On the Origin of Species

Abridged from the first edition (1859) by Richard Dawkins and Isabel Morgan. © CSA/Strathmore under Creative Commons.

Charles Darwin

Richard Dawkins

Charles Darwin had begun to write what was called his "big book" on species in 1856. However, in June of 1858, he received the famous letter from Alfred Russel Wallace outlining Wallace's independent exposition of evolution by natural selection. Advised by his friends Charles Lyell and Joseph Hooker, Darwin realized that he hadn't time to produce the big book, but had rather to write as soon as possible a version of his argument in a crisp and accessible form. The result was On the Origin of Species, which finally appeared on November 24, 1859.

Darwin had originally planned to title this work An Abstract of an Essay on the Origin of Species and Varieties through Natural Selection, indicating that the book was a short version of a much larger project. His publisher persuaded him to drop the "abstract," and in any event, On the Origin of Species turned out to be a very substantial work of some 150,000 words in the first edition. The book became larger still in later editions.

Although it does not amplify examples and evidence to make its case in the way that Darwin might have intended for the big book (which was in the end never written), On the Origin of Species is itself reasonably open to abridgement for modern readers. Particularly in university courses with reading lists that include other large works by Darwin as well as secondary authors on evolution, there is a need for an intelligently abridged version of the Origin.

This abridgement of the Origin of Species was created by Richard Dawkins with the help of Isabel Morgan for CSA Word. By carefully choosing the most compelling material from the book, they have reduced it to just under 59,000 words from the original 150,000 words of the first edition. Professor Dawkins's total reading time for this is 5hrs 53 mins.

Richard Dawkins is a gifted reader for the medium of audiobooks. In the case of the Origin of Species, however, his talent is especially notable. Backed by a knowledge of biology and a commitment to evolutionary science, his spoken stresses and sense of emotional animation are palpable. On the Origin of Species comes across not simply as a catalogue of facts, but as an argument. There is intellectual passion in Richard Dawkins's reading that perceptive listeners are bound to hear.

The audio version of this abridgement, produced by Nicholas Jones, may be purchased HERE. Also available is Richard Dawkins's abridgement of Darwin's The Voyage of the Beagle, as well as his own most recent book, The Greatest Show on Earth: the Evidence for Evolution, where he is joined in the reading by his wife, artist and actress Lalla Ward. Listening to Darwin's ideas come alive in the voice of these two Darwinians is an experience. I hope it will give you as much pleasure as it has brought to me. — Denis Dutton, University of Canterbury, New Zealand







When on board H.M.S. ‘Beagle,’ as naturalist, I was much struck with certain facts in the distribution of the inhabitants of South America, and in the geological relations of the present to the past inhabitants of that continent. These facts seemed to me to throw some light on the origin of species — that mystery of mysteries, as it has been called by one of our greatest philosophers. On my return home, ... [a]fter five years’ work I allowed myself to speculate on the subject, and drew up some short notes; these I enlarged in 1844 into a sketch of the conclusions, which then seemed to me probable: from that period to the present day I have steadily pursued the same object. I hope that I may be excused for entering on these personal details, as I give them to show that I have not been hasty in coming to a decision.

My work is now nearly finished; but as it will take me two or three more years to complete it, and as my health is far from strong, I have been urged to publish this Abstract. I have more especially been induced to do this, as Mr. Wallace, who is now studying the natural history of the Malay archipelago, has arrived at almost exactly the same general conclusions that I have on the origin of species. .... Sir C[harles] Lyell and Dr. Hooker, who both knew of my work ... honoured me by thinking it advisable to publish, with Mr. Wallace’s excellent memoir, some brief extracts from my manuscripts.

This Abstract, which I now publish, must necessarily be imperfect. I cannot here give references and authorities for my several statements; and I must trust to the reader reposing some confidence in my accuracy. .... I am well aware that scarcely a single point is discussed in this volume on which facts cannot be adduced, often apparently leading to conclusions directly opposite to those at which I have arrived. A fair result can be obtained only by fully stating and balancing the facts and arguments on both sides of each question; and this cannot possibly be here done.


In considering the Origin of Species, it is quite conceivable that a naturalist, reflecting on the mutual affinities of organic beings, on their embryological relations, their geographical distribution, geological succession, and other such facts, might come to the conclusion that [such] each species had not been independently created, but had descended, like varieties, from other species. Nevertheless, such a conclusion, even if well founded, would be unsatisfactory, until it could be shown how the innumerable species inhabiting this world have been modified, so as to acquire that perfection of structure and coadaptation which most justly excites our admiration. Naturalists continually refer to external conditions, such as climate, food, etc., as the only possible cause of variation. In one very limited sense, as we shall hereafter see, this may be true; but it is preposterous to attribute to mere external conditions, the structure, for instance, of the woodpecker, with its feet, tail, beak, and tongue, so admirably adapted to catch insects under the bark of trees. In the case of the mistletoe, which draws its nourishment from certain trees, which has seeds that must be transported by certain birds, and which has flowers with separate sexes absolutely requiring the agency of certain insects to bring pollen from one flower to the other, it is equally preposterous to account for the structure of this parasite, with its relations to several distinct organic beings, by the effects of external conditions, or of habit, or of the volition of the plant itself.

The author of the ‘Vestiges of Creation’ would, I presume, say that, after a certain unknown number of generations, some bird had given birth to a woodpecker, and some plant to the misseltoe, and that these had been produced perfect as we now see them; but this assumption seems to me to be no explanation, for it leaves the case of the coadaptations of organic beings to each other and to their physical conditions of life, untouched and unexplained.

It is, therefore, of the highest importance to gain a clear insight into the means of modification and coadaptation. At the commencement of my observations it seemed to me probable that a careful study of domesticated animals and of cultivated plants would offer the best chance of making out this obscure problem. Nor have I been disappointed; in this and in all other perplexing cases I have invariably found that our knowledge, imperfect though it be, of variation under domestication, afforded the best and safest clue. I may venture to express my conviction of the high value of such studies, although they have been very commonly neglected by naturalists.

From these considerations, I shall devote the first chapter of this Abstract to Variation under Domestication. We shall thus see that a large amount of hereditary modification is at least possible, and, what is equally or more important, we shall see how great is the power of man in accumulating by his Selection successive slight variations. I will then pass on to the variability of species in a state of nature; but I shall, unfortunately, be compelled to treat this subject far too briefly, as it can be treated properly only by giving long catalogues of facts. We shall, however, be enabled to discuss what circumstances are most favourable to variation. In the next chapter the Struggle for Existence amongst all organic beings throughout the world, which inevitably follows from their high geometrical powers of increase, will be treated of. This is the doctrine of Malthus, applied to the whole animal and vegetable kingdoms. As many more individuals of each species are born than can possibly survive; and as, consequently, there is a frequently recurring struggle for existence, it follows that any being, if it vary however slightly in any manner profitable to itself, under the complex and sometimes varying conditions of life, will have a better chance of surviving, and thus be naturally selected. From the strong principle of inheritance, any selected variety will tend to propagate its new and modified form.


Although much remains obscure, and will long remain obscure, I can entertain no doubt, after the most deliberate study and dispassionate judgment of which I am capable, that the view which most naturalists entertain, and which I formerly entertained — namely, that each species has been independently created — is erroneous. I am fully convinced that species are not immutable; but that those belonging to what are called the same genera are lineal descendants of some other and generally extinct species, in the same manner as the acknowledged varieties of any one species are the descendants of that species. Furthermore, I am convinced that Natural Selection has been the main but not exclusive means of modification.



Causes of Variability. Effects of Habit. Correlation of Growth. Inheritance. Character of Domestic Varieties. Difficulty of distinguishing between Varieties and Species. Origin of Domestic Varieties from one or more Species. Domestic Pigeons, their Differences and Origin. Principle of Selection anciently followed, its Effects. Methodical and Unconscious Selection. Unknown Origin of our Domestic Productions. Circumstances favourable to Man’s power of Selection.

When we look to the individuals of the same variety or sub-variety of our older cultivated plants and animals, one of the first points which strikes us, is, that they generally differ much more from each other, than do the individuals of any one species or variety in a state of nature. When we reflect on the vast diversity of the plants and animals which have been cultivated, and which have varied during all ages under the most different climates and treatment, I think we are driven to conclude that this greater variability is simply due to our domestic productions having been raised under conditions of life not so uniform as, and somewhat different from, those to which the parent-species have been exposed under nature. There is, also, I think, some probability in the view propounded by Andrew Knight, that this variability may be partly connected with excess of food. It seems pretty clear that organic beings must be exposed during several generations to the new conditions of life to cause any appreciable amount of variation; and that when the organisation has once begun to vary, it generally continues to vary for many generations. No case is on record of a variable being ceasing to be variable under cultivation. Our oldest cultivated plants, such as wheat, still often yield new varieties: our oldest domesticated animals are still capable of rapid improvement or modification.


There are many laws regulating variation, some few of which can be dimly seen, and will be hereafter briefly mentioned. I will here only allude to what may be called correlation of growth. Any change in the embryo or larva will almost certainly entail changes in the mature animal. In monstrosities, the correlations between quite distinct parts are very curious; ... . Breeders believe that long limbs are almost always accompanied by an elongated head. Some instances of correlation are quite whimsical; thus cats with blue eyes are invariably deaf; colour and constitutional peculiarities go together, ... it appears that white sheep and pigs are differently affected from coloured individuals by certain vegetable poisons. Hairless dogs have imperfect teeth. ... pigeons with feathered feet have skin between their outer toes; pigeons with short beaks have small feet, and those with long beaks large feet. Hence, if man goes on selecting, and thus augmenting, any peculiarity, he will almost certainly unconsciously modify other parts of the structure, owing to the mysterious laws of the correlation of growth.


Any variation which is not inherited is unimportant for us. But the number and diversity of inheritable deviations of structure, both those of slight and those of considerable physiological importance, is endless. ... No breeder doubts how strong is the tendency to inheritance: like produces like is his fundamental belief: doubts have been thrown on this principle by theoretical writers alone. When a deviation appears not unfrequently, and we see it in the father and child, we cannot tell whether it may not be due to the same original cause acting on both; but when amongst individuals, apparently exposed to the same conditions, any very rare deviation, due to some extraordinary combination of circumstances, appears in the parent — say, once amongst several million individuals — and it reappears in the child, the mere doctrine of chances almost compels us to attribute its reappearance to inheritance. Every one must have heard of cases of albinism, prickly skin, hairy bodies, etc., appearing in several members of the same family. If strange and rare deviations of structure are truly inherited, less strange and commoner deviations may be freely admitted to be inheritable. Perhaps the correct way of viewing the whole subject, would be, to look at the inheritance of every character whatever as the rule, and non-inheritance as the anomaly.

The laws governing inheritance are quite unknown; no one can say why the same peculiarity in different individuals of the same species, and in individuals of different species, is sometimes inherited and sometimes not so; why the child often reverts in certain characters to its grandfather or grandmother or other much more remote ancestor; why a peculiarity is often transmitted from one sex to both sexes or to one sex alone, more commonly but not exclusively to the like sex. It is a fact of some little importance to us, that peculiarities appearing in the males of our domestic breeds are often transmitted either exclusively, or in a much greater degree, to males alone. A much more important rule, which I think may be trusted, is that, at whatever period of life a peculiarity first appears, it tends to appear in the offspring at a corresponding age, though sometimes earlier. In many cases this could not be otherwise: thus the inherited peculiarities in the horns of cattle could appear only in the offspring when nearly mature; peculiarities in the silkworm are known to appear at the corresponding caterpillar or cocoon stage. But hereditary diseases and some other facts make me believe that the rule has a wider extension, and that when there is no apparent reason why a peculiarity should appear at any particular age, yet that it does tend to appear in the offspring at the same period at which it first appeared in the parent. I believe this rule to be of the highest importance in explaining the laws of embryology. These remarks are of course confined to the first appearance of the peculiarity, and not to its primary cause, which may have acted on the ovules or male element; in nearly the same manner as in the crossed offspring from a short-horned cow by a long-horned bull, the greater length of horn, though appearing late in life, is clearly due to the male element.

Having alluded to the subject of reversion, I may here refer to a statement often made by naturalists — namely, that our domestic varieties, when run wild, gradually but certainly revert in character to their aboriginal stocks. Hence it has been argued that no deductions can be drawn from domestic races to species in a state of nature. .... But there is not a shadow of evidence in favour of this view: to assert that we could not breed our cart and race-horses, long and short-horned cattle, and poultry of various breeds, and esculent vegetables, for an almost infinite number of generations, would be opposed to all experience. I may add, that when under nature the conditions of life do change, variations and reversions of character probably do occur; but natural selection, as will hereafter be explained, will determine how far the new characters thus arising shall be preserved.

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

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

It has often been assumed that man has chosen for domestication animals and plants having an extraordinary inherent tendency to vary, and likewise to withstand diverse climates. I do not dispute that these capacities have added largely to the value of most of our domesticated productions; but how could a savage possibly know, when he first tamed an animal, whether it would vary in succeeding generations, and whether it would endure other climates? Has the ... small power of endurance of warmth by the rein-deer, or of cold by the common camel, prevented their domestication? I cannot doubt that if other animals and plants, equal in number to our domesticated productions, and belonging to equally diverse classes and countries, were taken from a state of nature, and could be made to breed for an equal number of generations under domestication, they would vary on an average as largely as the parent species of our existing domesticated productions have varied.

In the case of most of our anciently domesticated animals and plants, I do not think it is possible to come to any definite conclusion, whether they have descended from one or several species. ....

The whole subject must, I think, remain vague; nevertheless, I may, without here entering on any details, state that, from geographical and other considerations, I think it highly probable that our domestic dogs have descended from several wild species. In regard to sheep and goats I can form no opinion. .... With respect to horses, from reasons which I cannot give here, I am doubtfully inclined to believe, in opposition to several authors, that all the races have descended from one wild stock. .... In regard to ducks and rabbits, the breeds of which differ considerably from each other in structure, I do not doubt that they all have descended from the common wild duck and rabbit.

The doctrine of the origin of our several domestic races from several aboriginal stocks, has been carried to an absurd extreme by some authors. They believe that every race which breeds true, let the distinctive characters be ever so slight, has had its wild prototype. At this rate there must have existed at least a score of species of wild cattle, as many sheep, and several goats in Europe alone, and several even within Great Britain. One author believes that there formerly existed in Great Britain eleven wild species of sheep peculiar to it! When we bear in mind that Britain has now hardly one peculiar mammal, and France but few distinct from those of Germany and conversely, and so with Hungary, Spain, etc., but that each of these kingdoms possesses several peculiar breeds of cattle, sheep, etc., we must admit that many domestic breeds have originated in Europe; for whence could they have been derived, as these several countries do not possess a number of peculiar species as distinct parent-stocks? So it is in India. Even in the case of the domestic dogs of the whole world, which I fully admit have probably descended from several wild species, I cannot doubt that there has been an immense amount of inherited variation. Who can believe that animals closely resembling the Italian greyhound, the bloodhound, the bull-dog, or Blenheim spaniel, etc. — so unlike all wild Canidae — ever existed freely in a state of nature? It has often been loosely said that all our races of dogs have been produced by the crossing of a few aboriginal species; but by crossing we can get only forms in some degree intermediate between their parents; and if we account for our several domestic races by this process, we must admit the former existence of the most extreme forms, as the Italian greyhound, bloodhound, bull-dog, etc., in the wild state. Moreover, the possibility of making distinct races by crossing has been greatly exaggerated. .... The offspring from the first cross between two pure breeds is tolerably and sometimes (as I have found with pigeons) extremely uniform, and everything seems simple enough; but when these mongrels are crossed one with another for several generations, hardly two of them will be alike, and then the extreme difficulty, or rather utter hopelessness, of the task becomes apparent. Certainly, a breed intermediate between two very distinct breeds could not be got without extreme care and long-continued

On the Breeds of the Domestic Pigeon Believing that it is always best to study some special group, I have, after deliberation, taken up domestic pigeons. I have kept every breed which I could purchase or obtain, and have been most kindly favoured with skins from several quarters of the world .... Many treatises in different languages have been published on pigeons, and some of them are very important, as being of considerable antiquity. I have associated with several eminent fanciers, and have been permitted to join two of the London Pigeon Clubs. The diversity of the breeds is something astonishing. Compare the English carrier and the short-faced tumbler, and see the wonderful difference in their beaks, entailing corresponding differences in their skulls. The carrier, more especially the male bird, is also remarkable from the wonderful development of the carunculated skin about the head, and this is accompanied by greatly elongated eyelids, very large external orifices to the nostrils, and a wide gape of mouth. The short-faced tumbler has a beak in outline almost like that of a finch; and the common tumbler has the singular and strictly inherited habit of flying at a great height in a compact flock, and tumbling in the air head over heels.... The barb is allied to the carrier, but, instead of a very long beak, has a very short and very broad one. The pouter has a much elongated body, wings, and legs; and its enormously developed crop, which it glories in inflating, may well excite astonishment and even laughter.... The Jacobin has the feathers so much reversed along the back of the neck that they form a hood, and it has, proportionally to its size, much elongated wing and tail feathers. The trumpeter and laugher, as their names express, utter a very different coo from the other breeds....

In the skeletons of the several breeds, the development of the bones of the face in length and breadth and curvature differs enormously. The shape, as well as the breadth and length of the ramus of the lower jaw, varies in a highly remarkable manner. The number of the caudal and sacral vertebrae vary; as does the number of the ribs, together with their relative breadth and the presence of processes.... The proportional width of the gape of mouth, the proportional length of the eyelids, of the orifice of the nostrils, of the tongue (not always in strict correlation with the length of beak), the size of the crop and of the upper part of the oesophagus.... the number of the primary wing and caudal feathers; the relative length of wing and tail to each other and to the body.... are all points of structure which are variable. The period at which the perfect plumage is acquired varies, as does the state of the down with which the nestling birds are clothed when hatched. The shape and size of the eggs vary. The manner of flight differs remarkably; as does in some breeds the voice and disposition. Lastly, in certain breeds, the males and females have come to differ to a slight degree from each other.


Great as the differences are between the breeds of pigeons, I am fully convinced that the common opinion of naturalists is correct, namely, that all have descended from the rock-pigeon (Columba livia), including under this term several geographical races or sub-species, which differ from each other in the most trifling respects....

Selection — Let us now briefly consider the steps by which domestic races have been produced, either from one or from several allied species. Some little effect may, perhaps, be attributed to the direct action of the external conditions of life, and some little to habit; but he would be a bold man who would account by such agencies for the differences of a dray and race horse, a greyhound and bloodhound, a carrier and tumbler pigeon. One of the most remarkable features in our domesticated races is that we see in them adaptation, not indeed to the animal’s or plant’s own good, but to man’s use or fancy. Some variations useful to him have probably arisen suddenly, or by one step; many botanists, for instance, believe that the fuller’s teazle, with its hooks, which cannot be rivalled by any mechanical contrivance, is only a variety of the wild Dipsacus; and this amount of change may have suddenly arisen in a seedling.... But when we compare the dray-horse and race-horse, the dromedary and camel, the various breeds of sheep fitted either for cultivated land or mountain pasture, with the wool of one breed good for one purpose, and that of another breed for another purpose; when we compare the many breeds of dogs, each good for man in very different ways; when we compare the game-cock, so pertinacious in battle, with other breeds so little quarrelsome, with “everlasting layers” which never desire to sit, and with the bantam so small and elegant; when we compare the host of agricultural, culinary, orchard, and flower-garden races of plants, most useful to man at different seasons and for different purposes, or so beautiful in his eyes, we must, I think, look further than to mere variability. We cannot suppose that all the breeds were suddenly produced as perfect and as useful as we now see them; indeed, in several cases, we know that this has not been their history. The key is man’s power of accumulative selection: nature gives successive variations; man adds them up in certain directions useful to him. In this sense he may be said to make for himself useful breeds.

The great power of this principle of selection is not hypothetical. It is certain that several of our eminent breeders have, even within a single lifetime, modified to a large extent some breeds of cattle and sheep.... Breeders habitually speak of an animal’s organisation as something quite plastic, which they can model almost as they please.... That most skilful breeder, Sir John Sebright, used to say, with respect to pigeons, that “he would produce any given feather in three years, but it would take him six years to obtain head and beak.” In Saxony the importance of the principle of selection in regard to merino sheep is so fully recognised, that men follow it as a trade: the sheep are placed on a table and are studied, like a picture by a connoisseur; this is done three times at intervals of months, and the sheep are each time marked and classed, so that the very best may ultimately be selected for breeding.

What English breeders have actually effected is proved by the enormous prices given for animals with a good pedigree; and these have now been exported to almost every quarter of the world. The improvement is by no means generally due to crossing different breeds; all the best breeders are strongly opposed to this practice, except sometimes amongst closely allied sub-breeds. And when a cross has been made, the closest selection is far more indispensable even than in ordinary cases. If selection consisted merely in separating some very distinct variety, and breeding from it, the principle would be so obvious as hardly to be worth notice; but its importance consists in the great effect produced by the accumulation in one direction, during successive generations, of differences absolutely inappreciable by an uneducated eye — differences which I for one have vainly attempted to appreciate. Not one man in a thousand has accuracy of eye and judgment sufficient to become an eminent breeder. If gifted with these qualities, and he studies his subject for years, and devotes his lifetime to it with indomitable perseverance, he will succeed, and may make great improvements; if he wants any of these qualities, he will assuredly fail. Few would readily believe in the natural capacity and years of practice requisite to become even a skilful pigeon-fancier.

