australian marsupials, which are divided into groups differing but little from each other,
and feebly representing, as Mr. Waterhouse and others have remarked, our carnivorous, ruminant, and rodent
mammals, could successfully compete with these well-pronounced orders. In the Australian mammals, we see
the process of diversification in an early and incomplete stage of development. After the foregoing discussion,
which ought to have been much amplified, we may, I think, assume that the modified descendants of any one
species will succeed by so much the better as they become more diversified in structure, and are thus enabled
to encroach on places occupied by other beings. Now let us see how this principle of great benefit being
derived from divergence of character, combined with the principles of natural selection and of extinction, will
tend to act.
The accompanying diagram will aid us in understanding this rather perplexing subject. Let A to L represent
the species of a genus large in its own country; these species are supposed to resemble each other in unequal
degrees, as is so generally the case in nature, and as is represented in the diagram by the letters standing at
unequal distances. I have said a large genus, because we have seen in the second chapter, that on an average
more of the species of large genera vary than of small genera; and the varying species of the large genera
present a greater number of varieties. We have, also, seen that the species, which are the commonest and the
most widely-diffused, vary more than rare species with restricted ranges. Let (A) be a common,
widely-diffused, and varying species, belonging to a genus large in its own country. The little fan of diverging
dotted lines of unequal lengths proceeding from (A), may represent its varying offspring. The variations are
supposed to be extremely slight, but of the most diversified nature; they are not supposed all to appear
simultaneously, but often after long intervals of time; nor are they all supposed to endure for equal periods.
Only those variations which are in some way profitable will be preserved or naturally selected. And here the
importance of the principle of benefit being derived from divergence of character comes in; for this will
generally lead to the most different or divergent variations (represented by the outer dotted lines) being
preserved and accumulated by natural selection. When a dotted line reaches one of the horizontal lines, and is
there marked by a small numbered letter, a sufficient amount of variation is supposed to have been
accumulated to have formed a fairly well-marked variety, such as would be thought worthy of record in a
systematic work.
The intervals between the horizontal lines in the diagram, may represent each a thousand generations; but it
would have been better if each had represented ten thousand generations. After a thousand generations,
species (A) is supposed to have produced two fairly well-marked varieties, namely a1 and m1. These two
varieties will generally continue to be exposed to the same conditions which made their parents variable, and
the tendency to variability is in itself hereditary, consequently they will tend to vary, and generally to vary in
nearly the same manner as their parents varied. Moreover, these two varieties, being only slightly modified
forms, will tend to inherit those advantages which made their common parent (A) more numerous than most
of the other inhabitants of the same country; they will likewise partake of those more general advantages
which made the genus to which the parent-species belonged, a large genus in its own country. And these
circumstances we know to be favourable to the production of new varieties.
If, then, these two varieties be variable, the most divergent of their variations will generally be preserved
during the next thousand generations. And after this interval, variety a1 is supposed in the diagram to have
produced variety a2, which will, owing to the principle of divergence, differ more from (A) than did variety
a1. Variety m1 is supposed to have produced two varieties, namely m2 and s2, differing from each other, and
more considerably from their common parent (A). We may continue the process by similar steps for any
length of time; some of the varieties, after each thousand generations, producing only a single variety, but in a
more and more modified condition, some producing two or three varieties, and some failing to produce any.
Thus the varieties or modified descendants, proceeding from the common parent (A), will generally go on
increasing in number and diverging in character. In the diagram the process is represented up to the
ten-thousandth generation, and under a condensed and simplified form up to the fourteen-thousandth
generation.
But I must here remark that I do not suppose that the process ever goes on so regularly as is represented in the
diagram, though in itself made somewhat irregular. I am far from thinking that the most divergent varieties
will invariably prevail and multiply: a medium form may often long endure, and may or may not produce
more than one modified descendant; for natural selection will always act according to the nature of the places
which are either unoccupied or not perfectly occupied by other beings; and this will depend on infinitely
complex relations. But as a general rule, the more diversified in structure the descendants from any one
species can be rendered, the more places they will be enabled to seize on, and the more their modified
progeny will be increased. In our diagram the line of succession is broken at regular intervals by small
numbered letters marking the successive forms which have become sufficiently distinct to be recorded as
varieties. But these breaks are imaginary, and might have been inserted anywhere, after intervals long enough
to have allowed the accumulation of a considerable amount of divergent variation.