The same principles are followed by horticulturists; but the variations are here often more abrupt. No one supposes that our choicest productions have been produced by a single variation from the aboriginal stock. We have proofs that this is not so in some cases, in which exact records have been kept; thus, to give a very trifling instance, the steadily-increasing size of the common gooseberry may be quoted. We see an astonishing improvement in many florists’ flowers, when the flowers of the present day are compared with drawings made only twenty or thirty years ago. When a race of plants is once pretty well established, the seed-raisers do not pick out the best plants, but merely go over their seed-beds, and pull up the “rogues,” as they call the plants that deviate from the proper standard. With animals this kind of selection is, in fact, also followed; for hardly any one is so careless as to allow his worst animals to breed.

In regard to plants, there is another means of observing the accumulated effects of selection — namely, by comparing the diversity of flowers in the different varieties of the same species in the flower-garden; the diversity of leaves, pods, or tubers, or whatever part is valued, in the kitchen-garden, in comparison with the flowers of the same varieties; and the diversity of fruit of the same species in the orchard, in comparison with the leaves and flowers of the same set of varieties. See how different the leaves of the cabbage are, and how extremely alike the flowers; how unlike the flowers of the heartsease are, and how alike the leaves; how much the fruit of the different kinds of gooseberries differ in size, colour, shape, and hairiness, and yet the flowers present very slight differences. It is not that the varieties which differ largely in some one point do not differ at all in other points; this is hardly ever, perhaps never, the case. The laws of correlation of growth, the importance of which should never be overlooked, will ensure some differences; but, as a general rule, I cannot doubt that the continued selection of slight variations, either in the leaves, the flowers, or the fruit, will produce races differing from each other chiefly in these characters.

It may be objected that the principle of selection has been reduced to methodical practice for scarcely more than three-quarters of a century; ... But it is very far from true that the principle is a modern discovery. .... In rude and barbarous periods of English history choice animals were often imported, and laws were passed to prevent their exportation: the destruction of horses under a certain size was ordered, and this may be compared to the “roguing” of plants by nurserymen. The principle of selection I find distinctly given in an ancient Chinese encyclopaedia. Explicit rules are laid down by some of the Roman classical writers. From passages in Genesis, it is clear that the colour of domestic animals was at that early period attended to. Savages now sometimes cross their dogs with wild canine animals, to improve the breed, and they formerly did so, as is attested by passages in Pliny. .... Livingstone shows how much good domestic breeds are valued by the negroes of the interior of Africa who have not associated with Europeans. Some of these facts do not show actual selection, but they show that the breeding of domestic animals was carefully attended to in ancient times, and is now attended to by the lowest savages. It would, indeed, have been a strange fact, had attention not been paid to breeding, for the inheritance of good and bad qualities is so obvious.

At the present time, eminent breeders try by methodical selection, with a distinct object in view, to make a new strain or sub-breed, superior to anything existing in the country. But, for our purpose, a kind of Selection, which may be called Unconscious, and which results from every one trying to possess and breed from the best individual animals, is more important. Thus, a man who intends keeping pointers naturally tries to get as good dogs as he can, and afterwards breeds from his own best dogs, but he has no wish or expectation of permanently altering the breed. Nevertheless I cannot doubt that this process, continued during centuries, would improve and modify any breed ... . Slow and insensible changes of this kind could never be recognised unless actual measurements or careful drawings of the breeds in question had been made long ago, which might serve for comparison. In some cases, however, unchanged or but little changed individuals of the same breed may be found in less civilised districts, where the breed has been less improved....

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


If there exist savages so barbarous as never to think of the inherited character of the offspring of their domestic animals, yet any one animal particularly useful to them, for any special purpose, would be carefully preserved during famines and other accidents, to which savages are so liable, and such choice animals would thus generally leave more offspring than the inferior ones; so that in this case there would be a kind of unconscious selection going on. We see the value set on animals even by the barbarians of Tierra del Fuego, by their killing and devouring their old women, in times of dearth, as of less value than their dogs.

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

A large amount of change in our cultivated plants, thus slowly and unconsciously accumulated, explains, as I believe, the well-known fact, that in a vast number of cases we cannot recognise, and therefore do not know, the wild parent-stocks of the plants which have been longest cultivated in our flower and kitchen gardens. If it has taken centuries or thousands of years to improve or modify most of our plants up to their present standard of usefulness to man, we can understand how it is that neither Australia, the Cape of Good Hope, nor any other region inhabited by quite uncivilised man, has afforded us a single plant worth culture. It is not that these countries, so rich in species, do not by a strange chance possess the aboriginal stocks of any useful plants, but that the native plants have not been improved by continued selection up to a standard of perfection comparable with that given to the plants in countries anciently civilised.


On the view here given of the all-important part which selection by man has played, it becomes at once obvious, how it is that our domestic races show adaptation in their structure or in their habits to man’s wants or fancies. We can, I think, further understand the frequently abnormal character of our domestic races, and likewise their differences being so great in external characters and relatively so slight in internal parts or organs. Man can hardly select, or only with much difficulty, any deviation of structure excepting such as is externally visible; and indeed he rarely cares for what is internal. He can never act by selection, except on variations which are first given to him in some slight degree by nature. No man would ever try to make a fantail, till he saw a pigeon with a tail developed in some slight degree in an unusual manner, or a pouter till he saw a pigeon with a crop of somewhat unusual size; and the more abnormal or unusual any character was when it first appeared, the more likely it would be to catch his attention. But to use such an expression as trying to make a fantail, is, I have no doubt, in most cases, utterly incorrect. The man who first selected a pigeon with a slightly larger tail, never dreamed what the descendants of that pigeon would become through long-continued, partly unconscious and partly methodical selection. ....

Nor let it be thought that some great deviation of structure would be necessary to catch the fancier’s eye: he perceives extremely small differences, and it is in human nature to value any novelty, however slight, in one’s own possession....

I think these views further explain what has sometimes been noticed — namely that we know nothing about the origin or history of any of our domestic breeds. But, in fact, a breed, like a dialect of a language, can hardly be said to have had a definite origin. A man preserves and breeds from an individual with some slight deviation of structure, or takes more care than usual in matching his best animals and thus improves them, and the improved individuals slowly spread in the immediate neighbourhood. But as yet they will hardly have a distinct name, and from being only slightly valued, their history will be disregarded. When further improved by the same slow and gradual process, they will spread more widely, and will get recognised as something distinct and valuable, and will then probably first receive a provincial name. In semi-civilised countries, with little free communication, the spreading and knowledge of any new sub-breed will be a slow process. As soon as the points of value of the new sub-breed are once fully acknowledged, the principle, as I have called it, of unconscious selection will always tend ... slowly to add to the characteristic features of the breed, whatever they may be. But the chance will be infinitely small of any record having been preserved of such slow, varying, and insensible changes.


To sum up on the origin of our Domestic Races of animals and plants. I believe that the conditions of life, from their action on the reproductive system, are so far of the highest importance as causing variability. I do not believe that variability is an inherent and necessary contingency, under all circumstances, with all organic beings, as some authors have thought. The effects of variability are modified by various degrees of inheritance and of reversion. Variability is governed by many unknown laws, more especially by that of correlation of growth. Something may be attributed to the direct action of the conditions of life. Something must be attributed to use and disuse. The final result is thus rendered infinitely complex. In some cases, I do not doubt that the intercrossing of species, aboriginally distinct, has played an important part in the origin of our domestic productions. When in any country several domestic breeds have once been established, their occasional intercrossing, with the aid of selection, has, no doubt, largely aided in the formation of new sub-breeds; but the importance of the crossing of varieties has, I believe, been greatly exaggerated, both in regard to animals and to those plants which are propagated by seed. In plants which are temporarily propagated by cuttings, buds, etc., the importance of the crossing both of distinct species and of varieties is immense; for the cultivator here quite disregards the extreme variability both of hybrids and mongrels, and the frequent sterility of hybrids; but the cases of plants not propagated by seed are of little importance to us, for their endurance is only temporary. Over all these causes of Change I am convinced that the accumulative action of Selection, whether applied methodically and more quickly, or unconsciously and more slowly, but more efficiently, is by far the predominant Power.


Variability. Individual differences. Doubtful species. Wide ranging, much diffused, and common species vary most. Species of the larger genera in any country vary more than the species of the smaller genera. Many of the species of the larger genera resemble varieties in being very closely, but unequally, related to each other, and in having restricted ranges.

Before applying the principles arrived at in the last chapter to organic beings in a state of nature, we must briefly discuss whether these latter are subject to any variation. To treat this subject at all properly, a long catalogue of dry facts should be given; but these I shall reserve for my future work. Nor shall I here discuss the various definitions which have been given of the term species. No one definition has as yet satisfied all naturalists; yet every naturalist knows vaguely what he means when he speaks of a species. Generally the term includes the unknown element of a distinct act of creation. The term “variety” is almost equally difficult to define; but here community of descent is almost universally implied, though it can rarely be proved. We have also what are called monstrosities; but they graduate into varieties. By a monstrosity I presume is meant some considerable deviation of structure in one part, either injurious to or not useful to the species, and not generally propagated. Some authors use the term “variation” in a technical sense, as implying a modification directly due to the physical conditions of life; and “variations” in this sense are supposed not to be inherited: but who can say that the dwarfed condition of shells in the brackish waters of the Baltic, or dwarfed plants on Alpine summits, or the thicker fur of an animal from far northwards, would not in some cases be inherited for at least some few generations? and in this case I presume that the form would be called a variety.

Again, we have many slight differences which may be called individual differences, such as are known frequently to appear in the offspring from the same parents, or which may be presumed to have thus arisen, from being frequently observed in the individuals of the same species inhabiting the same confined locality. No one supposes that all the individuals of the same species are cast in the very same mould. These individual differences are highly important for us, as they afford materials for natural selection to accumulate, in the same manner as man can accumulate in any given direction individual differences in his domesticated productions. These individual differences generally affect what naturalists consider unimportant parts; but I could show by a long catalogue of facts, that parts which must be called important, whether viewed under a physiological or classificatory point of view, sometimes vary in the individuals of the same species.... I should never have expected that the branching of the main nerves close to the great central ganglion of an insect would have been variable in the same species; I should have expected that changes of this nature could have been effected only by slow degrees: yet quite recently Mr. Lubbock has shown a degree of variability in these main nerves in Coccus, which may almost be compared to the irregular branching of the stem of a tree. This philosophical naturalist, I may add, has also quite recently shown that the muscles in the larvae of certain insects are very far from uniform. Authors sometimes argue in a circle when they state that important organs never vary; for these same authors practically rank that character as important (as some few naturalists have honestly confessed) which does not vary; and, under this point of view, no instance of an important part varying will ever be found: but under any other point of view many instances assuredly can be given.

There is one point connected with individual differences, which seems to me extremely perplexing: I refer to those genera which have sometimes been called “protean” or “polymorphic,” in which the species present an inordinate amount of variation; and hardly two naturalists can agree which forms to rank as species and which as varieties. We may instance Rubus, Rosa, and Hieracium amongst plants, several genera of insects, and several genera of Brachiopod shells. In most polymorphic genera some of the species have fixed and definite characters. Genera which are polymorphic in one country seem to be, with some few exceptions, polymorphic in other countries, and likewise, judging from Brachiopod shells, at former periods of time. ....

Those forms which possess in some considerable degree the character of species, but which are so closely similar to some other forms, or are so closely linked to them by intermediate gradations, that naturalists do not like to rank them as distinct species, are in several respects the most important for us. We have every reason to believe that many of these doubtful and closely-allied forms have permanently retained their characters in their own country for a long time; for as long, as far as we know, as have good and true species. Practically, when a naturalist can unite two forms together by others having intermediate characters, he treats the one as a variety of the other, ranking the most common, but sometimes the one first described, as the species, and the other as the variety. But cases of great difficulty, which I will not here enumerate, sometimes occur in deciding whether or not to rank one form as a variety of another, even when they are closely connected by intermediate links; nor will the commonly-assumed hybrid nature of the intermediate links always remove the difficulty. In very many cases, however, one form is ranked as a variety of another, not because the intermediate links have actually been found, but because analogy leads the observer to suppose either that they do now somewhere exist, or may formerly have existed; and here a wide door for the entry of doubt and conjecture is opened.

Hence, in determining whether a form should be ranked as a species or a variety, the opinion of naturalists having sound judgment and wide experience seems the only guide to follow. We must, however, in many cases, decide by a majority of naturalists, for few well-marked and well-known varieties can be named which have not been ranked as species by at least some competent judges.

That varieties of this doubtful nature are far from uncommon cannot be disputed... Mr. H. C. Watson, to whom I lie under deep obligation for assistance of all kinds, has marked for me 182 British plants, which are generally considered as varieties, but which have all been ranked by botanists as species; and in making this list he has omitted many trifling varieties, but which nevertheless have been ranked by some botanists as species, and he has entirely omitted several highly polymorphic genera. Under genera, including the most polymorphic forms, Mr. Babington gives 251 species, whereas Mr. Bentham gives only 112, — a difference of 139 doubtful forms! Amongst animals which unite for each birth, and which are highly locomotive, doubtful forms, ranked by one zoologist as a species and by another as a variety, can rarely be found within the same country, but are common in separated areas. How many of those birds and insects in North America and Europe, which differ very slightly from each other, have been ranked by one eminent naturalist as undoubted species, and by another as varieties, or, as they are often called, as geographical races! Many years ago, when comparing, and seeing others compare, the birds from the separate islands of the Galapagos Archipelago, both one with another, and with those from the American mainland, I was much struck how entirely vague and arbitrary is the distinction between species and varieties. On the islets of the little Madeira group there are many insects which are characterized as varieties in Mr. Wollaston’s admirable work, but which it cannot be doubted would be ranked as distinct species by many entomologists.... A wide distance between the homes of two doubtful forms leads many naturalists to rank both as distinct species; but what distance, it has been well asked, will suffice? if that between America and Europe is ample, will that between the Continent and the Azores, or Madeira, or the Canaries, or Ireland, be sufficient? It must be admitted that many forms, considered by highly-competent judges as varieties, have so perfectly the character of species that they are ranked by other highly-competent judges as good and true species. But to discuss whether they are rightly called species or varieties, before any definition of these terms has been generally accepted, is vainly to beat the air.

Many of the cases of strongly-marked varieties or doubtful species well deserve consideration; for several interesting lines of argument, from geographical distribution, analogical variation, hybridism, etc., have been brought to bear on the attempt to determine their rank. I will here give only a single instance, — the well-known one of the primrose and cowslip, or Primula veris and elatior. These plants differ considerably in appearance; they have a different flavour and emit a different odour; they flower at slightly different periods; they grow in somewhat different stations; they ascend mountains to different heights; they have different geographical ranges; and lastly, according to very numerous experiments made during several years by that most careful observer Gartner, they can be crossed only with much difficulty. We could hardly wish for better evidence of the two forms being specifically distinct. On the other hand, they are united by many intermediate links, and it is very doubtful whether these links are hybrids; and there is, as it seems to me, an overwhelming amount of experimental evidence, showing that they descend from common parents, and consequently must be ranked as varieties.

Close investigation, in most cases, will bring naturalists to an agreement how to rank doubtful forms. Yet it must be confessed, that it is in the best-known countries that we find the greatest number of forms of doubtful value. I have been struck with the fact, that if any animal or plant in a state of nature be highly useful to man, or from any cause closely attract his attention, varieties of it will almost universally be found recorded. These varieties, moreover, will be often ranked by some authors as species. Look at the common oak, how closely it has been studied; yet a German author makes more than a dozen species out of forms, which are very generally considered as varieties; and in this country the highest botanical authorities and practical men can be quoted to show that the sessile and pedunculated oaks are either good and distinct species or mere varieties.

When a young naturalist commences the study of a group of organisms quite unknown to him, he is at first much perplexed to determine what differences to consider as specific, and what as varieties; for he knows nothing of the amount and kind of variation to which the group is subject; and this shows, at least, how very generally there is some variation. But if he confine his attention to one class within one country, he will soon make up his mind how to rank most of the doubtful forms. His general tendency will be to make many species, for he will become impressed, ... with the amount of difference in the forms which he is continually studying; and he has little general knowledge of analogical variation in other groups and in other countries, by which to correct his first impressions. As he extends the range of his observations, he will meet with more cases of difficulty; for he will encounter a greater number of closely-allied forms. But if his observations be widely extended, he will in the end generally be enabled to make up his own mind which to call varieties and which species; but he will succeed in this at the expense of admitting much variation, — and the truth of this admission will often be disputed by other naturalists. When, moreover, he comes to study allied forms brought from countries not now continuous, in which case he can hardly hope to find the intermediate links between his doubtful forms, he will have to trust almost entirely to analogy, and his difficulties will rise to a climax.

Certainly no clear line of demarcation has as yet been drawn between species and sub-species — that is, the forms which in the opinion of some naturalists come very near to, but do not quite arrive at the rank of species; or, again, between sub-species and well-marked varieties, or between lesser varieties and individual differences. These differences blend into each other in an insensible series; and a series impresses the mind with the idea of an actual passage.

Hence I look at individual differences, though of small interest to the systematist, as of high importance for us, as being the first step towards such slight varieties as are barely thought worth recording in works on natural history. And I look at varieties which are in any degree more distinct and permanent, as steps leading to more strongly marked and more permanent varieties; and at these latter, as leading to sub-species, and to species. The passage from one stage of difference to another and higher stage may be, in some cases, due merely to the long-continued action of different physical conditions in two different regions; but I have not much faith in this view; and I attribute the passage of a variety, from a state in which it differs very slightly from its parent to one in which it differs more, to the action of natural selection in accumulating (as will hereafter be more fully explained) differences of structure in certain definite directions. Hence I believe a well-marked variety may be justly called an incipient species; but whether this belief be justifiable must be judged of by the general weight of the several facts and views given throughout this work.


From these remarks it will be seen that I look at the term species, as one arbitrarily given for the sake of convenience to a set of individuals closely resembling each other, and that it does not essentially differ from the term variety, which is given to less distinct and more fluctuating forms. The term variety, again, in comparison with mere individual differences, is also applied arbitrarily, and for mere convenience sake.

.... The whole subject, however, treated as it necessarily here is with much brevity, is rather perplexing, and allusions cannot be avoided to the “struggle for existence,” “divergence of character,” and other questions, hereafter to be discussed.

Alph. De Candolle and others have shown that plants which have very wide ranges generally present varieties; and this might have been expected, as they become exposed to diverse physical conditions, and as they come into competition (which, as we shall hereafter see, is a far more important circumstance) with different sets of organic beings. But my tables further show that, in any limited country, the species which are most common, that is abound most in individuals, and the species which are most widely diffused within their own country (and this is a different consideration from wide range, and to a certain extent from commonness), often give rise to varieties sufficiently well-marked to have been recorded in botanical works. Hence it is the most flourishing, or, as they may be called, the dominant species, — those which range widely over the world, are the most diffused in their own country, and are the most numerous in individuals, — which oftenest produce well-marked varieties, or, as I consider them, incipient species. And this, perhaps, might have been anticipated; for, as varieties, in order to become in any degree permanent, necessarily have to struggle with the other inhabitants of the country, the species which are already dominant will be the most likely to yield offspring which, though in some slight degree modified, will still inherit those advantages that enabled their parents to become dominant over their compatriots.

.... Moreover, the species of the large genera are related to each other, in the same manner as the varieties of any one species are related to each other. No naturalist pretends that all the species of a genus are equally distinct from each other; they may generally be divided into sub-genera, or sections, or lesser groups. As Fries has well remarked, little groups of species are generally clustered like satellites around certain other species. And what are varieties but groups of forms, unequally related to each other, and clustered round certain forms — that is, round their parent-species? Undoubtedly there is one most important point of difference between varieties and species; namely, that the amount of difference between varieties, when compared with each other or with their parent-species, is much less than that between the species of the same genus. But when we come to discuss the principle, as I call it, of Divergence of Character, we shall see how this may be explained, and how the lesser differences between varieties will tend to increase into the greater differences between species.

There is one other point which seems to me worth notice. Varieties generally have much restricted ranges: this statement is indeed scarcely more than a truism, for if a variety were found to have a wider range than that of its supposed parent-species, their denominations ought to be reversed. But there is also reason to believe, that those species which are very closely allied to other species, and in so far resemble varieties, often have much restricted ranges. For instance, Mr. H. C. Watson has marked for me in the well-sifted London Catalogue of plants (4th edition) 63 plants which are therein ranked as species, but which he considers as so closely allied to other species as to be of doubtful value: these 63 reputed species range on an average over 6.9 of the provinces into which Mr. Watson has divided Great Britain. Now, in this same catalogue, 53 acknowledged varieties are recorded, and these range over 7.7 provinces; whereas, the species to which these varieties belong range over 14.3 provinces. So that the acknowledged varieties have very nearly the same restricted average range, as have those very closely allied forms, marked for me by Mr. Watson as doubtful species, but which are almost universally ranked by British botanists as good and true species.

Finally, then, varieties have the same general characters as species, for they cannot be distinguished from species, — except, firstly, by the discovery of intermediate linking forms, and the occurrence of such links cannot affect the actual characters of the forms which they connect; and except, secondly, by a certain amount of difference, for two forms, if differing very little, are generally ranked as varieties, notwithstanding that intermediate linking forms have not been discovered; but the amount of difference considered necessary to give to two forms the rank of species is quite indefinite. In genera having more than the average number of species in any country, the species of these genera have more than the average number of varieties. In large genera the species are apt to be closely, but unequally, allied together, forming little clusters round certain species. Species very closely allied to other species apparently have restricted ranges. In all these several respects the species of large genera present a strong analogy with varieties. And we can clearly understand these analogies, if species have once existed as varieties, and have thus originated: whereas, these analogies are utterly inexplicable if each species has been independently created.

We have, also, seen that it is the most flourishing and dominant species of the larger genera which on an average vary most; and varieties, as we shall hereafter see, tend to become converted into new and distinct species. The larger genera thus tend to become larger; and throughout nature the forms of life which are now dominant tend to become still more dominant by leaving many modified and dominant descendants. But by steps hereafter to be explained, the larger genera also tend to break up into smaller genera. And thus, the forms of life throughout the universe become divided into groups subordinate to groups.


Bears on natural selection. The term used in a wide sense. Geometrical powers of increase. Rapid increase of naturalised animals and plants. Nature of the checks to increase. Competition universal. Effects of climate. Protection from the number of individuals. Complex relations of all animals and plants throughout nature. Struggle for life most severe between individuals and varieties of the same species; often severe between species of the same genus. The relation of organism to organism the most important of all relations.

Before entering on the subject of this chapter, I must make a few preliminary remarks, to show how the struggle for existence bears on Natural Selection. It has been seen in the last chapter that amongst organic beings in a state of nature there is some individual variability; indeed I am not aware that this has ever been disputed. It is immaterial for us whether a multitude of doubtful forms be called species or sub-species or varieties; what rank, for instance, the two or three hundred doubtful forms of British plants are entitled to hold, if the existence of any well-marked varieties be admitted. But the mere existence of individual variability and of some few well-marked varieties, though necessary as the foundation for the work, helps us but little in understanding how species arise in nature. How have all those exquisite adaptations of one part of the organisation to another part, and to the conditions of life, and of one distinct organic being to another being, been perfected? We see these beautiful co-adaptations most plainly in the woodpecker and missletoe; and only a little less plainly in the humblest parasite which clings to the hairs of a quadruped or feathers of a bird; in the structure of the beetle which dives through the water; in the plumed seed which is wafted by the gentlest breeze; in short, we see beautiful adaptations everywhere and in every part of the organic world.

Again, it may be asked, how is it that varieties, which I have called incipient species, become ultimately converted into good and distinct species, which in most cases obviously differ from each other far more than do the varieties of the same species? How do those groups of species, which constitute what are called distinct genera, and which differ from each other more than do the species of the same genus, arise? All these results, as we shall more fully see in the next chapter, follow inevitably from the struggle for life. Owing to this struggle for life, any variation, however slight and from whatever cause proceeding, if it be in any degree profitable to an individual of any species, in its infinitely complex relations to other organic beings and to external nature, will tend to the preservation of that individual, and will generally be inherited by its offspring. The offspring, also, will thus have a better chance of surviving, for, of the many individuals of any species which are periodically born, but a small number can survive. I have called this principle, by which each slight variation, if useful, is preserved, by the term of Natural Selection, in order to mark its relation to man’s power of selection. We have seen that man by selection can certainly produce great results, and can adapt organic beings to his own uses, through the accumulation of slight but useful variations, given to him by the hand of Nature. But Natural Selection, as we shall hereafter see, is a power incessantly ready for action, and is as immeasurably superior to man’s feeble efforts, as the works of Nature are to those of Art.

.... Nothing is easier than to admit in words the truth of the universal struggle for life, or more difficult — at least I have found it so — than constantly to bear this conclusion in mind. Yet unless it be thoroughly engrained in the mind, I am convinced that the whole economy of nature, with every fact on distribution, rarity, abundance, extinction, and variation, will be dimly seen or quite misunderstood. We behold the face of nature bright with gladness, we often see superabundance of food; we do not see, or we forget, that the birds which are idly singing round us mostly live on insects or seeds, and are thus constantly destroying life; or we forget how largely these songsters, or their eggs, or their nestlings, are destroyed by birds and beasts of prey; we do not always bear in mind, that though food may be now superabundant, it is not so at all seasons of each recurring year.

I should premise that I use the term Struggle for Existence in a large and metaphorical sense, including dependence of one being on another, and including (which is more important) not only the life of the individual, but success in leaving progeny. Two canine animals in a time of dearth, may be truly said to struggle with each other which shall get food and live. But a plant on the edge of a desert is said to struggle for life against the drought, though more properly it should be said to be dependent on the moisture. A plant which annually produces a thousand seeds, of which on an average only one comes to maturity, may be more truly said to struggle with the plants of the same and other kinds which already clothe the ground. The mistletoe is dependent on the apple and a few other trees, but can only in a far-fetched sense be said to struggle with these trees, for if too many of these parasites grow on the same tree, it will languish and die. But several seedling mistletoes, growing close together on the same branch, may more truly be said to struggle with each other. As the mistletoe is disseminated by birds, its existence depends on birds; and it may metaphorically be said to struggle with other fruit-bearing plants, in order to tempt birds to devour and thus disseminate its seeds rather than those of other plants. In these several senses, which pass into each other, I use for convenience sake the general term of struggle for existence.

A struggle for existence inevitably follows from the high rate at which all organic beings tend to increase. Every being, which during its natural lifetime produces several eggs or seeds, must suffer destruction during some period of its life, and during some season or occasional year, otherwise, on the principle of geometrical increase, its numbers would quickly become so inordinately great that no country could support the product. Hence, as more individuals are produced than can possibly survive, there must in every case be a struggle for existence, either one individual with another of the same species, or with the individuals of distinct species, or with the physical conditions of life. It is the doctrine of Malthus applied with manifold force to the whole animal and vegetable kingdoms; for in this case there can be no artificial increase of food, and no prudential restraint from marriage. Although some species may be now increasing, more or less rapidly, in numbers, all cannot do so, for the world would not hold them.

There is no exception to the rule that every organic being naturally increases at so high a rate, that if not destroyed, the earth would soon be covered by the progeny of a single pair. Even slow-breeding man has doubled in twenty-five years, and at this rate, in a few thousand years, there would literally not be standing room for his progeny. Linnaeus has calculated that if an annual plant produced only two seeds — and there is no plant so unproductive as this — and their seedlings next year produced two, and so on, then in twenty years there would be a million plants. The elephant is reckoned to be the slowest breeder of all known animals, and I have taken some pains to estimate its probable minimum rate of natural increase: it will be under the mark to assume that it breeds when thirty years old, and goes on breeding till ninety years old, bringing forth three pair of young in this interval; if this be so, at the end of the fifth century there would be alive fifteen million elephants, descended from the first pair.

But we have better evidence on this subject than mere theoretical calculations, namely, the numerous recorded cases of the astonishingly rapid increase of various animals in a state of nature, when circumstances have been favourable to them during two or three following seasons. Still more striking is the evidence from our domestic animals of many kinds which have run wild in several parts of the world: if the statements of the rate of increase of slow-breeding cattle and horses in South America, and latterly in Australia, had not been well authenticated, they would have been quite incredible. So it is with plants: cases could be given of introduced plants which have become common throughout whole islands in a period of less than ten years. Several of the plants now most numerous over the wide plains of La Plata, clothing square leagues of surface almost to the exclusion of all other plants, have been introduced from Europe; and there are plants which now range in India, as I hear from Dr. Falconer, from Cape Comorin to the Himalaya, which have been imported from America since its discovery. In such cases, and endless instances could be given, no one supposes that the fertility of these animals or plants has been suddenly and temporarily increased in any sensible degree. The obvious explanation is that the conditions of life have been very favourable, and that there has consequently been less destruction of the old and young, and that nearly all the young have been enabled to breed. In such cases the geometrical ratio of increase, the result of which never fails to be surprising, simply explains the extraordinarily rapid increase and wide diffusion of naturalised productions in their new homes.

In a state of nature almost every plant produces seed, and amongst animals there are very few which do not annually pair. Hence we may confidently assert, that all plants and animals are tending to increase at a geometrical ratio, that all would most rapidly stock every station in which they could any how exist, and that the geometrical tendency to increase must be checked by destruction at some period of life. Our familiarity with the larger domestic animals tends, I think, to mislead us: we see no great destruction falling on them, and we forget that thousands are annually slaughtered for food, and that in a state of nature an equal number would have somehow to be disposed of.

The only difference between organisms which annually produce eggs or seeds by the thousand, and those which produce extremely few, is, that the slow-breeders would require a few more years to people, under favourable conditions, a whole district, let it be ever so large. The condor lays a couple of eggs and the ostrich a score, and yet in the same country the condor may be the more numerous of the two: the Fulmar petrel lays but one egg, yet it is believed to be the most numerous bird in the world. One fly deposits hundreds of eggs, and another, like the hippobosca, a single one; but this difference does not determine how many individuals of the two species can be supported in a district. A large number of eggs is of some importance to those species, which depend on a rapidly fluctuating amount of food, for it allows them rapidly to increase in number. But the real importance of a large number of eggs or seeds is to make up for much destruction at some period of life; and this period in the great majority of cases is an early one. If an animal can in any way protect its own eggs or young, a small number may be produced, and yet the average stock be fully kept up; but if many eggs or young are destroyed, many must be produced, or the species will become extinct. It would suffice to keep up the full number of a tree, which lived on an average for a thousand years, if a single seed were produced once in a thousand years, supposing that this seed were never destroyed, and could be ensured to germinate in a fitting place. So that in all cases, the average number of any animal or plant depends only indirectly on the number of its eggs or seeds.

In looking at Nature, it is most necessary to keep the foregoing considerations always in mind — never to forget that every single organic being around us may be said to be striving to the utmost to increase in numbers; that each lives by a struggle at some period of its life; that heavy destruction inevitably falls either on the young or old, during each generation or at recurrent intervals. Lighten any check, mitigate the destruction ever so little, and the number of the species will almost instantaneously increase to any amount. The face of Nature may be compared to a yielding surface, with ten thousand sharp wedges packed close together and driven inwards by incessant blows, sometimes one wedge being struck, and then another with greater force.

What checks the natural tendency of each species to increase in number is most obscure. Look at the most vigorous species; by as much as it swarms in numbers, by so much will its tendency to increase be still further increased. We know not exactly what the checks are in even one single instance. Nor will this surprise any one who reflects how ignorant we are on this head, even in regard to mankind, so incomparably better known than any other animal.... Here I will make only a few remarks, just to recall to the reader’s mind some of the chief points. Eggs or very young animals seem generally to suffer most, but this is not invariably the case. With plants there is a vast destruction of seeds, but, from some observations which I have made, I believe that it is the seedlings which suffer most from germinating in ground already thickly stocked with other plants. Seedlings, also, are destroyed in vast numbers by various enemies; for instance, on a piece of ground three feet long and two wide, dug and cleared, and where there could be no choking from other plants, I marked all the seedlings of our native weeds as they came up, and out of the 357 no less than 295 were destroyed, chiefly by slugs and insects. If turf which has long been mown, and the case would be the same with turf closely browsed by quadrupeds, be let to grow, the more vigorous plants gradually kill the less vigorous, though fully grown, plants: thus out of twenty species growing on a little plot of turf (three feet by four) nine species perished from the other species being allowed to grow up freely.

The amount of food for each species of course gives the extreme limit to which each can increase; but very frequently it is not the obtaining food, but the serving as prey to other animals, which determines the average numbers of a species. Thus, there seems to be little doubt that the stock of partridges, grouse, and hares on any large estate depends chiefly on the destruction of vermin. If not one head of game were shot during the next twenty years in England, and, at the same time, if no vermin were destroyed, there would, in all probability, be less game than at present, although hundreds of thousands of game animals are now annually killed. On the other hand, in some cases, as with the elephant and rhinoceros, none are destroyed by beasts of prey: even the tiger in India most rarely dares to attack a young elephant protected by its dam.

Climate plays an important part in determining the average numbers of a species, and periodical seasons of extreme cold or drought, I believe to be the most effective of all checks. I estimated that the winter of 1854-55 destroyed four-fifths of the birds in my own grounds; and this is a tremendous destruction, when we remember that ten per cent. is an extraordinarily severe mortality from epidemics with man. The action of climate seems at first sight to be quite independent of the struggle for existence; but in so far as climate chiefly acts in reducing food, it brings on the most severe struggle between the individuals, whether of the same or of distinct species, which subsist on the same kind of food. Even when climate, for instance extreme cold, acts directly, it will be the least vigorous, or those which have got least food through the advancing winter, which will suffer most. When we travel from south to north, or from a damp region to a dry, we invariably see some species gradually getting rarer and rarer, and finally disappearing; and the change of climate being conspicuous, we are tempted to attribute the whole effect to its direct action. But this is a very false view: we forget that each species, even where it most abounds, is constantly suffering enormous destruction at some period of its life, from enemies or from competitors for the same place and food; and if these enemies or competitors be in the least degree favoured by any slight change of climate, they will increase in numbers, and, as each area is already fully stocked with inhabitants, the other species will decrease. When we travel southward and see a species decreasing in numbers, we may feel sure that the cause lies quite as much in other species being favoured, as in this one being hurt. So it is when we travel northward, but in a somewhat lesser degree, for the number of species of all kinds, and therefore of competitors, decreases northwards; hence in going northward, or in ascending a mountain, we far oftener meet with stunted forms, due to the directly injurious action of climate, than we do in proceeding southwards or in descending a mountain. When we reach the Arctic regions, or snow-capped summits, or absolute deserts, the struggle for life is almost exclusively with the elements.

That climate acts in main part indirectly by favouring other species, we may clearly see in the prodigious number of plants in our gardens which can perfectly well endure our climate, but which never become naturalised, for they cannot compete with our native plants, nor resist destruction by our native animals.


Many cases are on record showing how complex and unexpected are the checks and relations between organic beings, which have to struggle together in the same country. I will give only a single instance, which, though a simple one, has interested me. In Staffordshire, on the estate of a relation where I had ample means of investigation, there was a large and extremely barren heath, which had never been touched by the hand of man; but several hundred acres of exactly the same nature had been enclosed twenty-five years previously and planted with Scotch fir. The change in the native vegetation of the planted part of the heath was most remarkable, more than is generally seen in passing from one quite different soil to another: not only the proportional numbers of the heath-plants were wholly changed, but twelve species of plants (not counting grasses and carices) flourished in the plantations, which could not be found on the heath. The effect on the insects must have been still greater, for six insectivorous birds were very common in the plantations, which were not to be seen on the heath; and the heath was frequented by two or three distinct insectivorous birds. Here we see how potent has been the effect of the introduction of a single tree, nothing whatever else having been done, with the exception that the land had been enclosed, so that cattle could not enter. But how important an element enclosure is, I plainly saw near Farnham, in Surrey. Here there are extensive heaths, with a few clumps of old Scotch firs on the distant hill-tops: within the last ten years large spaces have been enclosed, and self-sown firs are now springing up in multitudes, so close together that all cannot live.

When I ascertained that these young trees had not been sown or planted, I was so much surprised at their numbers that I went to several points of view, whence I could examine hundreds of acres of the unenclosed heath, and literally I could not see a single Scotch fir, except the old planted clumps. But on looking closely between the stems of the heath, I found a multitude of seedlings and little trees, which had been perpetually browsed down by the cattle. In one square yard, at a point some hundred yards distant from one of the old clumps, I counted thirty-two little trees; and one of them, judging from the rings of growth, had during twenty-six years tried to raise its head above the stems of the heath, and had failed. No wonder that, as soon as the land was enclosed, it became thickly clothed with vigorously growing young firs. Yet the heath was so extremely barren and so extensive that no one would ever have imagined that cattle would have so closely and effectually searched it for food.

Here we see that cattle absolutely determine the existence of the Scotch fir; but in several parts of the world insects determine the existence of cattle. Perhaps Paraguay offers the most curious instance of this; for here neither cattle nor horses nor dogs have ever run wild, though they swarm southward and northward in a feral state; and Azara and Rengger have shown that this is caused by the greater number in Paraguay of a certain fly, which lays its eggs in the navels of these animals when first born. The increase of these flies, numerous as they are, must be habitually checked by some means, probably by birds. Hence, if certain insectivorous birds (whose numbers are probably regulated by hawks or beasts of prey) were to increase in Paraguay, the flies would decrease — then cattle and horses would become feral, and this would certainly greatly alter (as indeed I have observed in parts of South America) the vegetation: this again would largely affect the insects; and this, as we just have seen in Staffordshire, the insectivorous birds, and so onwards in ever-increasing circles of complexity. We began this series by insectivorous birds, and we have ended with them. Not that in nature the relations can ever be as simple as this. Battle within battle must ever be recurring with varying success; and yet in the long-run the forces are so nicely balanced, that the face of nature remains uniform for long periods of time, though assuredly the merest trifle would often give the victory to one organic being over another. Nevertheless so profound is our ignorance, and so high our presumption, that we marvel when we hear of the extinction of an organic being; and as we do not see the cause, we invoke cataclysms to desolate the world, or invent laws on the duration of the forms of life!

I am tempted to give one more instance showing how plants and animals, most remote in the scale of nature, are bound together by a web of complex relations. I shall hereafter have occasion to show that the exotic Lobelia fulgens, in this part of England, is never visited by insects, and consequently, from its peculiar structure, never can set a seed. Many of our orchidaceous plants absolutely require the visits of moths to remove their pollen-masses and thus to fertilise them. I have, also, reason to believe that humble-bees are indispensable to the fertilisation of the heartsease (Viola tricolor), for other bees do not visit this flower. From experiments which I have tried, I have found that the visits of bees, if not indispensable, are at least highly beneficial to the fertilisation of our clovers; but humble-bees alone visit the common red clover (Trifolium pratense), as other bees cannot reach the nectar. Hence I have very little doubt, that if the whole genus of humble-bees became extinct or very rare in England, the heartsease and red clover would become very rare, or wholly disappear. The number of humble-bees in any district depends in a great degree on the number of field-mice, which destroy their combs and nests; and Mr. H. Newman, who has long attended to the habits of humble-bees, believes that “more than two thirds of them are thus destroyed all over England.” Now the number of mice is largely dependent, as every one knows, on the number of cats; and Mr. Newman says, “Near villages and small towns I have found the nests of humble-bees more numerous than elsewhere, which I attribute to the number of cats that destroy the mice.” Hence it is quite credible that the presence of a feline animal in large numbers in a district might determine, through the intervention first of mice and then of bees, the frequency of certain flowers in that district!

In the case of every species, many different checks, acting at different periods of life, and during different seasons or years, probably come into play; some one check or some few being generally the most potent, but all concurring in determining the average number or even the existence of the species. In some cases it can be shown that widely-different checks act on the same species in different districts. When we look at the plants and bushes clothing an entangled bank, we are tempted to attribute their proportional numbers and kinds to what we call chance. But how false a view is this! Every one has heard that when an American forest is cut down, a very different vegetation springs up; but it has been observed that the trees now growing on the ancient Indian mounds, in the Southern United States, display the same beautiful diversity and proportion of kinds as in the surrounding virgin forests. What a struggle between the several kinds of trees must here have gone on during long centuries, each annually scattering its seeds by the thousand; what war between insect and insect — between insects, snails, and other animals with birds and beasts of prey — all striving to increase, and all feeding on each other or on the trees or their seeds and seedlings, or on the other plants which first clothed the ground and thus checked the growth of the trees! Throw up a handful of feathers, and all must fall to the ground according to definite laws; but how simple is this problem compared to the action and reaction of the innumerable plants and animals which have determined, in the course of centuries, the proportional numbers and kinds of trees now growing on the old Indian ruins!

The dependency of one organic being on another, as of a parasite on its prey, lies generally between beings remote in the scale of nature. This is often the case with those which may strictly be said to struggle with each other for existence, as in the case of locusts and grass-feeding quadrupeds. But the struggle almost invariably will be most severe between the individuals of the same species, for they frequent the same districts, require the same food, and are exposed to the same dangers. In the case of varieties of the same species, the struggle will generally be almost equally severe, and we sometimes see the contest soon decided: for instance, if several varieties of wheat be sown together, and the mixed seed be resown, some of the varieties which best suit the soil or climate, or are naturally the most fertile, will beat the others and so yield more seed, and will consequently in a few years quite supplant the other varieties. To keep up a mixed stock of even such extremely close varieties as the variously coloured sweet-peas, they must be each year harvested separately, and the seed then mixed in due proportion, otherwise the weaker kinds will steadily decrease in numbers and disappear. So again with the varieties of sheep: it has been asserted that certain mountain-varieties will starve out other mountain-varieties, so that they cannot be kept together. .... It may even be doubted whether the varieties of any one of our domestic plants or animals have so exactly the same strength, habits, and constitution, that the original proportions of a mixed stock could be kept up for half a dozen generations, if they were allowed to struggle together, like beings in a state of nature, and if the seed or young were not annually sorted.