As all the modified descendants from a common and widely-diffused species, belonging to a large genus, will
tend to partake of the same advantages which made their parent successful in life, they will generally go on
multiplying in number as well as diverging in character: this is represented in the diagram by the several
divergent branches proceeding from (A). The modified offspring from the later and more highly improved
branches in the lines of descent, will, it is probable, often take the place of, and so destroy, the earlier and less
improved branches: this is represented in the diagram by some of the lower branches not reaching to the upper
horizontal lines. In some cases I do not doubt that the process of modification will be confined to a single line
of descent, and the number of the descendants will not be increased; although the amount of divergent
modification may have been increased in the successive generations. This case would be represented in the
diagram, if all the lines proceeding from (A) were removed, excepting that from a1 to a10. In the same way,
for instance, the English race-horse and English pointer have apparently both gone on slowly diverging in
character from their original stocks, without either having given off any fresh branches or races.
After ten thousand generations, species (A) is supposed to have produced three forms, a10, f10, and m10,
which, from having diverged in character during the successive generations, will have come to differ largely,
but perhaps unequally, from each other and from their common parent. If we suppose the amount of change
between each horizontal line in our diagram to be excessively small, these three forms may still be only
well-marked varieties; or they may have arrived at the doubtful category of sub-species; but we have only to
suppose the steps in the process of modification to be more numerous or greater in amount, to convert these
three forms into well-defined species: thus the diagram illustrates the steps by which the small differences
distinguishing varieties are increased into the larger differences distinguishing species. By continuing the
same process for a greater number of generations (as shown in the diagram in a condensed and simplified
manner), we get eight species, marked by the letters between a14 and m14, all descended from (A). Thus, as I
believe, species are multiplied and genera are formed.
In a large genus it is probable that more than one species would vary. In the diagram I have assumed that a
second species (I) has produced, by analogous steps, after ten thousand generations, either two well-marked
varieties (w10 and z10) or two species, according to the amount of change supposed to be represented
between the horizontal lines. After fourteen thousand generations, six new species, marked by the letters n14
to z14, are supposed to have been produced. In each genus, the species, which are already extremely different
in character, will generally tend to produce the greatest number of modified descendants; for these will have
the best chance of filling new and widely different places in the polity of nature: hence in the diagram I have
chosen the extreme species (A), and the nearly extreme species (I), as those which have largely varied, and
have given rise to new varieties and species. The other nine species (marked by capital letters) of our original
genus, may for a long period continue transmitting unaltered descendants; and this is shown in the diagram by
the dotted lines not prolonged far upwards from want of space.
But during the process of modification, represented in the diagram, another of our principles, namely that of
extinction, will have played an important part. As in each fully stocked country natural selection necessarily
acts by the selected form having some advantage in the struggle for life over other forms, there will be a
constant tendency in the improved descendants of any one species to supplant and exterminate in each stage
of descent their predecessors and their original parent. For it should be remembered that the competition will
generally be most severe between those forms which are most nearly related to each other in habits,
constitution, and structure. Hence all the intermediate forms between the earlier and later states, that is
between the less and more improved state of a species, as well as the original parent-species itself, will
generally tend to become extinct. So it probably will be with many whole collateral lines of descent, which
will be conquered by later and improved lines of descent. If, however, the modified offspring of a species get
into some distinct country, or become quickly adapted to some quite new station, in which child and parent do
not come into competition, both may continue to exist.
If then our diagram be assumed to represent a considerable amount of modification, species (A) and all the
earlier varieties will have become extinct, having been replaced by eight new species (a14 to m14); and (I)
will have been replaced by six (n14 to z14) new species.
But we may go further than this. The original species of our genus were supposed to resemble each other in
unequal degrees, as is so generally the case in nature; species (A) being more nearly related to B, C, and D,
than to the other species; and species (I) more to G, H, K, L, than to the others. These two species (A) and (I),
were also supposed to be very common and widely diffused species, so that they must originally have had
some advantage over most of the other species of the genus. Their modified descendants, fourteen in number
at the fourteen-thousandth generation, will probably have inherited some of the same advantages: they have
also been modified and improved in a diversified manner at each stage of descent, so as to have become
adapted to many related places in the natural economy of their country. It seems, therefore, to me extremely
probable that they will have taken the places of, and thus exterminated, not only their parents (A) and (I), but
likewise some of the original species which were most nearly related to their parents. Hence very few of the
original species will have transmitted offspring to the fourteen-thousandth generation. We may suppose that
only one (F), of the two species which were least closely related to the other nine original species, has
transmitted descendants to this late stage of descent.
The new species in our diagram descended from the original eleven species, will now be fifteen in number.
Owing to the divergent tendency of natural selection, the extreme amount of difference in character between
species a14 and z14 will be much greater than that between the most different of the original eleven species.
The new species, moreover, will be allied to each other in a widely different manner. Of the eight descendants
from (A) the three marked a14, q14, p14, will be nearly related from having recently branched off from a10;
b14 and f14, from having diverged at an earlier period from a5, will be in some degree distinct from the three
first-named species; and lastly, o14, e14, and m14, will be nearly related one to the other, but from having
diverged at the first commencement of the process of modification, will be widely different from the other
five species, and may constitute a sub-genus or even a distinct genus.