As species of the same genus have usually, though by no means invariably, some similarity in habits and constitution, and always in structure, the struggle will generally be more severe between species of the same genus, when they come into competition with each other, than between species of distinct genera. We see this in the recent extension over parts of the United States of one species of swallow having caused the decrease of another species. The recent increase of the missel-thrush in parts of Scotland has caused the decrease of the song-thrush. How frequently we hear of one species of rat taking the place of another species under the most different climates! .... We can dimly see why the competition should be most severe between allied forms, which fill nearly the same place in the economy of nature; but probably in no one case could we precisely say why one species has been victorious over another in the great battle of life.

A corollary of the highest importance may be deduced from the foregoing remarks, namely, that the structure of every organic being is related, in the most essential yet often hidden manner, to that of all other organic beings, with which it comes into competition for food or residence, or from which it has to escape, or on which it preys. This is obvious in the structure of the teeth and talons of the tiger; and in that of the legs and claws of the parasite which clings to the hair on the tiger’s body. But in the beautifully plumed seed of the dandelion, and in the flattened and fringed legs of the water-beetle, the relation seems at first confined to the elements of air and water. Yet the advantage of plumed seeds no doubt stands in the closest relation to the land being already thickly clothed by other plants; so that the seeds may be widely distributed and fall on unoccupied ground. In the water-beetle, the structure of its legs, so well adapted for diving, allows it to compete with other aquatic insects, to hunt for its own prey, and to escape serving as prey to other animals.

The store of nutriment laid up within the seeds of many plants seems at first sight to have no sort of relation to other plants. But from the strong growth of young plants produced from such seeds (as peas and beans), when sown in the midst of long grass, I suspect that the chief use of the nutriment in the seed is to favour the growth of the young seedling, whilst struggling with other plants growing vigorously all around.

Look at a plant in the midst of its range, why does it not double or quadruple its numbers? We know that it can perfectly well withstand a little more heat or cold, dampness or dryness, for elsewhere it ranges into slightly hotter or colder, damper or drier districts. In this case we can clearly see that if we wished in imagination to give the plant the power of increasing in number, we should have to give it some advantage over its competitors, or over the animals which preyed on it. On the confines of its geographical range, a change of constitution with respect to climate would clearly be an advantage to our plant; but we have reason to believe that only a few plants or animals range so far, that they are destroyed by the rigour of the climate alone. Not until we reach the extreme confines of life, in the arctic regions or on the borders of an utter desert, will competition cease. The land may be extremely cold or dry, yet there will be competition between some few species, or between the individuals of the same species, for the warmest or dampest spots.

Hence, also, we can see that when a plant or animal is placed in a new country amongst new competitors, though the climate may be exactly the same as in its former home, yet the conditions of its life will generally be changed in an essential manner. If we wished to increase its average numbers in its new home, we should have to modify it in a different way to what we should have done in its native country; for we should have to give it some advantage over a different set of competitors or enemies.

It is good thus to try in our imagination to give any form some advantage over another. Probably in no single instance should we know what to do, so as to succeed. It will convince us of our ignorance on the mutual relations of all organic beings; a conviction as necessary, as it seems to be difficult to acquire. All that we can do, is to keep steadily in mind that each organic being is striving to increase at a geometrical ratio; that each at some period of its life, during some season of the year, during each generation or at intervals, has to struggle for life, and to suffer great destruction. When we reflect on this struggle, we may console ourselves with the full belief, that the war of nature is not incessant, that no fear is felt, that death is generally prompt, and that the vigorous, the healthy, and the happy survive and multiply.



Natural Selection: its power compared with man’s selection, its power on characters of trifling importance, its power at all ages and on both sexes. Sexual Selection. On the generality of intercrosses between individuals of the same species. Circumstances favourable and unfavourable to Natural Selection, namely, intercrossing, isolation, number of individuals. Slow action. Extinction caused by Natural Selection. Divergence of Character, related to the diversity of inhabitants of any small area, and to naturalisation. Action of Natural Selection, through Divergence of Character and Extinction, on the descendants from a common parent. Explains the Grouping of all organic beings.

How will the struggle for existence, discussed too briefly in the last chapter, act in regard to variation? Can the principle of selection, which we have seen is so potent in the hands of man, apply in nature? I think we shall see that it can act most effectually. Let it be borne in mind in what an endless number of strange peculiarities our domestic productions, and, in a lesser degree, those under nature, vary; and how strong the hereditary tendency is. Under domestication, it may be truly said that the whole organisation becomes in some degree plastic. Let it be borne in mind how infinitely complex and close-fitting are the mutual relations of all organic beings to each other and to their physical conditions of life. Can it, then, be thought improbable, seeing that variations useful to man have undoubtedly occurred, that other variations useful in some way to each being in the great and complex battle of life, should sometimes occur in the course of thousands of generations? If such do occur, can we doubt (remembering that many more individuals are born than can possibly survive) that individuals having any advantage, however slight, over others, would have the best chance of surviving and of procreating their kind? On the other hand, we may feel sure that any variation in the least degree injurious would be rigidly destroyed. This preservation of favourable variations and the rejection of injurious variations, I call Natural Selection. Variations neither useful nor injurious would not be affected by natural selection, and would be left a fluctuating element, as perhaps we see in the species called polymorphic.

We shall best understand the probable course of natural selection by taking the case of a country undergoing some physical change, for instance, of climate. The proportional numbers of its inhabitants would almost immediately undergo a change, and some species might become extinct. We may conclude, from what we have seen of the intimate and complex manner in which the inhabitants of each country are bound together, that any change in the numerical proportions of some of the inhabitants, independently of the change of climate itself, would most seriously affect many of the others. If the country were open on its borders, new forms would certainly immigrate, and this also would seriously disturb the relations of some of the former inhabitants. Let it be remembered how powerful the influence of a single introduced tree or mammal has been shown to be. But in the case of an island, or of a country partly surrounded by barriers, into which new and better adapted forms could not freely enter, we should then have places in the economy of nature which would assuredly be better filled up, if some of the original inhabitants were in some manner modified; for, had the area been open to immigration, these same places would have been seized on by intruders. In such case, every slight modification, which in the course of ages chanced to arise, and which in any way favoured the individuals of any of the species, by better adapting them to their altered conditions, would tend to be preserved; and natural selection would thus have free scope for the work of improvement.

 . . . As man can produce and certainly has produced a great result by his methodical and unconscious means of selection, what may not nature effect? Man can act only on external and visible characters: nature cares nothing for appearances, except in so far as they may be useful to any being. She can act on every internal organ, on every shade of constitutional difference, on the whole machinery of life. Man selects only for his own good; Nature only for that of the being which she tends. Every selected character is fully exercised by her; and the being is placed under well-suited conditions of life. Man keeps the natives of many climates in the same country; he seldom exercises each selected character in some peculiar and fitting manner; he feeds a long and a short beaked pigeon on the same food; he does not exercise a long-backed or long-legged quadruped in any peculiar manner; he exposes sheep with long and short wool to the same climate. He does not allow the most vigorous males to struggle for the females. He does not rigidly destroy all inferior animals, but protects during each varying season, as far as lies in his power, all his productions. He often begins his selection by some half-monstrous form; or at least by some modification prominent enough to catch his eye, or to be plainly useful to him. Under nature, the slightest difference of structure or constitution may well turn the nicely-balanced scale in the struggle for life, and so be preserved. How fleeting are the wishes and efforts of man! how short his time! and consequently how poor will his products be, compared with those accumulated by nature during whole geological periods. Can we wonder, then, that nature’s productions should be far “truer” in character than man’s productions; that they should be infinitely better adapted to the most complex conditions of life, and should plainly bear the stamp of far higher workmanship?

It may be said that natural selection is daily and hourly scrutinising, throughout the world, every variation, even the slightest; rejecting that which is bad, preserving and adding up all that is good; silently and insensibly working, whenever and wherever opportunity offers, at the improvement of each organic being in relation to its organic and inorganic conditions of life. We see nothing of these slow changes in progress, until the hand of time has marked the long lapse of ages, and then so imperfect is our view into long past geological ages, that we only see that the forms of life are now different from what they formerly were.

Although natural selection can act only through and for the good of each being, yet characters and structures, which we are apt to consider as of very trifling importance, may thus be acted on. When we see leaf-eating insects green, and bark-feeders mottled-grey; the alpine ptarmigan white in winter, the red-grouse the colour of heather, and the black-grouse that of peaty earth, we must believe that these tints are of service to these birds and insects in preserving them from danger. .... In plants the down on the fruit and the colour of the flesh are considered by botanists as characters of the most trifling importance: yet we hear from an excellent horticulturist, Downing, that in the United States smooth-skinned fruits suffer far more from a beetle, a curculio, than those with down; that purple plums suffer far more from a certain disease than yellow plums; whereas another disease attacks yellow-fleshed peaches far more than those with other coloured flesh. If, with all the aids of art, these slight differences make a great difference in cultivating the several varieties, assuredly, in a state of nature, where the trees would have to struggle with other trees and with a host of enemies, such differences would effectually settle which variety, whether a smooth or downy, a yellow or purple fleshed fruit, should succeed.


As we see that those variations which under domestication appear at any particular period of life, tend to reappear in the offspring at the same period; — for instance, in the seeds of the many varieties of our culinary and agricultural plants; in the caterpillar and cocoon stages of the varieties of the silkworm; in the eggs of poultry, and in the colour of the down of their chickens; in the horns of our sheep and cattle when nearly adult; — so in a state of nature, natural selection will be enabled to act on and modify organic beings at any age, by the accumulation of profitable variations at that age, and by their inheritance at a corresponding age. If it profit a plant to have its seeds more and more widely disseminated by the wind, I can see no greater difficulty in this being effected through natural selection, than in the cotton-planter increasing and improving by selection the down in the pods on his cotton-trees. Natural selection may modify and adapt the larva of an insect to a score of contingencies, wholly different from those which concern the mature insect. These modifications will no doubt affect, through the laws of correlation, the structure of the adult; and probably in the case of those insects which live only for a few hours, and which never feed, a large part of their structure is merely the correlated result of successive changes in the structure of their larvae. So, conversely, modifications in the adult will probably often affect the structure of the larva; but in all cases natural selection will ensure that modifications consequent on other modifications at a different period of life, shall not be in the least degree injurious: for if they became so, they would cause the extinction of the species.

Natural selection will modify the structure of the young in relation to the parent, and of the parent in relation to the young. In social animals it will adapt the structure of each individual for the benefit of the community; if each in consequence profits by the selected change. What natural selection cannot do, is to modify the structure of one species, without giving it any advantage, for the good of another species; and though statements to this effect may be found in works of natural history, I cannot find one case which will bear investigation. A structure used only once in an animal’s whole life, if of high importance to it, might be modified to any extent by natural selection; for instance, the great jaws possessed by certain insects, and used exclusively for opening the cocoon — or the hard tip to the beak of nestling birds, used for breaking the egg. It has been asserted, that of the best short-beaked tumbler-pigeons more perish in the egg than are able to get out of it; so that fanciers assist in the act of hatching. Now, if nature had to make the beak of a full-grown pigeon very short for the bird’s own advantage, the process of modification would be very slow, and there would be simultaneously the most rigorous selection of the young birds within the egg, which had the most powerful and hardest beaks, for all with weak beaks would inevitably perish: or, more delicate and more easily broken shells might be selected, the thickness of the shell being known to vary like every other structure.

Sexual Selection. — Inasmuch as peculiarities often appear under domestication in one sex and become hereditarily attached to that sex, the same fact probably occurs under nature, and if so, natural selection will be able to modify one sex in its functional relations to the other sex, or in relation to wholly different habits of life in the two sexes, as is sometimes the case with insects. And this leads me to say a few words on what I call Sexual Selection. This depends, not on a struggle for existence, but on a struggle between the males for possession of the females; the result is not death to the unsuccessful competitor, but few or no offspring. Sexual selection is, therefore, less rigorous than natural selection. Generally, the most vigorous males, those which are best fitted for their places in nature, will leave most progeny. But in many cases, victory will depend not on general vigour, but on having special weapons, confined to the male sex. A hornless stag or spurless cock would have a poor chance of leaving offspring. Sexual selection by always allowing the victor to breed might surely give indomitable courage, length to the spur, and strength to the wing to strike in the spurred leg, as well as the brutal cock-fighter, who knows well that he can improve his breed by careful selection of the best cocks. How low in the scale of nature this law of battle descends, I know not; male alligators have been described as fighting, bellowing, and whirling round, like Indians in a war-dance, for the possession of the females; male salmons have been seen fighting all day long; male stag-beetles often bear wounds from the huge mandibles of other males. The war is, perhaps, severest between the males of polygamous animals, and these seem oftenest provided with special weapons. The males of carnivorous animals are already well armed; though to them and to others, special means of defence may be given through means of sexual selection, as the mane to the lion, the shoulder-pad to the boar, and the hooked jaw to the male salmon; for the shield may be as important for victory, as the sword or spear.

Amongst birds, the contest is often of a more peaceful character. All those who have attended to the subject, believe that there is the severest rivalry between the males of many species to attract by singing the females. The rock-thrush of Guiana, birds of Paradise, and some others, congregate; and successive males display their gorgeous plumage and perform strange antics before the females, which standing by as spectators, at last choose the most attractive partner. .... [I]f man can in a short time give elegant carriage and beauty to his bantams, according to his standard of beauty, I can see no good reason to doubt that female birds, by selecting, during thousands of generations, the most melodious or beautiful males, according to their standard of beauty, might produce a marked effect.


In order to make it clear how, as I believe, natural selection acts, I must beg permission to give one or two imaginary illustrations. Let us take the case of a wolf, which preys on various animals, securing some by craft, some by strength, and some by fleetness; and let us suppose that the fleetest prey, a deer for instance, had from any change in the country increased in numbers, or that other prey had decreased in numbers, during that season of the year when the wolf is hardest pressed for food. I can under such circumstances see no reason to doubt that the swiftest and slimmest wolves would have the best chance of surviving, and so be preserved or selected, — provided always that they retained strength to master their prey at this or at some other period of the year, when they might be compelled to prey on other animals. I can see no more reason to doubt this, than that man can improve the fleetness of his greyhounds by careful and methodical selection, or by that unconscious selection which results from each man trying to keep the best dogs without any thought of modifying the breed.

Even without any change in the proportional numbers of the animals on which our wolf preyed, a cub might be born with an innate tendency to pursue certain kinds of prey. Nor can this be thought very improbable; for we often observe great differences in the natural tendencies of our domestic animals; one cat, for instance, taking to catch rats, another mice ... The tendency to catch rats rather than mice is known to be inherited. Now, if any slight innate change of habit or of structure benefited an individual wolf, it would have the best chance of surviving and of leaving offspring. Some of its young would probably inherit the same habits or structure, and by the repetition of this process, a new variety might be formed which would either supplant or coexist with the parent-form of wolf. Or, again, the wolves inhabiting a mountainous district, and those frequenting the lowlands, would naturally be forced to hunt different prey; and from the continued preservation of the individuals best fitted for the two sites, two varieties might slowly be formed. .... I may add, that, according to Mr. Pierce, there are two varieties of the wolf inhabiting the Catskill Mountains in the United States, one with a light greyhound-like form, which pursues deer, and the other more bulky, with shorter legs, which more frequently attacks the shepherd’s flocks.

Let us now take a more complex case. Certain plants excrete a sweet juice, apparently for the sake of eliminating something injurious from their sap ... . This juice, though small in quantity, is greedily sought by insects. Let us now suppose a little sweet juice or nectar to be excreted by the inner bases of the petals of a flower. In this case insects in seeking the nectar would get dusted with pollen, and would certainly often transport the pollen from one flower to the stigma of another flower. The flowers of two distinct individuals of the same species would thus get crossed; and the act of crossing, we have good reason to believe (as will hereafter be more fully alluded to), would produce very vigorous seedlings, which consequently would have the best chance of flourishing and surviving. Some of these seedlings would probably inherit the nectar-excreting power. Those individual flowers which had the largest glands or nectaries, and which excreted most nectar, would be oftenest visited by insects, and would be oftenest crossed; and so in the long-run would gain the upper hand. Those flowers, also, which had their stamens and pistils placed, in relation to the size and habits of the particular insects which visited them, so as to favour in any degree the transportal of their pollen from flower to flower, would likewise be favoured or selected. ....

When our plant, by this process of the continued preservation or natural selection of more and more attractive flowers, had been rendered highly attractive to insects, they would, unintentionally on their part, regularly carry pollen from flower to flower; and that they can most effectually do this, I could easily show by many striking instances. I will give only one — not as a very striking case, but as likewise illustrating one step in the separation of the sexes of plants.... Some holly-trees bear only male flowers, which have four stamens producing rather a small quantity of pollen, and a rudimentary pistil; other holly-trees bear only female flowers; these have a full-sized pistil, and four stamens with shrivelled anthers, in which not a grain of pollen can be detected. Having found a female tree exactly sixty yards from a male tree, I put the stigmas of twenty flowers, taken from different branches, under the microscope, and on all, without exception, there were pollen-grains, and on some a profusion of pollen. As the wind had set for several days from the female to the male tree, the pollen could not thus have been carried. The weather had been cold and boisterous, and therefore not favourable to bees, nevertheless every female flower which I examined had been effectually fertilised by the bees, accidentally dusted with pollen, having flown from tree to tree in search of nectar. But to return to our imaginary case: as soon as the plant had been rendered so highly attractive to insects that pollen was regularly carried from flower to flower, another process might commence. No naturalist doubts the advantage of what has been called the “physiological division of labour;” hence we may believe that it would be advantageous to a plant to produce stamens alone in one flower or on one whole plant, and pistils alone in another flower or on another plant. In plants under culture and placed under new conditions of life, sometimes the male organs and sometimes the female organs become more or less impotent; now if we suppose this to occur in ever so slight a degree under nature, then as pollen is already carried regularly from flower to flower, and as a more complete separation of the sexes of our plant would be advantageous on the principle of the division of labour, individuals with this tendency more and more increased, would be continually favoured or selected, until at last a complete separation of the sexes would be effected.

Let us now turn to the nectar-feeding insects in our imaginary case: we may suppose the plant of which we have been slowly increasing the nectar by continued selection, to be a common plant; and that certain insects depended in main part on its nectar for food. I could give many facts, showing how anxious bees are to save time; for instance, their habit of cutting holes and sucking the nectar at the bases of certain flowers, which they can, with a very little more trouble, enter by the mouth. Bearing such facts in mind, I can see no reason to doubt that an accidental deviation in the size and form of the body, or in the curvature and length of the proboscis, etc., far too slight to be appreciated by us, might profit a bee or other insect, so that an individual so characterised would be able to obtain its food more quickly, and so have a better chance of living and leaving descendants. Its descendants would probably inherit a tendency to a similar slight deviation of structure. The tubes of the corollas of the common red and incarnate clovers (Trifolium pratense and incarnatum) do not on a hasty glance appear to differ in length; yet the hive-bee can easily suck the nectar out of the incarnate clover, but not out of the common red clover, which is visited by humble-bees alone; so that whole fields of the red clover offer in vain an abundant supply of precious nectar to the hive-bee. Thus it might be a great advantage to the hive-bee to have a slightly longer or differently constructed proboscis. On the other hand, I have found by experiment that the fertility of clover greatly depends on bees visiting and moving parts of the corolla, so as to push the pollen on to the stigmatic surface. Hence, again, if humble-bees were to become rare in any country, it might be a great advantage to the red clover to have a shorter or more deeply divided tube to its corolla, so that the hive-bee could visit its flowers. Thus I can understand how a flower and a bee might slowly become, either simultaneously or one after the other, modified and adapted in the most perfect manner to each other, by the continued preservation of individuals presenting mutual and slightly favourable deviations of structure.

I am well aware that this doctrine of natural selection, exemplified in the above imaginary instances, is open to the same objections which were at first urged against Sir Charles Lyell’s noble views on “the modern changes of the earth, as illustrative of geology;” but we now very seldom hear the action, for instance, of the coast-waves, called a trifling and insignificant cause, when applied to the excavation of gigantic valleys or to the formation of the longest lines of inland cliffs. Natural selection can act only by the preservation and accumulation of infinitesimally small inherited modifications [variations], each profitable to the preserved being; and as modern geology has almost banished such views as the excavation of a great valley by a single diluvial wave, so will natural selection, if it be a true principle, banish the belief of the continued creation of new organic beings, or of any great and sudden modification in their structure....

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.

 . . . 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.

 . . . 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 ... 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 ... . 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?... 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. .... [W]e 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. .... 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.


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. .... [O]n 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.

If during the long course of ages and under varying conditions of life, organic beings vary at all in the several parts of their organisation, and I think this cannot be disputed; if there be, owing to the high geometrical powers of increase of [in] each species, at some age, season, or year, a severe struggle for life, and this certainly cannot be disputed; then, considering the infinite complexity of the relations of all organic beings to each other and to their conditions of existence, causing an infinite diversity in structure, constitution, and habits, to be advantageous to them, I think it would be a most extraordinary fact if no variation ever had occurred useful to each being’s own welfare, in the same way as so many variations have occurred useful to man. But if variations useful to any organic being do occur, assuredly individuals thus characterised will have the best chance of being preserved in the struggle for life; and from the strong principle of inheritance they will tend to produce offspring similarly characterised. This principle of preservation, I have called, for the sake of brevity, Natural Selection. Natural selection, on the principle of qualities being inherited at corresponding ages, can modify the egg, seed, or young, as easily as the adult. Amongst many animals, sexual selection will give its aid to ordinary selection, by assuring to the most vigorous and best adapted males the greatest number of offspring. Sexual selection will also give characters useful to the males alone, in their struggles with other males.

Whether natural selection has really thus acted in nature, in modifying and adapting the various forms of life to their several conditions and stations, must be judged of by the general tenour and balance of evidence given in the following chapters. But we already see how it entails extinction; and how largely extinction has acted in the world’s history, geology plainly declares. Natural selection, also, leads to divergence of character; for more living beings can be supported on the same area the more they diverge in structure, habits, and constitution, of which we see proof by looking at the inhabitants of any small spot or at naturalised productions. Therefore during the modification of the descendants of any one species, and during the incessant struggle of all species to increase in numbers, the more diversified these descendants become, the better will be their chance of succeeding in the battle of life. Thus the small differences distinguishing varieties of the same species, will steadily tend to increase till they come to equal the greater differences between species of the same genus, or even of distinct genera.

We have seen that it is the common, the widely-diffused, and widely-ranging species, belonging to the larger genera, which vary most; and these will tend to transmit to their modified offspring that superiority which now makes them dominant in their own countries. Natural selection, as has just been remarked, leads to divergence of character and to much extinction of the less improved and intermediate forms of life. On these principles, I believe, the nature of the affinities of all organic beings may be explained. It is a truly wonderful fact — the wonder of which we are apt to overlook from familiarity — that all animals and all plants throughout all time and space should be related to each other in group subordinate to group, in the manner which we everywhere behold — namely, varieties of the same species most closely related together, species of the same genus less closely and unequally related together, forming sections and sub-genera, species of distinct genera much less closely related, and genera related in different degrees, forming sub-families, families, orders, sub-classes, and classes. The several subordinate groups in any class cannot be ranked in a single file, but seem rather to be clustered round points, and these round other points, and so on in almost endless cycles. On the view that each species has been independently created, I can see no explanation of this great fact in the classification of all organic beings; but, to the best of my judgment, it is explained through inheritance and the complex action of natural selection, entailing extinction and divergence of character ... .

The affinities of all the beings of the same class have sometimes been represented by a great tree. I believe this simile largely speaks the truth. The green and budding twigs may represent existing species; and those produced during each former year may represent the long succession of extinct species. At each period of growth all the growing twigs have tried to branch out on all sides, and to overtop and kill the surrounding twigs and branches, in the same manner as species and groups of species have tried to overmaster other species in the great battle for life. The limbs divided into great branches, and these into lesser and lesser branches, were themselves once, when the tree was small, budding twigs; and this connexion of the former and present buds by ramifying branches may well represent the classification of all extinct and living species in groups subordinate to groups. Of the many twigs which flourished when the tree was a mere bush, only two or three, now grown into great branches, yet survive and bear all the other branches; so with the species which lived during long-past geological periods, very few now have living and modified descendants. From the first growth of the tree, many a limb and branch has decayed and dropped off; and these lost branches of various sizes may represent those whole orders, families, and genera which have now no living representatives, and which are known to us only from having been found in a fossil state. As we here and there see a thin straggling branch springing from a fork low down in a tree, and which by some chance has been favoured and is still alive on its summit, so we occasionally see an animal like the Ornithorhynchus [platypus] or Lepidosiren [lungfish], which in some small degree connects by its affinities two large branches of life, and which has apparently been saved from fatal competition by having inhabited a protected station. As buds give rise by growth to fresh buds, and these, if vigorous, branch out and overtop on all sides many a feebler branch, so by generation I believe it has been with the great Tree of Life, which fills with its dead and broken branches the crust of the earth, and covers the surface with its ever branching and beautiful ramifications.



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

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

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

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

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

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

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


On the origin and transitions of organic beings with peculiar habits and structure.— It has been asked by the opponents of such views as I hold, how, for instance, a land carnivorous animal could have been converted into one with aquatic habits; for how could the animal in its transitional state have subsisted? It would be easy to show that within the same group carnivorous animals exist having every intermediate grade between truly aquatic and strictly terrestrial habits; and as each exists by a struggle for life, it is clear that each is well adapted in its habits to its place in nature. Look at the Mustela vison of North America, which has webbed feet and which resembles an otter in its fur, short legs, and form of tail; during summer this animal dives for and preys on fish, but during the long winter it leaves the frozen waters, and preys like other polecats on mice and land animals. If a different case had been taken, and it had been asked how an insectivorous quadruped could possibly have been converted into a flying bat, the question would have been far more difficult, and I could have given no answer. Yet I think such difficulties have very little weight.

Here, as on other occasions, I lie under a heavy disadvantage, for out of the many striking cases which I have collected, I can give only one or two instances of transitional habits and structures in closely allied species of the same genus; and of diversified habits, either constant or occasional, in the same species. And it seems to me that nothing less than a long list of such cases is sufficient to lessen the difficulty in any particular case like that of the bat.


[L]ook at the Galeopithecus or flying lemur, which formerly was falsely ranked amongst bats. It has an extremely wide flank-membrane, stretching from the corners of the jaw to the tail, and including the limbs and the elongated fingers: the flank membrane is, also, furnished with an extensor muscle. Although no graduated links of structure, fitted for gliding through the air, now connect the Galeopithecus with the other Lemuridae, yet I can see no difficulty in supposing that such links formerly existed ... and that each grade of structure had been useful to its possessor. [Editor’s note: Modern classification does not regard this animal as a lemur at all.] Nor can I see any insuperable difficulty in further believing it possible that the membrane-connected fingers and fore-arm of the Galeopithecus might be greatly lengthened by natural selection; and this, as far as the organs of flight are concerned, would convert it into a bat. In bats which have the wing-membrane extended from the top of the shoulder to the tail, including the hind-legs, we perhaps see traces of an apparatus originally constructed for gliding through the air rather than for flight.

If about a dozen genera of birds had become extinct or were unknown, who would have ventured to have surmised that birds might have existed which used their wings solely as flappers, like the logger-headed duck (Micropterus of Eyton); as fins in the water and front legs on the land, like the penguin; as sails, like the ostrich; and functionally for no purpose, like the Apteryx kiwi. Yet the structure of each of these birds is good for it, under the conditions of life to which it is exposed, for each has to live by a struggle; but it is not necessarily the best possible under all possible conditions. It must not be inferred from these remarks that any of the grades of wing-structure here alluded to, which perhaps may all have resulted from disuse, indicate the natural steps by which birds have acquired their perfect power of flight; but they serve, at least, to show what diversified means of transition are possible.

Seeing that a few members of such water-breathing classes as the Crustacea and Mollusca are adapted to live on the land, and seeing that we have flying birds and mammals, flying insects of the most diversified types, and formerly had flying reptiles, it is conceivable that flying-fish, which now glide far through the air, slightly rising and turning by the aid of their fluttering fins, might have been modified into perfectly winged animals. If this had been effected, who would have ever imagined that in an early transitional state they had been inhabitants of the open ocean, and had used their incipient organs of flight exclusively, as far as we know, to escape being devoured by other fish?

When we see any structure highly perfected for any particular habit, as the wings of a bird for flight, we should bear in mind that animals displaying early transitional grades of the structure will seldom continue to exist to the present day, for they will have been supplanted by the very process of perfection through natural selection. Furthermore, we may conclude that transitional grades between structures fitted for very different habits of life will rarely have been developed at an early period in great numbers and under many subordinate forms. Thus, to return to our imaginary illustration of the flying-fish, it does not seem probable that fishes capable of true flight would have been developed under many subordinate forms, for taking prey of many kinds in many ways, on the land and in the water, until their organs of flight had come to a high stage of perfection, so as to have given them a decided advantage over other animals in the battle for life. Hence the chance of discovering species with transitional grades of structure in a fossil condition will always be less, from their having existed in lesser numbers, than in the case of species with fully developed structures.

I will now give two or three instances of diversified and of changed habits in the individuals of the same species. When either case occurs, it would be easy for natural selection to fit the animal, by some modification of its structure, for its changed habits, or exclusively for one of its several different habits. But it is difficult to tell, and immaterial for us, whether habits generally change first and structure afterwards; or whether slight modifications of structure lead to changed habits; both probably often change almost simultaneously. Of cases of changed habits it will suffice merely to allude to that of the many British insects which now feed on exotic plants, or exclusively on artificial substances. Of diversified habits innumerable instances could be given: I have often watched a tyrant flycatcher (Saurophagus sulphuratus) in South America, hovering over one spot and then proceeding to another, like a kestrel, and at other times standing stationary on the margin of water, and then dashing like a kingfisher at a fish. In our own country the larger titmouse (Parus major) may be seen climbing branches, almost like a creeper; it often, like a shrike, kills small birds by blows on the head; and I have many times seen and heard it hammering the seeds of the yew on a branch, and thus breaking them like a nuthatch. In North America the black bear was seen by Hearne swimming for hours with widely open mouth, thus catching, like a whale, insects in the water. Even in so extreme a case as this, if the supply of insects were constant, and if better adapted competitors did not already exist in the country, I can see no difficulty in a race of bears being rendered, by natural selection, more and more aquatic in their structure and habits, with larger and larger mouths, till a creature was produced as monstrous as a whale.

As we sometimes see individuals of a species following habits widely different from those both of their own species and of the other species of the same genus, we might expect, on my theory, that such individuals would occasionally have given rise to new species, having anomalous habits, and with their structure either slightly or considerably modified from that of their proper type. And such instances do occur in nature. Can a more striking instance of adaptation be given than that of a woodpecker for climbing trees and for seizing insects in the chinks of the bark? Yet in North America there are woodpeckers which feed largely on fruit, and others with elongated wings which chase insects on the wing; and on the plains of La Plata, where not a tree grows, there is a woodpecker, which in every essential part of its organisation, even in its colouring, in the harsh tone of its voice, and undulatory flight, told me plainly of its close blood-relationship to our common species; yet it is a woodpecker which never climbs a tree!


He who believes that each being has been created as we now see it, must occasionally have felt surprise when he has met with an animal having habits and structure not at all in agreement. What can be plainer than that the webbed feet of ducks and geese are formed for swimming? yet there are upland geese with webbed feet which rarely or never go near the water; and no one except Audubon has seen the frigate-bird, which has all its four toes webbed, alight on the surface of the sea. On the other hand, grebes and coots are eminently aquatic, although their toes are only bordered by membrane.... In such cases ... habits have changed without a corresponding change of structure. The webbed feet of the upland goose may be said to have become rudimentary in function, though not in structure. In the frigate-bird, the deeply-scooped membrane between the toes shows that structure has begun to change.

He who believes in separate and innumerable acts of creation will say, that in these cases it has pleased the Creator to cause a being of one type to take the place of one of another type; but this seems to me only restating the fact in dignified language. He who believes in the struggle for existence and in the principle of natural selection, will acknowledge that every organic being is constantly endeavouring to increase in numbers; and that if any one being vary ever so little, either in habits or structure, and thus gain an advantage over some other inhabitant of the country, it will seize on the place of that inhabitant, however different it may be from its own place. Hence it will cause him no surprise that there should be geese and frigate-birds with webbed feet, either living on the dry land or most rarely alighting on the water; that there should be long-toed corncrakes living in meadows instead of in swamps; that there should be woodpeckers where not a tree grows; that there should be diving thrushes, and petrels with the habits of auks.

Organs of extreme perfection and complication.— To suppose that the eye, with all its inimitable contrivances for adjusting the focus to different distances, for admitting different amounts of light, and for the correction of spherical and chromatic aberration, could have been formed by natural selection, seems, I freely confess, absurd in the highest possible degree. Yet reason tells me, that if numerous gradations from a perfect and complex eye to one very imperfect and simple, each grade being useful to its possessor, can be shown to exist; if further, the eye does vary ever so slightly, and the variations be inherited, which is certainly the case; and if any variation or modification in the organ be ever useful to an animal under changing conditions of life, then the difficulty of believing that a perfect and complex eye could be formed by natural selection, though insuperable by our imagination, can hardly be considered real. How a nerve comes to be sensitive to light, hardly concerns us more than how life itself first originated; but I may remark that several facts make me suspect that any sensitive nerve may be rendered sensitive to light, and likewise to those coarser vibrations of the air which produce sound.

In looking for the gradations by which an organ in any species has been perfected, we ought to look exclusively to its lineal ancestors; but this is scarcely ever possible, and we are forced in each case to look to species of the same group, that is to the collateral descendants from the same original parent-form, in order to see what gradations are possible, and for the chance of some gradations having been transmitted from the earlier stages of descent, in an unaltered or little altered condition. Amongst existing Vertebrata, we find but a small amount of gradation in the structure of the eye, and from fossil species we can learn nothing on this head. In this great class we should probably have to descend far beneath the lowest known fossiliferous stratum to discover the earlier stages, by which the eye has been perfected.

In the Articulata we can commence a series with an optic nerve merely coated with pigment, and without any other mechanism; and from this low stage, numerous gradations of structure, branching off in two fundamentally different lines, can be shown to exist, until we reach a moderately high stage of perfection. In certain crustaceans, for instance, there is a double cornea, the inner one divided into facets, within each of which there is a lens-shaped swelling. In other crustaceans the transparent cones which are coated by pigment, and which properly act only by excluding lateral pencils of light, are convex at their upper ends and must act by convergence; and at their lower ends there seems to be an imperfect vitreous substance. With these facts, here far too briefly and imperfectly given, which show that there is much graduated diversity in the eyes of living crustaceans, and bearing in mind how small the number of living animals is in proportion to those which have become extinct, I can see no very great difficulty (not more than in the case of many other structures) in believing that natural selection has converted the simple apparatus of an optic nerve merely coated with pigment and invested by transparent membrane, into an optical instrument as perfect as is possessed by any member of the great Articulate class.

He who will go thus far, if he find on finishing this treatise that large bodies of facts, otherwise inexplicable, can be explained by the theory of descent, ought not to hesitate to go further, and to admit that a structure even as perfect as the eye of an eagle might be formed by natural selection, although in this case he does not know any of the transitional grades. His reason ought to conquer his imagination; though I have felt the difficulty far too keenly to be surprised at any degree of hesitation in extending the principle of natural selection to such startling lengths.

It is scarcely possible to avoid comparing the eye to a telescope. We know that this instrument has been perfected by the long-continued efforts of the highest human intellects; and we naturally infer that the eye has been formed by a somewhat analogous process. But may not this inference be presumptuous? Have we any right to assume that the Creator works by intellectual powers like those of man? If we must compare the eye to an optical instrument, we ought in imagination to take a thick layer of transparent tissue, with a nerve sensitive to light beneath, and then suppose every part of this layer to be continually changing slowly in density, so as to separate into layers of different densities and thicknesses, placed at different distances from each other, and with the surfaces of each layer slowly changing in form. Further we must suppose that there is a power always intently watching each slight accidental alteration in the transparent layers; and carefully selecting each alteration which, under varied circumstances, may in any way, or in any degree, tend to produce a distincter image. We must suppose each new state of the instrument to be multiplied by the million; and each to be preserved till a better be produced, and then the old ones to be destroyed. In living bodies, variation will cause the slight alterations, generation will multiply them almost infinitely, and natural selection will pick out with unerring skill each improvement. Let this process go on for millions on millions of years; and during each year on millions of individuals of many kinds; and may we not believe that a living optical instrument might thus be formed as superior to one of glass, as the works of the Creator are to those of man?

If it could be demonstrated that any complex organ existed, which could not possibly have been formed by numerous, successive, slight modifications, my theory would absolutely break down. But I can find out no such case....

 Two distinct organs sometimes perform simultaneously the same function in the same individual; to give one instance, there are fish with gills or branchiae that breathe the air dissolved in the water, at the same time that they breathe free air in their swimbladders, this latter organ having a ductus pneumaticus for its supply, and being divided by highly vascular partitions. In these cases, one of the two organs might with ease be modified and perfected so as to perform all the work by itself, being aided during the process of modification by the other organ; and then this other organ might be modified for some other and quite distinct purpose, or be quite obliterated.


We can thus ... understand the strange fact that every particle of food and drink which we swallow has to pass over the orifice of the trachea, with some risk of falling into the lungs, notwithstanding the beautiful contrivance by which the glottis is closed. In the higher Vertebrata the branchiae have wholly disappeared — the slits on the sides of the neck and the loop-like course of the arteries still marking in the embryo their former position. But it is conceivable that the now utterly lost branchiae might have been gradually worked in by natural selection for some quite distinct purpose: in the same manner as, on the view entertained by some naturalists that the branchiae and dorsal scales of Annelids are homologous with the wings and wing-covers of insects, it is probable that organs which at a very ancient period served for respiration have been actually converted into organs of flight.

 . . . [U]ndoubtedly, grave cases of difficulty occur, some of which will be discussed in my future work.

One of the gravest is that of neuter insects, which are often very differently constructed from either the males or fertile females; but this case will be treated of in the next chapter. ....

Although in many cases it is most difficult to conjecture by what transitions an organ could have arrived at its present state; yet, considering that the proportion of living and known forms to the extinct and unknown is very small, I have been astonished how rarely an organ can be named, towards which no transitional grade is known to lead. The truth of this remark is indeed shown by that old canon in natural history of “Natura non facit saltum [nature does not make a jump].” We meet with this admission in the writings of almost every experienced naturalist; or, as Milne Edwards has well expressed it, nature is prodigal in variety, but niggard in innovation. Why, on the theory of Creation, should this be so? Why should all the parts and organs of many independent beings, each supposed to have been separately created for its proper place in nature, be so invariably linked together by graduated steps? Why should not Nature have taken a leap from structure to structure? On the theory of natural selection, we can clearly understand why she should not; for natural selection can act only by taking advantage of slight successive variations; she can never take a leap, but must advance by the shortest and slowest steps.

Organs of little apparent importance.— As natural selection acts by life and death, — by the preservation of individuals with any favourable variation, and by the destruction of those with any unfavourable deviation of structure, — I have sometimes felt much difficulty in understanding the origin of simple parts, of which the importance does not seem sufficient to cause the preservation of successively varying individuals. I have sometimes felt as much difficulty, though of a very different kind, on this head, as in the case of an organ as perfect and complex as the eye.

In the first place, we are much too ignorant in regard to the whole economy of any one organic being, to say what slight modifications would be of importance or not. In a former chapter I have given instances of most trifling characters, such as the down on fruit and the colour of the flesh, which, from determining the attacks of insects or from being correlated with constitutional differences, might assuredly be acted on by natural selection. The tail of the giraffe looks like an artificially constructed fly-flapper; and it seems at first incredible that this could have been adapted for its present purpose by successive slight modifications, each better and better, for so trifling an object as driving away flies; yet we should pause before being too positive even in this case, for we know that the distribution and existence of cattle and other animals in South America absolutely depends on their power of resisting the attacks of insects: so that individuals which could by any means defend themselves from these small enemies, would be able to range into new pastures and thus gain a great advantage. It is not that the larger quadrupeds are actually destroyed (except in some rare cases) by the flies, but they are incessantly harassed and their strength reduced, so that they are more subject to disease, or not so well enabled in a coming dearth to search for food, or to escape from beasts of prey.

Organs now of trifling importance have probably in some cases been of high importance to an early progenitor, and, after having been slowly perfected at a former period, have been transmitted in nearly the same state, although now become of very slight use; and any actually injurious deviations in their structure will always have been checked by natural selection. Seeing how important an organ of locomotion the tail is in most aquatic animals, its general presence and use for many purposes in so many land animals, which in their lungs or modified swim-bladders betray their aquatic origin, may perhaps be thus accounted for. A well-developed tail having been formed in an aquatic animal, it might subsequently come to be worked in for all sorts of purposes, as a fly-flapper, an organ of prehension, or as an aid in turning, as with the dog, though the aid must be slight, for the hare, with hardly any tail, can double quickly enough.

In the second place, we may sometimes attribute importance to characters which are really of very little importance, and which have originated from quite secondary causes, independently of natural selection. We should remember that ... correlation of growth will have had a most important influence in modifying various structures; and finally, that sexual selection will often have largely modified the external characters of animals having a will, to give one male an advantage in fighting with another or in charming the females. Moreover when a modification of structure has primarily arisen from the above or other unknown causes, it may at first have been of no advantage to the species, but may subsequently have been taken advantage of by the descendants of the species under new conditions of life and with newly acquired habits.

To give a few instances to illustrate these latter remarks. If green woodpeckers alone had existed, and we did not know that there were many black and pied kinds, I dare say that we should have thought that the green colour was a beautiful adaptation to hide this tree-frequenting bird from its enemies; and consequently that it was a character of importance and might have been acquired through natural selection; as it is, I have no doubt that the colour is due to some quite distinct cause, probably to sexual selection. A trailing bamboo in the Malay Archipelago climbs the loftiest trees by the aid of exquisitely constructed hooks clustered around the ends of the branches, and this contrivance, no doubt, is of the highest service to the plant; but as we see nearly similar hooks on many trees which are not climbers, the hooks on the bamboo may have arisen from unknown laws of growth, and have been subsequently taken advantage of by the plant undergoing further modification and becoming a climber.