The six descendants from (I) will form two sub-genera or even genera. But as the original species (I) differed
largely from (A), standing nearly at the extreme points of the original genus, the six descendants from (I) will,
owing to inheritance, differ considerably from the eight descendants from (A); the two groups, moreover, are
supposed to have gone on diverging in different directions. The intermediate species, also (and this is a very
important consideration), which connected the original species (A) and (I), have all become, excepting (F),
extinct, and have left no descendants. Hence the six new species descended from (I), and the eight descended
from (A), will have to be ranked as very distinct genera, or even as distinct sub-families.
Thus it is, as I believe, that two or more genera are produced by descent, with modification, from two or more
species of the same genus. And the two or more parent-species are supposed to have descended from some
one species of an earlier genus. In our diagram, this is indicated by the broken lines, beneath the capital
letters, converging in sub-branches downwards towards a single point; this point representing a single species,
the supposed single parent of our several new sub-genera and genera.
It is worth while to reflect for a moment on the character of the new species F14, which is supposed not to
have diverged much in character, but to have retained the form of (F), either unaltered or altered only in a
slight degree. In this case, its affinities to the other fourteen new species will be of a curious and circuitous
nature. Having descended from a form which stood between the two parent-species (A) and (I), now supposed
to be extinct and unknown, it will be in some degree intermediate in character between the two groups
descended from these species. But as these two groups have gone on diverging in character from the type of
their parents, the new species (F14) will not be directly intermediate between them, but rather between types
of the two groups; and every naturalist will be able to bring some such case before his mind.
In the diagram, each horizontal line has hitherto been supposed to represent a thousand generations, but each
may represent a million or hundred million generations, and likewise a section of the successive strata of the
earth's crust including extinct remains. We shall, when we come to our chapter on Geology, have to refer
again to this subject, and I think we shall then see that the diagram throws light on the affinities of extinct
beings, which, though generally belonging to the same orders, or families, or genera, with those now living,
yet are often, in some degree, intermediate in character between existing groups; and we can understand this
fact, for the extinct species lived at very ancient epochs when the branching lines of descent had diverged less.
I see no reason to limit the process of modification, as now explained, to the formation of genera alone. If, in
our diagram, we suppose the amount of change represented by each successive group of diverging dotted lines
to be very great, the forms marked a14 to p14, those marked b14 and f14, and those marked o14 to m14, will
form three very distinct genera. We shall also have two very distinct genera descended from (I) and as these
latter two genera, both from continued divergence of character and from inheritance from a different parent,
will differ widely from the three genera descended from (A), the two little groups of genera will form two
distinct families, or even orders, according to the amount of divergent modification supposed to be
represented in the diagram. And the two new families, or orders, will have descended from two species of the
original genus; and these two species are supposed to have descended from one species of a still more ancient
and unknown genus.
We have seen that in each country it is the species of the larger genera which oftenest present varieties or
incipient species. This, indeed, might have been expected; for as natural selection acts through one form
having some advantage over other forms in the struggle for existence, it will chiefly act on those which
already have some advantage; and the largeness of any group shows that its species have inherited from a
common ancestor some advantage in common. Hence, the struggle for the production of new and modified
descendants, will mainly lie between the larger groups, which are all trying to increase in number. One large
group will slowly conquer another large group, reduce its numbers, and thus lessen its chance of further
variation and improvement. Within the same large group, the later and more highly perfected sub-groups,
from branching out and seizing on many new places in the polity of Nature, will constantly tend to supplant
and destroy the earlier and less improved sub-groups. Small and broken groups and sub-groups will finally
tend to disappear. Looking to the future, we can predict that the groups of organic beings which are now large
and triumphant, and which are least broken up, that is, which as yet have suffered least extinction, will for a
long period continue to increase. But which groups will ultimately prevail, no man can predict; for we well
know that many groups, formerly most extensively developed, have now become extinct. Looking still more
remotely to the future, we may predict that, owing to the continued and steady increase of the larger groups, a
multitude of smaller groups will become utterly extinct, and leave no modified descendants; and consequently
that of the species living at any one period, extremely few will transmit descendants to a remote futurity. I
shall have to return to this subject in the chapter on Classification, but I may add that on this view of
extremely few of the more ancient species having transmitted descendants, and on the view of all the
descendants of the same species making a class, we can understand how it is that there exist but very few
classes in each main division of the animal and vegetable kingdoms. Although extremely few of the most
ancient species may now have living and modified descendants, yet at the most remote geological period, the
earth may have been as well peopled with many species of many genera, families, orders, and classes, as at
the present day.