The foregoing remarks lead me to say a few words on the protest lately made by some naturalists, against the utilitarian doctrine that every detail of structure has been produced for the good of its possessor. They believe that very many structures have been created for beauty in the eyes of man, or for mere variety. This doctrine, if true, would be absolutely fatal to my theory. Yet I fully admit that many structures are of no direct use to their possessors. Physical conditions probably have had some little effect on structure, quite independently of any good thus gained. Correlation of growth has no doubt played a most important part, and a useful modification of one part will often have entailed on other parts diversified changes of no direct use. So again characters which formerly were useful, or which formerly had arisen from correlation of growth, or from other unknown cause, may reappear from the law of reversion, though now of no direct use. The effects of sexual selection, when displayed in beauty to charm the females, can be called useful only in rather a forced sense. But by far the most important consideration is that the chief part of the organisation of every being is simply due to inheritance; and consequently, though each being assuredly is well fitted for its place in nature, many structures now have no direct relation to the habits of life of each species. Thus, we can hardly believe that the webbed feet of the upland goose or of the frigate-bird are of special use to these birds; we cannot believe that the same bones in the arm of the monkey, in the fore leg of the horse, in the wing of the bat, and in the flipper of the seal, are of special use to these animals. We may safely attribute these structures to inheritance. But to the progenitor of the upland goose and of the frigate-bird, webbed feet no doubt were as useful as they now are to the most aquatic of existing birds. So we may believe that the progenitor of the seal had not a flipper, but a foot with five toes fitted for walking or grasping; and we may further venture to believe that the several bones in the limbs of the monkey, horse, and bat, which have been inherited from a common progenitor, were formerly of more special use to that progenitor, or its progenitors, than they now are to these animals having such widely diversified habits. Therefore we may infer that these several bones might have been acquired through natural selection, subjected formerly, as now, to the several laws of inheritance, reversion, correlation of growth, etc. Hence every detail of structure in every living creature (making some little allowance for the direct action of physical conditions) may be viewed, either as having been of special use to some ancestral form, or as being now of special use to the descendants of this form — either directly, or indirectly through the complex laws of growth.

Natural selection cannot possibly produce any modification in any one species exclusively for the good of another species; though throughout nature one species incessantly takes advantage of, and profits by, the structure of another.... If it could be proved that any part of the structure of any one species had been formed for the exclusive good of another species, it would annihilate my theory, for such could not have been produced through natural selection. Although many statements may be found in works on natural history to this effect, I cannot find even one which seems to me of any weight. It is admitted that the rattlesnake has a poison-fang for its own defence and for the destruction of its prey; but some authors suppose that at the same time this snake is furnished with a rattle for its own injury, namely, to warn its prey to escape. I would almost as soon believe that the cat curls the end of its tail when preparing to spring, in order to warn the doomed mouse. But I have not space here to enter on this and other such cases.

Natural selection will never produce in a being anything injurious to itself, for natural selection acts solely by and for the good of each. No organ will be formed, as Paley has remarked, for the purpose of causing pain or for doing an injury to its possessor. If a fair balance be struck between the good and evil caused by each part, each will be found on the whole advantageous. After the lapse of time, under changing conditions of life, if any part comes to be injurious, it will be modified; or if it be not so, the being will become extinct, as myriads have become extinct.

Natural selection tends only to make each organic being as perfect as, or slightly more perfect than, the other inhabitants of the same country with which it has to struggle for existence. And we see that this is the degree of perfection attained under nature. The endemic productions of New Zealand, for instance, are perfect one compared with another; but they are now rapidly yielding before the advancing legions of plants and animals introduced from Europe. Natural selection will not produce absolute perfection, nor do we always meet, as far as we can judge, with this high standard under nature. The correction for the aberration of light is said, on high authority, not to be perfect even in that most perfect organ, the eye. If our reason leads us to admire with enthusiasm a multitude of inimitable contrivances in nature, this same reason tells us, though we may easily err on both sides, that some other contrivances are less perfect. Can we consider the sting of the wasp or of the bee as perfect, which, when used against many attacking animals, cannot be withdrawn, owing to the backward serratures, and so inevitably causes the death of the insect by tearing out its viscera?


On the theory of natural selection we can clearly understand the full meaning of that old canon in natural history, “Natura non facit saltum.” This canon, if we look only to the present inhabitants of the world, is not strictly correct, but if we include all those of past times, it must by my theory be strictly true.


Instincts comparable with habits, but different in their origin. Instincts graduated. Aphides and ants. Instincts variable. Domestic instincts, their origin. Natural instincts of the cuckoo, ostrich, and parasitic bees. Slave-making ants. Hive-bee, its cell-making instinct. Difficulties on the theory of the Natural Selection of instincts. Neuter or sterile insects. Summary.

The subject of instinct might have been worked into the previous chapters; but I have thought that it would be more convenient to treat the subject separately ... . I must premise, that I have nothing to do with the origin of the primary mental powers, any more than I have with that of life itself. We are concerned only with the diversities of instinct and of the other mental qualities of animals within the same class.

I will not attempt any definition of instinct. It would be easy to show that several distinct mental actions are commonly embraced by this term; but every one understands what is meant, when it is said that instinct impels the cuckoo to migrate and to lay her eggs in other birds’ nests. An action, which we ourselves should require experience to enable us to perform, when performed by an animal, more especially by a very young one, without any experience, and when performed by many individuals in the same way, without their knowing for what purpose it is performed, is usually said to be instinctive. But I could show that none of these characters of instinct are universal. A little dose, as Pierre Huber expresses it, of judgment or reason, often comes into play, even in animals very low in the scale of nature.

Frederick Cuvier and several of the older metaphysicians have compared instinct with habit. This comparison gives, I think, a remarkably accurate notion of the frame of mind under which an instinctive action is performed, but not of its origin. How unconsciously many habitual actions are performed, indeed not rarely in direct opposition to our conscious will! yet they may be modified by the will or reason. Habits easily become associated with other habits, and with certain periods of time and states of the body. When once acquired, they often remain constant throughout life. Several other points of resemblance between instincts and habits could be pointed out. As in repeating a well-known song, so in instincts, one action follows another by a sort of rhythm; if a person be interrupted in a song, or in repeating anything by rote, he is generally forced to go back to recover the habitual train of thought: so P. Huber found it was with a caterpillar, which makes a very complicated hammock; for if he took a caterpillar which had completed its hammock up to, say, the sixth stage of construction, and put it into a hammock completed up only to the third stage, the caterpillar simply re-performed the fourth, fifth, and sixth stages of construction. If, however, a caterpillar were taken out of a hammock made up, for instance, to the third stage, and were put into one finished up to the sixth stage, so that much of its work was already done for it, far from feeling the benefit of this, it was much embarrassed, and, in order to complete its hammock, seemed forced to start from the third stage, where it had left off, and thus tried to complete the already finished work.

If we suppose any habitual action to become inherited — and I think it can be shown that this does sometimes happen — then the resemblance between what originally was a habit and an instinct becomes so close as not to be distinguished. If Mozart, instead of playing the pianoforte at three years old with wonderfully little practice, had played a tune with no practice at all, he might truly be said to have done so instinctively. But it would be the most serious error to suppose that the greater number of instincts have been acquired by habit in one generation, and then transmitted by inheritance to succeeding generations. It can be clearly shown that the most wonderful instincts with which we are acquainted, namely, those of the hive-bee and of many ants, could not possibly have been thus acquired.

It will be universally admitted that instincts are as important as corporeal structure for the welfare of each species, under its present conditions of life. Under changed conditions of life, it is at least possible that slight modifications of instinct might be profitable to a species; and if it can be shown that instincts do vary ever so little, then I can see no difficulty in natural selection preserving and continually accumulating variations of instinct to any extent that may be profitable. It is thus, as I believe, that all the most complex and wonderful instincts have originated. As modifications of corporeal structure arise from, and are increased by, use or habit, and are diminished or lost by disuse, so I do not doubt it has been with instincts. But I believe that the effects of habit are of quite subordinate importance to the effects of the natural selection of what may be called accidental variations of instincts; — that is of variations produced by the same unknown causes which produce slight deviations of bodily structure.

No complex instinct can possibly be produced through natural selection, except by the slow and gradual accumulation of numerous, slight, yet profitable, variations. Hence, as in the case of corporeal structures, we ought to find in nature, not the actual transitional gradations by which each complex instinct has been acquired — for these could be found only in the lineal ancestors of each species — but we ought to find in the collateral lines of descent some evidence of such gradations; or we ought at least to be able to show that gradations of some kind are possible; and this we certainly can do. I have been surprised to find ... how very generally gradations, leading to the most complex instincts, can be discovered. The canon of “Natura non facit saltum” applies with almost equal force to instincts as to bodily organs. Changes of instinct may sometimes be facilitated by the same species having different instincts at different periods of life, or at different seasons of the year, or when placed under different circumstances, etc.; in which case either one or the other instinct might be preserved by natural selection. And such instances of diversity of instinct in the same species can be shown to occur in nature.

Again as in the case of corporeal structure, and conformably with my theory, the instinct of each species is good for itself, but has never, as far as we can judge, been produced for the exclusive good of others. One of the strongest instances of an animal apparently performing an action for the sole good of another, with which I am acquainted, is that of aphides voluntarily yielding their sweet excretion to ants: that they do so voluntarily, the following facts show. I removed all the ants from a group of about a dozen aphides on a dock-plant, and prevented their attendance during several hours. After this interval, I felt sure that the aphides would want to excrete. I watched them for some time through a lens, but not one excreted; I then tickled and stroked them with a hair in the same manner, as well as I could, as the ants do with their antennae; but not one excreted. Afterwards I allowed an ant to visit them, and it immediately seemed, by its eager way of running about, to be well aware what a rich flock it had discovered; it then began to play with its antennae on the abdomen first of one aphis and then of another; and each aphis, as soon as it felt the antennae, immediately lifted up its abdomen and excreted a limpid drop of sweet juice, which was eagerly devoured by the ant. Even the quite young aphides behaved in this manner, showing that the action was instinctive, and not the result of experience. But as the excretion is extremely viscid, it is probably a convenience to the aphides to have it removed; and therefore probably the aphides do not instinctively excrete for the sole good of the ants. [Editor’s note: It is now thought that the aphids gain protection from the ants that they feed.] Although I do not believe that any animal in the world performs an action for the exclusive good of another of a distinct species, yet each species tries to take advantage of the instincts of others, as each takes advantage of the weaker bodily structure of others. So again, in some few cases, certain instincts cannot be considered as absolutely perfect; but as details on this and other such points are not indispensable, they may be here passed over.

As some degree of variation in instincts under a state of nature, and the inheritance of such variations, are indispensable for the action of natural selection, as many instances as possible ought to have been here given; but want of space prevents me. I can only assert, that instincts certainly do vary — for instance, the migratory instinct, both in extent and direction, and in its total loss. So it is with the nests of birds, which vary partly in dependence on the situations chosen, and on the nature and temperature of the country inhabited, but often from causes wholly unknown to us: Audubon has given several remarkable cases of differences in nests of the same species in the northern and southern United States. Fear of any particular enemy is certainly an instinctive quality, as may be seen in nestling birds, though it is strengthened by experience, and by the sight of fear of the same enemy in other animals. But fear of man is slowly acquired, as I have elsewhere shown, by various animals inhabiting desert islands; and we may see an instance of this, even in England, in the greater wildness of all our large birds than of our small birds; for the large birds have been most persecuted by man. We may safely attribute the greater wildness of our large birds to this cause; for in uninhabited islands large birds are not more fearful than small; and the magpie, so wary in England, is tame in Norway, as is the hooded crow in Egypt.

That the general disposition of individuals of the same species, born in a state of nature, is extremely diversified, can be shown by a multitude of facts. Several cases also, could be given, of occasional and strange habits in certain species, which might, if advantageous to the species, give rise, through natural selection, to quite new instincts. But I am well aware that these general statements, without facts given in detail, can produce but a feeble effect on the reader’s mind. I can only repeat my assurance, that I do not speak without good evidence.

The possibility, or even probability, of inherited variations of instinct in a state of nature will be strengthened by briefly considering a few cases under domestication. We shall thus also be enabled to see the respective parts which habit and the selection of so-called accidental variations have played in modifying the mental qualities of our domestic animals. .... [L]et us look to the familiar case of the several breeds of dogs: it cannot be doubted that young pointers (I have myself seen a striking instance) will sometimes point and even back other dogs the very first time that they are taken out; retrieving is certainly in some degree inherited by retrievers; and a tendency to run round, instead of at, a flock of sheep, by shepherd-dogs. I cannot see that these actions, performed without experience by the young, and in nearly the same manner by each individual, performed with eager delight by each breed, and without the end being known, — for the young pointer can no more know that he points to aid his master, than the white butterfly knows why she lays her eggs on the leaf of the cabbage, — I cannot see that these actions differ essentially from true instincts. If we were to see one kind of wolf, when young and without any training, as soon as it scented its prey, stand motionless like a statue, and then slowly crawl forward with a peculiar gait; and another kind of wolf rushing round, instead of at, a herd of deer, and driving them to a distant point, we should assuredly call these actions instinctive. Domestic instincts, as they may be called, are certainly far less fixed or invariable than natural instincts; but they have been acted on by far less rigorous selection, and have been transmitted for an incomparably shorter period, under less fixed conditions of life.


We shall, perhaps, best understand how instincts in a state of nature have become modified by selection, by considering a few cases. I will select only three, out of the several which I shall have to discuss in my future work, — namely, the instinct which leads the cuckoo to lay her eggs in other birds’ nests; the slave-making instinct of certain ants; and the comb-making power of the hive-bee: these two latter instincts have generally, and most justly, been ranked by naturalists as the most wonderful of all known instincts.

[Instincts of the cuckoo.—] It is now commonly admitted that the more immediate and final cause of the cuckoo’s instinct is, that she lays her eggs, not daily, but at intervals of two or three days; so that, if she were to make her own nest and sit on her own eggs, those first laid would have to be left for some time unincubated, or there would be eggs and young birds of different ages in the same nest. If this were the case, the process of laying and hatching might be inconveniently long, more especially as she has to migrate at a very early period; and the first hatched young would probably have to be fed by the male alone. But the American cuckoo is in this predicament; for she makes her own nest and has eggs and young successively hatched, all at the same time. .... Now let us suppose that the ancient progenitor of our European cuckoo had the habits of the American cuckoo; but that occasionally she laid an egg in another bird’s nest. If the old bird profited by this occasional habit, or if the young were made more vigorous by advantage having been taken of the mistaken maternal instinct of another bird, than by their own mother’s care, encumbered as she can hardly fail to be by having eggs and young of different ages at the same time; then the old birds or the fostered young would gain an advantage. And analogy would lead me to believe, that the young thus reared would be apt to follow by inheritance the occasional and aberrant habit of their mother, and in their turn would be apt to lay their eggs in other birds’ nests, and thus be successful in rearing their young. By a continued process of this nature, I believe that the strange instinct of our cuckoo could be, and has been, generated. I may add that, according to Dr. Gray and to some other observers, the European cuckoo has not utterly lost all maternal love and care for her own offspring....

Slave-making instinct.— This remarkable instinct was first discovered in the Formica (Polyerges) rufescens by Pierre Huber, a better observer even than his celebrated father. This ant is absolutely dependent on its slaves; without their aid, the species would certainly become extinct in a single year. The males and fertile females do no work. The workers or sterile females, though most energetic and courageous in capturing slaves, do no other work. They are incapable of making their own nests, or of feeding their own larvae. When the old nest is found inconvenient, and they have to migrate, it is the slaves which determine the migration, and actually carry their masters in their jaws. So utterly helpless are the masters, that when Huber shut up thirty of them without a slave, but with plenty of the food which they like best, and with their larvae and pupae to stimulate them to work, they did nothing; they could not even feed themselves, and many perished of hunger. Huber then introduced a single slave (F. fusca), and she instantly set to work, fed and saved the survivors; made some cells and tended the larvae, and put all to rights. What can be more extraordinary than these well-ascertained facts? If we had not known of any other slave-making ant, it would have been hopeless to have speculated how so wonderful an instinct could have been perfected.

Formica sanguinea was likewise first discovered by P. Huber to be a slave-making ant. This species is found in the southern parts of England, and its habits have been attended to by Mr. F. Smith, of the British Museum, to whom I am much indebted for information on this and other subjects. Although fully trusting to the statements of Huber and Mr. Smith, I tried to approach the subject in a sceptical frame of mind, as any one may well be excused for doubting the truth of so extraordinary and odious an instinct as that of making slaves. Hence I will give the observations which I have myself made, in some little detail. I opened fourteen nests of F. sanguinea, and found a few slaves in all. Males and fertile females of the slave-species are found only in their own proper communities, and have never been observed in the nests of F. sanguinea. The slaves are black and not above half the size of their red masters, so that the contrast in their appearance is very great. When the nest is slightly disturbed, the slaves occasionally come out, and like their masters are much agitated and defend the nest: when the nest is much disturbed and the larvae and pupae are exposed, the slaves work energetically with their masters in carrying them away to a place of safety. Hence, it is clear, that the slaves feel quite at home. During the months of June and July, on three successive years, I have watched for many hours several nests in Surrey and Sussex, and never saw a slave either leave or enter a nest. As, during these months, the slaves are very few in number, I thought that they might behave differently when more numerous; but Mr. Smith informs me that he has watched the nests at various hours during May, June and August, both in Surrey and Hampshire, and has never seen the slaves, though present in large numbers in August, either leave or enter the nest. Hence he considers them as strictly household slaves. The masters, on the other hand, may be constantly seen bringing in materials for the nest, and food of all kinds. During the present year, however, in the month of July, I came across a community with an unusually large stock of slaves, and I observed a few slaves mingled with their masters leaving the nest, and marching along the same road to a tall Scotch-fir-tree, twenty-five yards distant, which they ascended together, probably in search of aphides or cocci. According to Huber, ... in Switzerland the slaves habitually work with their masters in making the nest, and they alone open and close the doors in the morning and evening; and, as Huber expressly states, their principal office is to search for aphides. This difference in the usual habits of the masters and slaves in the two countries, probably depends merely on the slaves being captured in greater numbers in Switzerland than in England.

One day I fortunately chanced to witness a migration from one nest to another, and it was a most interesting spectacle to behold the masters carefully carrying, as Huber has described, their slaves in their jaws. [Editor’s comment: Note that this is very different from Formica rufescens, as Darwin will soon discuss.] Another day my attention was struck by about a score of the slave-makers haunting the same spot, and evidently not in search of food; they approached and were vigorously repulsed by an independent community of the slave species (F. fusca); sometimes as many as three of these ants clinging to the legs of the slave-making F. sanguinea. The latter ruthlessly killed their small opponents, and carried their dead bodies as food to their nest, twenty-nine yards distant; but they were prevented from getting any pupae to rear as slaves. I then dug up a small parcel of the pupae of F. fusca from another nest, and put them down on a bare spot near the place of combat; they were eagerly seized, and carried off by the tyrants, who perhaps fancied that, after all, they had been victorious in their late combat.

At the same time I laid on the same place a small parcel of the pupae of another species, F. flava, with a few of these little yellow ants still clinging to the fragments of the nest. This species is sometimes, though rarely, made into slaves, as has been described by Mr. Smith. Although so small a species, it is very courageous, and I have seen it ferociously attack other ants. In one instance I found to my surprise an independent community of F. flava under a stone beneath a nest of the slave-making F. sanguinea; and when I had accidentally disturbed both nests, the little ants attacked their big neighbours with surprising courage. Now I was curious to ascertain whether F. sanguinea could distinguish the pupae of F. fusca, which they habitually make into slaves, from those of the little and furious F. flava, which they rarely capture, and it was evident that they did at once distinguish them: for we have seen that they eagerly and instantly seized the pupae of F. fusca, whereas they were much terrified when they came across the pupae, or even the earth from the nest of F. flava, and quickly ran away; but in about a quarter of an hour, shortly after all the little yellow ants had crawled away, they took heart and carried off the pupae.

One evening I visited another community of F. sanguinea, and found a number of these ants entering their nest, carrying the dead bodies of F. fusca (showing that it was not a migration) and numerous pupae. I traced the returning file burthened with booty, for about forty yards, to a very thick clump of heath, whence I saw the last individual of F. sanguinea emerge, carrying a pupa; but I was not able to find the desolated nest in the thick heath. The nest, however, must have been close at hand, for two or three individuals of F. fusca were rushing about in the greatest agitation, and one was perched motionless with its own pupa in its mouth on the top of a spray of heath over its ravaged home.

Such are the facts, though they did not need confirmation by me, in regard to the wonderful instinct of making slaves. Let it be observed what a contrast the instinctive habits of F. sanguinea present with those of the F. rufescens. The latter does not build its own nest, does not determine its own migrations, does not collect food for itself or its young, and cannot even feed itself: it is absolutely dependent on its numerous slaves. Formica sanguinea, on the other hand, possesses much fewer slaves, and in the early part of the summer extremely few. The masters determine when and where a new nest shall be formed, and when they migrate, the masters carry the slaves. Both in Switzerland and England the slaves seem to have the exclusive care of the larvae, and the masters alone go on slave-making expeditions. In Switzerland the slaves and masters work together, making and bringing materials for the nest: both, but chiefly the slaves, tend, and milk as it may be called, their aphides; and thus both collect food for the community. In England the masters alone usually leave the nest to collect building materials and food for themselves, their slaves and larvae. So that the masters in this country receive much less service from their slaves than they do in Switzerland.

By what steps the instinct of F. sanguinea originated I will not pretend to conjecture. But as ants, which are not slave-makers, will, as I have seen, carry off pupae of other species, if scattered near their nests, it is possible that pupae originally stored as food might become developed; and the ants thus unintentionally reared would then follow their proper instincts, and do what work they could. If their presence proved useful to the species which had seized them — if it were more advantageous to this species to capture workers than to procreate them — the habit of collecting pupae originally for food might by natural selection be strengthened and rendered permanent for the very different purpose of raising slaves. When the instinct was once acquired, if carried out to a much less extent even than in our British F. sanguinea, which, as we have seen, is less aided by its slaves than the same species in Switzerland, I can see no difficulty in natural selection increasing and modifying the instinct — always supposing each modification to be of use to the species — until an ant was formed as abjectly dependent on its slaves as is the Formica rufescens.

Cell-making instinct of the hive-bee.— .... He must be a dull man who can examine the exquisite structure of a comb, so beautifully adapted to its end, without enthusiastic admiration. We hear from mathematicians that bees have practically solved a recondite problem, and have made their cells of the proper shape to hold the greatest possible amount of honey, with the least possible consumption of precious wax in their construction. It has been remarked that a skilful workman, with fitting tools and measures, would find it very difficult to make cells of wax of the true form, though this is perfectly effected by a crowd of bees working in a dark hive. Grant whatever instincts you please, and it seems at first quite inconceivable how they can make all the necessary angles and planes, or even perceive when they are correctly made. But the difficulty is not nearly so great as it at first appears: all this beautiful work can be .... explained by natural selection having taken advantage of numerous, successive, slight modifications of simpler instincts; natural selection having by slow degrees, more and more perfectly, led the bees to sweep equal spheres at a given distance from each other in a double layer, and to build up and excavate the wax along the planes of intersection. The bees, of course, no more knowing that they swept their spheres at one particular distance from each other, than they know what are the several angles of the hexagonal prisms and of the basal rhombic plates. The motive power of the process of natural selection having been economy of wax; that individual swarm which wasted least honey in the secretion of wax, having succeeded best, and having transmitted by inheritance its newly acquired economical instinct to new swarms, which in their turn will have had the best chance of succeeding in the struggle for existence.


No doubt many instincts of very difficult explanation could be opposed to the theory of natural selection, — cases, in which we cannot see how an instinct could possibly have originated; cases, in which no intermediate gradations are known to exist; cases of instinct of apparently such trifling importance, that they could hardly have been acted on by natural selection; cases of instincts almost identically the same in animals so remote in the scale of nature, that we cannot account for their similarity by inheritance from a common parent, and must therefore believe that they have been acquired by independent acts of natural selection. I will not here enter on these several cases, but will confine myself to one special difficulty, which at first appeared to me insuperable, and actually fatal to my whole theory. I allude to the neuters or sterile females in insect-communities: for these neuters often differ widely in instinct and in structure from both the males and fertile females, and yet, from being sterile, they cannot propagate their kind.

The subject well deserves to be discussed at great length, but I will here take only a single case, that of working or sterile ants. How the workers have been rendered sterile is a difficulty; but not much greater than that of any other striking modification of structure; for it can be shown that some insects and other articulate animals in a state of nature occasionally become sterile; and if such insects had been social, and it had been profitable to the community that a number should have been annually born capable of work, but incapable of procreation, I can see no very great difficulty in this being effected by natural selection. But I must pass over this preliminary difficulty. The great difficulty lies in the working ants differing widely from both the males and the fertile females in structure, as in the shape of the thorax and in being destitute of wings and sometimes of eyes, and in instinct. As far as instinct alone is concerned, the prodigious difference in this respect between the workers and the perfect females, would have been far better exemplified by the hive-bee. If a working ant or other neuter insect had been an animal in the ordinary state, I should have unhesitatingly assumed that all its characters had been slowly acquired through natural selection; namely, by an individual having been born with some slight profitable modification of structure, this being inherited by its offspring, which again varied and were again selected, and so onwards. But with the working ant we have an insect differing greatly from its parents, yet absolutely sterile; so that it could never have transmitted successively acquired modifications of structure or instinct to its progeny. It may well be asked how is it possible to reconcile this case with the theory of natural selection?

First, let it be remembered that we have innumerable instances, both in our domestic productions and in those in a state of nature, of all sorts of differences of structure which have become correlated to certain ages, and to either sex. We have differences correlated not only to one sex, but to that short period alone when the reproductive system is active, as in the nuptial plumage of many birds, and in the hooked jaws of the male salmon. We have even slight differences in the horns of different breeds of cattle in relation to an artificially imperfect state of the male sex; for oxen of certain breeds have longer horns than in other breeds, in comparison with the horns of the bulls or cows of these same breeds. Hence I can see no real difficulty in any character having become correlated with the sterile condition of certain members of insect-communities: the difficulty lies in understanding how such correlated modifications of structure could have been slowly accumulated by natural selection.

This difficulty, though appearing insuperable, is lessened, or, as I believe, disappears, when it is remembered that selection may be applied to the family, as well as to the individual, and may thus gain the desired end. Thus, a well-flavoured vegetable is cooked, and the individual is destroyed; but the horticulturist sows seeds of the same stock, and confidently expects to get nearly the same variety; breeders of cattle wish the flesh and fat to be well marbled together; the animal has been slaughtered, but the breeder goes with confidence to the same family. I have such faith in the powers of selection, that I do not doubt that a breed of cattle, always yielding oxen with extraordinarily long horns, could be slowly formed by carefully watching which individual bulls and cows, when matched, produced oxen with the longest horns; and yet no one ox could ever have propagated its kind. Thus I believe it has been with social insects: a slight modification of structure, or instinct, correlated with the sterile condition of certain members of the community, has been advantageous to the community: consequently the fertile males and females of the same community flourished, and transmitted to their fertile offspring a tendency to produce sterile members having the same modification. And I believe that this process has been repeated, until that prodigious amount of difference between the fertile and sterile females of the same species has been produced, which we see in many social insects.

But we have not as yet touched on the climax of the difficulty; namely, the fact that the neuters of several ants differ, not only from the fertile females and males, but from each other, sometimes to an almost incredible degree, and are thus divided into two or even three castes. The castes, moreover, do not generally graduate into each other, but are perfectly well defined; being as distinct from each other, as are any two species of the same genus, or rather as any two genera of the same family. Thus in Eciton, there are working and soldier neuters, with jaws and instincts extraordinarily different: in Cryptocerus, the workers of one caste alone carry a wonderful sort of shield on their heads, the use of which is quite unknown: in the Mexican Myrmecocystus, the workers of one caste never leave the nest; they are fed by the workers of another caste, and they have an enormously developed abdomen which secretes a sort of honey, supplying the place of that excreted by the aphides, or the domestic cattle as they may be called, which our European ants guard or imprison.

It will indeed be thought that I have an overweening confidence in the principle of natural selection, when I do not admit that such wonderful and well-established facts at once annihilate my theory. In the simpler case of neuter insects all of one caste or of the same kind, which have been rendered by natural selection, as I believe to be quite possible, different from the fertile males and females, — in this case, we may safely conclude from the analogy of ordinary variations, that each successive, slight, profitable modification did not probably at first appear in all the individual neuters in the same nest, but in a few alone; and that by the long-continued selection of the fertile parents which produced most neuters with the profitable modification, all the neuters ultimately came to have the desired character. On this view we ought occasionally to find neuter-insects of the same species, in the same nest, presenting gradations of structure; and this we do find, even often, considering how few neuter-insects out of Europe have been carefully examined. .... It often happens that the larger or the smaller sized workers are the most numerous; or that both large and small are numerous, with those of an intermediate size scanty in numbers. Formica flava has larger and smaller workers, with some of intermediate size; and, in this species, as Mr. F. Smith has observed, the larger workers have simple eyes (ocelli), which though small can be plainly distinguished, whereas the smaller workers have their ocelli rudimentary. Having carefully dissected several specimens of these workers, I can affirm that the eyes are far more rudimentary in the smaller workers than can be accounted for merely by their proportionally lesser size; and I fully believe, though I dare not assert so positively, that the workers of intermediate size have their ocelli in an exactly intermediate condition. So that we here have two bodies of sterile workers in the same nest, differing not only in size, but in their organs of vision, yet connected by some few members in an intermediate condition. ....

I may give one other case: so confidently did I expect to find gradations in important points of structure between the different castes of neuters in the same species, that I gladly availed myself of Mr. F. Smith’s offer of numerous specimens from the same nest of the driver ant (Anomma) of West Africa. The reader will perhaps best appreciate the amount of difference in these workers, by my giving not the actual measurements, but a strictly accurate illustration: the difference was the same as if we were to see a set of workmen building a house of whom many were five feet four inches high, and many sixteen feet high; but we must suppose that the larger workmen had heads four instead of three times as big as those of the smaller men, and jaws nearly five times as big. The jaws, moreover, of the working ants of the several sizes differed wonderfully in shape, and in the form and number of the teeth. But the important fact for us is, that though the workers can be grouped into castes of different sizes, yet they graduate insensibly into each other, as does the widely-different structure of their jaws. I speak confidently on this latter point, as Mr. Lubbock made drawings for me with the camera lucida of the jaws which I had dissected from the workers of the several sizes.

With these facts before me, I believe that natural selection, by acting on the fertile parents, could form a species which should regularly produce neuters, either all of large size with one form of jaw, or all of small size with jaws having a widely different structure; or lastly, and this is our climax of difficulty, one set of workers of one size and structure, and simultaneously another set of workers of a different size and structure; — a graduated series having been first formed, as in the case of the driver ant, and then the extreme forms, from being the most useful to the community, having been produced in greater and greater numbers through the natural selection of the parents which generated them; until none with an intermediate structure were produced.

Thus, as I believe, the wonderful fact of two distinctly defined castes of sterile workers existing in the same nest, both widely different from each other and from their parents, has originated. We can see how useful their production may have been to a social community of insects, on the same principle that the division of labour is useful to civilised man. As ants work by inherited instincts and by inherited tools or weapons, and not by acquired knowledge and manufactured instruments, a perfect division of labour could be effected with them only by the workers being sterile; for had they been fertile, they would have intercrossed, and their instincts and structure would have become blended. And nature has, as I believe, effected this admirable division of labour in the communities of ants, by the means of natural selection. But I am bound to confess, that, with all my faith in this principle, I should never have anticipated that natural selection could have been efficient in so high a degree, had not the case of these neuter insects convinced me of the fact. I have, therefore, discussed this case, at some little but wholly insufficient length, in order to show the power of natural selection, and likewise because this is by far the most serious special difficulty, which my theory has encountered. The case, also, is very interesting, as it proves that with animals, as with plants, any amount of modification in structure can be effected by the accumulation of numerous, slight, and as we must call them accidental, variations, which are in any manner profitable, without exercise or habit having come into play. For no amount of exercise, or habit, or volition, in the utterly sterile members of a community could possibly have affected the structure or instincts of the fertile members, which alone leave descendants. I am surprised that no one has advanced this demonstrative case of neuter insects, against the well-known doctrine of Lamarck. [Editor’s note: Lamarck is less well-known today than he was in Darwin’s time. He proposed a theory of evolution according to which characteristics enhanced by exercise were inherited.]


Present distribution cannot be accounted for by differences in physical conditions. Importance of barriers. Affinity of the productions of the same continent. Centres of creation. Means of dispersal, by changes of climate and of the level of the land, and by occasional means. Dispersal during the Glacial period co-extensive with the world.

In considering the distribution of organic beings over the face of the globe, the first great fact which strikes us is, that neither the similarity nor the dissimilarity of the inhabitants of various regions can be accounted for by their climatal and other physical conditions. .... There is hardly a climate or condition in the Old World which cannot be paralleled in the New — at least as closely as the same species generally require; for it is a most rare case to find a group of organisms confined to any small spot, having conditions peculiar in only a slight degree; for instance, small areas in the Old World could be pointed out hotter than any in the New World, yet these are not inhabited by a peculiar fauna or flora. Notwithstanding this parallelism in the conditions of the Old and New Worlds, how widely different are their living productions!

In the southern hemisphere, if we compare large tracts of land in Australia, South Africa, and western South America, between latitudes 25 deg and 35 deg, we shall find parts extremely similar in all their conditions, yet it would not be possible to point out three faunas and floras more utterly dissimilar. Or again we may compare the productions of South America south of lat. 35 deg with those north of 25 deg, which consequently inhabit a considerably different climate, and they will be found incomparably more closely related to each other, than they are to the productions of Australia or Africa under nearly the same climate. Analogous facts could be given with respect to the inhabitants of the sea.

A second great fact which strikes us in our general review is, that barriers of any kind, or obstacles to free migration, are related in a close and important manner to the differences between the productions of various regions. We see this in the great difference of nearly all the terrestrial productions of the New and Old Worlds, excepting in the northern parts, where the land almost joins, and where, under a slightly different climate, there might have been free migration for the northern temperate forms, as there now is for the strictly arctic productions. We see the same fact in the great difference between the inhabitants of Australia, Africa, and South America under the same latitude: for these countries are almost as much isolated from each other as is possible. On each continent, also, we see the same fact; for on the opposite sides of lofty and continuous mountain-ranges, and of great deserts, and sometimes even of large rivers, we find different productions; though as mountain chains, deserts, etc., are not as impassable, or likely to have endured so long as the oceans separating continents, the differences are very inferior in degree to those characteristic of distinct continents.

Turning to the sea, we find the same law. No two marine faunas are more distinct, with hardly a fish, shell, or crab in common, than those of the eastern and western shores of South and Central America; yet these great faunas are separated only by the narrow, but impassable, isthmus of Panama. Westward of the shores of America, a wide space of open ocean extends, with not an island as a halting-place for emigrants; here we have a barrier of another kind, and as soon as this is passed we meet in the eastern islands of the Pacific, with another and totally distinct fauna. So that here three marine faunas range far northward and southward, in parallel lines not far from each other, under corresponding climates; but from being separated from each other by impassable barriers, either of land or open sea, they are wholly distinct. On the other hand, proceeding still further westward from the eastern islands of the tropical parts of the Pacific, we encounter no impassable barriers, and we have innumerable islands as halting-places, until after travelling over a hemisphere we come to the shores of Africa; and over this vast space we meet with no well-defined and distinct marine faunas. Although hardly one shell, crab or fish is common to the above-named three approximate faunas of Eastern and Western America and the eastern Pacific islands, yet many fish range from the Pacific into the Indian Ocean, and many shells are common to the eastern islands of the Pacific and the eastern shores of Africa, on almost exactly opposite meridians of longitude.

A third great fact, partly included in the foregoing statements, is the affinity of the productions of the same continent or sea, though the species themselves are distinct at different points and stations. It is a law of the widest generality, and every continent offers innumerable instances. Nevertheless the naturalist in travelling, for instance, from north to south never fails to be struck by the manner in which successive groups of beings, specifically distinct, yet clearly related, replace each other. He hears from closely allied, yet distinct kinds of birds, notes nearly similar, and sees their nests similarly constructed, but not quite alike, with eggs coloured in nearly the same manner. The plains near the Straits of Magellan are inhabited by one species of Rhea (American ostrich), and northward the plains of La Plata by another species of the same genus; and not by a true ostrich or emeu, like those found in Africa and Australia under the same latitude. On these same plains of La Plata, we see the agouti and bizcacha, animals having nearly the same habits [habitats] as our hares and rabbits ... but they plainly display an American type of structure. We ascend the lofty peaks of the Cordillera and we find an alpine species of bizcacha; we look to the waters, and we do not find the beaver or musk-rat, but the coypu and capybara, rodents of the American type. Innumerable other instances could be given. If we look to the islands off the American shore, however much they may differ in geological structure, the inhabitants, though they may be all peculiar species, are essentially American. ....

This bond, on my theory, is simply inheritance, that cause which alone, as far as we positively know, produces organisms quite like, or, as we see in the case of varieties nearly like each other. The dissimilarity of the inhabitants of different regions may be attributed to modification through natural selection, and in a quite subordinate degree to the direct influence of different physical conditions. The degree of dissimilarity will depend on the migration of the more dominant forms of life from one region into another having been effected with more or less ease, at periods more or less remote; — on the nature and number of the former immigrants; — and on their action and reaction, in their mutual struggles for life; — the relation of organism to organism being, as I have already often remarked, the most important of all relations. Thus the high importance of barriers comes into play by checking migration; as does time for the slow process of modification through natural selection. Widely-ranging species, abounding in individuals, which have already triumphed over many competitors in their own widely-extended homes will have the best chance of seizing on new places, when they spread into new countries. In their new homes they will be exposed to new conditions, and will frequently undergo further modification and improvement; and thus they will become still further victorious, and will produce groups of modified descendants. On this principle of inheritance with modification, we can understand how it is that sections of genera, whole genera, and even families are confined to the same areas, as is so commonly and notoriously the case.


On these views, it is obvious, that the several species of the same genus, though inhabiting the most distant quarters of the world, must originally have proceeded from the same source, as they have descended from the same progenitor. In the case of those species, which have undergone during whole geological periods but little modification, there is not much difficulty in believing that they may have migrated from the same region; for during the vast geographical and climatal changes which will have supervened since ancient times, almost any amount of migration is possible. But in many other cases, in which we have reason to believe that the species of a genus have been produced within comparatively recent times, there is great difficulty on this head. It is also obvious that the individuals of the same species, though now inhabiting distant and isolated regions, must have proceeded from one spot, where their parents were first produced: for, as explained in the last chapter, it is incredible that individuals identically the same should ever have been produced through natural selection from parents specifically distinct.

We are thus brought to the question which has been largely discussed by naturalists, namely, whether species have been created at one or more points of the earth’s surface. Undoubtedly there are very many cases of extreme difficulty, in understanding how the same species could possibly have migrated from some one point to the several distant and isolated points, where now found. Nevertheless the simplicity of the view that each species was first produced within a single region captivates the mind. He who rejects it, rejects the vera causa of ordinary generation with subsequent migration, and calls in the agency of a miracle. It is universally admitted, that in most cases the area inhabited by a species is continuous; and when a plant or animal inhabits two points so distant from each other, or with an interval of such a nature, that the space could not be easily passed over by migration, the fact is given as something remarkable and exceptional. The capacity of migrating across the sea is more distinctly limited in terrestrial mammals, than perhaps in any other organic beings; and, accordingly, we find no inexplicable cases of the same mammal inhabiting distant points of the world. No geologist will feel any difficulty in such cases as Great Britain having been formerly united to Europe, and consequently possessing the same quadrupeds. But if the same species can be produced at two separate points, why do we not find a single mammal common to Europe and Australia or South America? The conditions of life are nearly the same, so that a multitude of European animals and plants have become naturalised in America and Australia; and some of the aboriginal plants are identically the same at these distant points of the northern and southern hemispheres? The answer, as I believe, is, that mammals have not been able to migrate, whereas some plants, from their varied means of dispersal, have migrated across the vast and broken interspace. The great and striking influence which barriers of every kind have had on distribution, is intelligible only on the view that the great majority of species have been produced on one side alone, and have not been able to migrate to the other side. Some few families, many sub-families, very many genera, and a still greater number of sections of genera are confined to a single region; and it has been observed by several naturalists, that the most natural genera, or those genera in which the species are most closely related to each other, are generally local, or confined to one area. What a strange anomaly it would be, if, when coming one step lower in the series, to the individuals of the same species, a directly opposite rule prevailed; and species were not local, but had been produced in two or more distinct areas!

Hence it seems to me, as it has to many other naturalists, that the view of each species having been produced in one area alone, and having subsequently migrated from that area as far as its powers of migration and subsistence under past and present conditions permitted, is the most probable. Undoubtedly many cases occur, in which we cannot explain how the same species could have passed from one point to the other. But the geographical and climatal changes, which have certainly occurred within recent geological times, must have interrupted or rendered discontinuous the formerly continuous range of many species. So that we are reduced to consider whether the exceptions to continuity of range are so numerous and of so grave a nature, that we ought to give up the belief, rendered probable by general considerations, that each species has been produced within one area, and has migrated thence as far as it could. It would be hopelessly tedious to discuss all the exceptional cases of the same species, now living at distant and separated points; nor do I for a moment pretend that any explanation could be offered of many such cases. But after some preliminary remarks, I will discuss a few of the most striking classes of facts; namely, the existence of the same species on the summits of distant mountain-ranges, and at distant points in the arctic and antarctic regions; and secondly ..., the wide distribution of freshwater productions; and thirdly, the occurrence of the same terrestrial species on islands and on the mainland, though separated by hundreds of miles of open sea. If the existence of the same species at distant and isolated points of the earth’s surface, can in many instances be explained on the view of each species having migrated from a single birthplace; then, considering our ignorance with respect to former climatal and geographical changes and various occasional means of transport, the belief that this has been the universal law, seems to me incomparably the safest.

In discussing this subject, we shall be enabled at the same time to consider a point equally important for us, namely, whether the several distinct species of a genus, which on my theory have all descended from a common progenitor, can have migrated (undergoing modification during some part of their migration) from the area inhabited by their progenitor. If it can be shown to be almost invariably the case, that a region, of which most of its inhabitants are closely related to, or belong to the same genera with the species of a second region, has probably received at some former period immigrants from this other region, my theory will be strengthened; for we can clearly understand, on the principle of modification, why the inhabitants of a region should be related to those of another region, whence it has been stocked. A volcanic island, for instance, upheaved and formed at the distance of a few hundreds of miles from a continent, would probably receive from it in the course of time a few colonists, and their descendants, though modified, would still be plainly related by inheritance to the inhabitants of the continent. Cases of this nature are common, and are, as we shall hereafter more fully see, inexplicable on the theory of independent creation. This view of the relation of species in one region to those in another, does not differ much (by substituting the word variety for species) from that lately advanced in an ingenious paper by Mr. Wallace, in which he concludes, that “every species has come into existence coincident both in space and time with a pre-existing closely allied species.” And I now know from correspondence, that this coincidence he attributes to generation with modification.


Before discussing the three classes of facts, which I have selected as presenting the greatest amount of difficulty on the theory of “single centres of creation,” I must say a few words on the means of dispersal.

Means of dispersal.— .... Change of climate must have had a powerful influence on migration: a region when its climate was different may have been a high road for migration, but now be impassable.... Changes of level in the land must also have been highly influential: a narrow isthmus now separates two marine faunas; submerge it, or let it formerly have been submerged, and the two faunas will now blend or may formerly have blended: where the sea now extends, land may at a former period have connected islands or possibly even continents together, and thus have allowed terrestrial productions to pass from one to the other. No geologist will dispute that great mutations of level have occurred within the period of existing organisms. Edward Forbes insisted that all the islands in the Atlantic must recently have been connected with Europe or Africa, and Europe likewise with America. Other authors have thus hypothetically bridged over every ocean, and have united almost every island to some mainland. If indeed the arguments used by Forbes are to be trusted, it must be admitted that scarcely a single island exists which has not recently been united to some continent. This view cuts the Gordian knot of the dispersal of the same species to the most distant points, and removes many a difficulty: but to the best of my judgment we are not authorized in admitting such enormous geographical changes within the period of existing species. It seems to me that we have abundant evidence of great oscillations of level in our continents; but not of such vast changes in their position and extension, as to have united them within the recent period to each other and to the several intervening oceanic islands. I freely admit the former existence of many islands, now buried beneath the sea, which may have served as halting places for plants and for many animals during their migration. In the coral-producing oceans such sunken islands are now marked, as I believe, by rings of coral or atolls standing over them. Whenever it is fully admitted, as I believe it will some day be, that each species has proceeded from a single birthplace, and when in the course of time we know something definite about the means of distribution, we shall be enabled to speculate with security on the former extension of the land....

[Editor’s note: Darwin would have been overjoyed by the modern theory of plate tectonics, which achieved currency 100 years after he was writing. Massive evidence shows that the Earth’s crust is composed of a set of gigantic plates which move relative to each other, bearing the continents aloft. Africa and America were once attached and are drifting apart with the speed with which fingernails grow. India was once attached to Madagascar, before the plate on which it sits migrated north to collide with Asia and raise the Himalayas.]

But I do not believe that it will ever be proved that within the recent period continents which are now quite separate, have been continuously, or almost continuously, united with each other, and with the many existing oceanic islands. Several facts in distribution, — such as the great difference in the marine faunas on the opposite sides of almost every continent, — the close relation of the tertiary inhabitants of several lands and even seas to their present inhabitants, — a certain degree of relation (as we shall hereafter see) between the distribution of mammals and the depth of the sea, — these and other such facts seem to me opposed to the admission of such prodigious geographical revolutions within the recent period, as are necessitated on the view advanced by Forbes and admitted by his many followers. The nature and relative proportions of the inhabitants of oceanic islands likewise seem to me opposed to the belief of their former continuity with continents. Nor does their almost universally volcanic composition favour the admission that they are the wrecks of sunken continents; — if they had originally existed as mountain-ranges on the land, some at least of the islands would have been formed, like other mountain-summits, of granite, metamorphic schists, old fossiliferous or other such rocks, instead of consisting of mere piles of volcanic matter.

I must now say a few words on what are called accidental means, but which more properly might be called occasional means of distribution. I shall here confine myself to plants. In botanical works, this or that plant is stated to be ill adapted for wide dissemination; but for transport across the sea, the greater or less facilities may be said to be almost wholly unknown. Until I tried, with Mr. Berkeley’s aid, a few experiments, it was not even known how far seeds could resist the injurious action of sea-water. To my surprise I found that out of 87 kinds, 64 germinated after an immersion of 28 days, and a few survived an immersion of 137 days. For convenience sake I chiefly tried small seeds, without the capsule or fruit; and as all of these sank in a few days, they could not be floated across wide spaces of the sea, whether or not they were injured by the salt-water. Afterwards I tried some larger fruits, capsules, etc., and some of these floated for a long time. It is well known what a difference there is in the buoyancy of green and seasoned timber; and it occurred to me that floods might wash down plants or branches, and that these might be dried on the banks, and then by a fresh rise in the stream be washed into the sea. Hence I was led to dry stems and branches of 94 plants with ripe fruit, and to place them on sea water. The majority sank quickly, but some which whilst green floated for a very short time, when dried floated much longer; for instance, ripe hazel-nuts sank immediately, but when dried, they floated for 90 days and afterwards when planted they germinated; an asparagus plant with ripe berries floated for 23 days, when dried it floated for 85 days, and the seeds afterwards germinated: the ripe seeds of Helosciadium sank in two days, when dried they floated for above 90 days, and afterwards germinated. Altogether out of the 94 dried plants, 18 floated for above 28 days, and some of the 18 floated for a very much longer period. So that as 64/87 seeds germinated after an immersion of 28 days; and as 18/94 plants with ripe fruit (but not all the same species as in the foregoing experiment) floated, after being dried, for above 28 days, as far as we may infer anything from these scanty facts, we may conclude that the seeds of 14/100 plants of any country might be floated by sea-currents during 28 days, and would retain their power of germination. In Johnston’s Physical Atlas, the average rate of the several Atlantic currents is 33 miles per diem (some currents running at the rate of 60 miles per diem); on this average, the seeds of 14/100 plants belonging to one country might be floated across 924 miles of sea to another country; and when stranded, if blown to a favourable spot by an inland gale, they would germinate.

.... But I do not doubt that plants exposed to the waves would float for a less time than those protected from violent movement as in our experiments. Therefore it would perhaps be safer to assume that the seeds of about 10/100 plants of a flora, after having been dried, could be floated across a space of sea 900 miles in width, and would then germinate. The fact of the larger fruits often floating longer than the small, is interesting; as plants with large seeds or fruit could hardly be transported by any other means; and Alph. de Candolle has shown that such plants generally have restricted ranges.

But seeds may be occasionally transported in another manner. Drift timber is thrown up on most islands, even on those in the midst of the widest oceans; and the natives of the coral-islands in the Pacific, procure stones for their tools, solely from the roots of drifted trees, these stones being a valuable royal tax. I find on examination, that when irregularly shaped stones are embedded in the roots of trees, small parcels of earth are very frequently enclosed in their interstices and behind them, — so perfectly that not a particle could be washed away in the longest transport: out of one small portion of earth thus completely enclosed by wood in an oak about 50 years old, three dicotyledonous plants germinated: I am certain of the accuracy of this observation. Again, I can show that the carcasses of birds, when floating on the sea, sometimes escape being immediately devoured; and seeds of many kinds in the crops of floating birds long retain their vitality: peas and vetches, for instance, are killed by even a few days’ immersion in sea-water; but some taken out of the crop of a pigeon, which had floated on artificial salt-water for 30 days, to my surprise nearly all germinated.

Living birds can hardly fail to be highly effective agents in the transportation of seeds. I could give many facts showing how frequently birds of many kinds are blown by gales to vast distances across the ocean. We may I think safely assume that under such circumstances their rate of flight would often be 35 miles an hour; and some authors have given a far higher estimate. I have never seen an instance of nutritious seeds passing through the intestines of a bird; but hard seeds of fruit will pass uninjured through even the digestive organs of a turkey. In the course of two months, I picked up in my garden 12 kinds of seeds, out of the excrement of small birds, and these seemed perfect, and some of them, which I tried, germinated. But the following fact is more important: the crops of birds do not secrete gastric juice, and do not in the least injure, as I know by trial, the germination of seeds; now after a bird has found and devoured a large supply of food, it is positively asserted that all the grains do not pass into the gizzard for 12 or even 18 hours. A bird in this interval might easily be blown to the distance of 500 miles, and hawks are known to look out for tired birds, and the contents of their torn crops might thus readily get scattered. .... Some hawks and owls bolt their prey whole, and after an interval of from twelve to twenty hours, disgorge pellets, which, as I know from experiments made in the Zoological Gardens, include seeds capable of germination. .... Freshwater fish, I find, eat seeds of many land and water plants: fish are frequently devoured by birds, and thus the seeds might be transported from place to place. I forced many kinds of seeds into the stomachs of dead fish, and then gave their bodies to fishing-eagles, storks, and pelicans; these birds after an interval of many hours, either rejected the seeds in pellets or passed them in their excrement; and several of these seeds retained their power of germination. Certain seeds, however, were always killed by this process.

Although the beaks and feet of birds are generally quite clean, I can show that earth sometimes adheres to them: in one instance I removed twenty-two grains of dry argillaceous earth from one foot of a partridge, and in this earth there was a pebble quite as large as the seed of a vetch. .... Reflect for a moment on the millions of quails which annually cross the Mediterranean; and can we doubt that the earth adhering to their feet would sometimes include a few minute seeds? ....

As icebergs are known to be sometimes loaded with earth and stones, and have even carried brushwood, bones, and the nest of a land-bird, I can hardly doubt that they must occasionally have transported seeds from one part to another of the arctic and antarctic regions ... and during the Glacial period from one part of the now temperate regions to another. In the Azores, from the large number of the species of plants common to Europe, in comparison with the plants of other oceanic islands nearer to the mainland, and ... from the somewhat northern character of the flora in comparison with the latitude, I suspected that these islands had been partly stocked by ice-borne seeds, during the Glacial epoch.... M. Hartung ... had found large fragments of granite and other rocks, which do not occur in the archipelago. Hence we may safely infer that icebergs formerly landed their rocky burthens on the shores of these mid-ocean islands, and it is at least possible that they may have brought thither the seeds of northern plants.

Considering that the several above means of transport, and that several other means, which without doubt remain to be discovered, have been in action year after year, for centuries and tens of thousands of years, it would I think be a marvellous fact if many plants had not thus become widely transported. .... [I]t would be a great error to argue that because a well-stocked island, like Great Britain, has not, as far as is known (and it would be very difficult to prove this), received within the last few centuries, through occasional means of transport, immigrants from Europe or any other continent, that a poorly-stocked island, though standing more remote from the mainland, would not receive colonists by similar means. I do not doubt that out of twenty seeds or animals transported to an island, even if far less well-stocked than Britain, scarcely more than one would be so well fitted to its new home, as to become naturalised. But this, as it seems to me, is no valid argument against what would be effected by occasional means of transport, during the long lapse of geological time, whilst an island was being upheaved and formed, and before it had become fully stocked with inhabitants. On almost bare land, with few or no destructive insects or birds living there, nearly every seed, which chanced to arrive, would be sure to germinate and survive.



Distribution of fresh-water productions. On the inhabitants of oceanic islands. Absence of Batrachians and of terrestrial Mammals. On the relation of the inhabitants of islands to those of the nearest mainland. On colonisation from the nearest source with subsequent modification. Summary of the last and present chapters.

Distribution of fresh-water productions.— As lakes and river-systems are separated from each other by barriers of land, it might have been thought that fresh-water productions would not have ranged widely within the same country, and as the sea is apparently a still more impassable barrier, that they never would have extended to distant countries. But the case is exactly the reverse. Not only have many fresh-water species, belonging to quite different classes, an enormous range, but allied species prevail in a remarkable manner throughout the world. I well remember, when first collecting in the fresh waters of Brazil, feeling much surprise at the similarity of the fresh-water insects, shells, etc., and at the dissimilarity of the surrounding terrestrial beings, compared with those of Britain.

But this power in fresh-water productions of ranging widely, though so unexpected, can, I think, in most cases be explained by their having become fitted, in a manner highly useful to them, for short and frequent migrations from pond to pond, or from stream to stream; and liability to wide dispersal would follow from this capacity as an almost necessary consequence. We can here consider only a few cases. In regard to fish, I believe that the same species never occur in the fresh waters of distant continents. But on the same continent the species often range widely and almost capriciously; for two river-systems will have some fish in common and some different. A few facts seem to favour the possibility of their occasional transport by accidental means; like that of the live fish not rarely dropped by whirlwinds in India, and the vitality of their ova when removed from the water. But I am inclined to attribute the dispersal of fresh-water fish mainly to slight changes within the recent period in the level of the land, having caused rivers to flow into each other. .... The wide difference of the fish on opposite sides of continuous mountain-ranges, which from an early period must have parted river-systems and completely prevented their inosculation, seems to lead to this same conclusion. With respect to allied fresh-water fish occurring at very distant points of the world, no doubt there are many cases which cannot at present be explained: but some fresh-water fish belong to very ancient forms, and in such cases there will have been ample time for great geographical changes, and consequently time and means for much migration. In the second place, salt-water fish can with care be slowly accustomed to live in fresh water; and, according to Valenciennes, there is hardly a single group of fishes confined exclusively to fresh water, so that we may imagine that a marine member of a fresh-water group might travel far along the shores of the sea, and subsequently become modified and adapted to the fresh waters of a distant land.

Some species of fresh-water shells have a very wide range, and allied species, which, on my theory, are descended from a common parent and must have proceeded from a single source, prevail throughout the world. Their distribution at first perplexed me much, as their ova are not likely to be transported by birds, and they are immediately killed by sea water, as are the adults. I could not even understand how some naturalised species have rapidly spread throughout the same country. But two facts, which I have observed — and no doubt many others remain to be observed — throw some light on this subject. When a duck suddenly emerges from a pond covered with duck-weed, I have twice seen these little plants adhering to its back; and it has happened to me, in removing a little duck-weed from one aquarium to another, that I have quite unintentionally stocked the one with fresh-water shells from the other. But another agency is perhaps more effectual: I suspended a duck’s feet, which might represent those of a bird sleeping in a natural pond, in an aquarium, where many ova of fresh-water shells were hatching; and I found that numbers of the extremely minute and just hatched shells crawled on the feet, and clung to them so firmly that when taken out of the water they could not be jarred off, though at a somewhat more advanced age they would voluntarily drop off. These just hatched molluscs, though aquatic in their nature, survived on the duck’s feet, in damp air, from twelve to twenty hours; and in this length of time a duck or heron might fly at least six or seven hundred miles, and would be sure to alight on a pool or rivulet, if blown across sea to an oceanic island or to any other distant point. Sir Charles Lyell also informs me that a Dyticus has been caught with an Ancylus (a fresh-water shell like a limpet) firmly adhering to it; and a water-beetle of the same family, a Colymbetes, once flew on board the ‘Beagle,’ when forty-five miles distant from the nearest land: how much farther it might have flown with a favouring gale no one can tell.

With respect to plants, it has long been known what enormous ranges many fresh-water and even marsh-species have, both over continents and to the most remote oceanic islands. This is strikingly shown, as remarked by Alph. de Candolle, in large groups of terrestrial plants, which have only a very few aquatic members; for these latter seem immediately to acquire, as if in consequence, a very wide range. I think favourable means of dispersal explain this fact. I have before mentioned that earth occasionally, though rarely, adheres in some quantity to the feet and beaks of birds. Wading birds, which frequent the muddy edges of ponds, if suddenly flushed, would be the most likely to have muddy feet. Birds of this order I can show are the greatest wanderers, and are occasionally found on the most remote and barren islands in the open ocean; they would not be likely to alight on the surface of the sea, so that the dirt would not be washed off their feet; when making land, they would be sure to fly to their natural fresh-water haunts. I do not believe that botanists are aware how charged the mud of ponds is with seeds: I have tried several little experiments, but will here give only the most striking case: I took in February three table-spoonfuls of mud from three different points, beneath water, on the edge of a little pond; this mud when dry weighed only 6 3/4 ounces; I kept it covered up in my study for six months, pulling up and counting each plant as it grew; the plants were of many kinds, and were altogether 537 in number; and yet the viscid mud was all contained in a breakfast cup! Considering these facts, I think it would be an inexplicable circumstance if water-birds did not transport the seeds of fresh-water plants to vast distances, and if consequently the range of these plants was not very great. The same agency may have come into play with the eggs of some of the smaller fresh-water animals.

Other and unknown agencies probably have also played a part.... [F]resh-water fish eat some kinds of seeds ... Herons and other birds, century after century, have gone on daily devouring fish; they then take flight and go to other waters, or are blown across the sea; and we have seen that seeds retain their power of germination, when rejected in pellets or in excrement, many hours afterwards....

In considering these several means of distribution, it should be remembered that when a pond or stream is first formed, for instance, on a rising islet, it will be unoccupied; and a single seed or egg will have a good chance of succeeding. Although there will always be a struggle for life between the individuals of the species, however few, already occupying any pond, yet as the number of kinds is small, compared with those on the land, the competition will probably be less severe between aquatic than between terrestrial species; consequently an intruder from the waters of a foreign country, would have a better chance of seizing on a place, than in the case of terrestrial colonists. .... We should not forget the probability of many species having formerly ranged as continuously as fresh-water productions ever can range, over immense areas, and having subsequently become extinct in intermediate regions. But the wide distribution of fresh-water plants and of the lower animals, whether retaining the same identical form or in some degree modified, I believe mainly depends on the wide dispersal of their seeds and eggs by animals, more especially by fresh-water birds, which have large powers of flight, and naturally travel from one to another and often distant piece of water. Nature, like a careful gardener, thus takes her seeds from a bed of a particular nature, and drops them in another equally well fitted for them.

On the inhabitant of oceanic islands.— We now come to the last of the three classes of facts, which I have selected as presenting the greatest amount of difficulty, on the view that all the individuals both of the same and of allied species have descended from a single parent; and therefore have all proceeded from a common birthplace, notwithstanding that in the course of time they have come to inhabit distant points of the globe. I have already stated that I cannot honestly admit Forbes’s view on continental extensions, which, if legitimately followed out, would lead to the belief that within the recent period all existing islands have been nearly or quite joined to some continent. This view would remove many difficulties, but it would not, I think, explain all the facts in regard to insular productions. In the following remarks I shall not confine myself to the mere question of dispersal; but shall consider some other facts, which bear on the truth of the two theories of independent creation and of descent with modification.

The species of all kinds which inhabit oceanic islands are few in number compared with those on equal continental areas: Alph. de Candolle admits this for plants, and Wollaston for insects. If we look to the large size and varied stations of New Zealand, extending over 780 miles of latitude, and compare its flowering plants, only 750 in number, with those on an equal area at the Cape of Good Hope or in Australia, we must, I think, admit that something quite independently of any difference in physical conditions has caused so great a difference in number. Even the uniform county of Cambridge has 847 plants, and the little island of Anglesea 764, but a few ferns and a few introduced plants are included in these numbers, and the comparison in some other respects is not quite fair. We have evidence that the barren island of Ascension aboriginally possessed under half-a-dozen flowering plants; yet many have become naturalised on it, as they have on New Zealand and on every other oceanic island which can be named. In St. Helena there is reason to believe that the naturalised plants and animals have nearly or quite exterminated many native productions. He who admits the doctrine of the creation of each separate species, will have to admit, that a sufficient number of the best adapted plants and animals have not been created on oceanic islands; for man has unintentionally stocked them from various sources far more fully and perfectly than has nature.

Although in oceanic islands the number of kinds of inhabitants is scanty, the proportion of endemic species (i.e. those found nowhere else in the world) is often extremely large. If we compare, for instance, the number of the endemic land-shells in Madeira, or of the endemic birds in the Galapagos Archipelago, with the number found on any continent, and then compare the area of the islands with that of the continent, we shall see that this is true. This fact might have been expected on my theory, for, as already explained, species occasionally arriving after long intervals in a new and isolated district, and having to compete with new associates, will be eminently liable to modification, and will often produce groups of modified descendants. But it by no means follows, that, because in an island nearly all the species of one class are peculiar, those of another class, or of another section of the same class, are peculiar; and this difference seems to depend on the species which do not become modified having immigrated with facility and in a body, so that their mutual relations have not been much disturbed. Thus in the Galapagos Islands nearly every land-bird, but only two out of the eleven marine birds, are peculiar; and it is obvious that marine birds could arrive at these islands more easily than land-birds. Bermuda, on the other hand, which lies at about the same distance from North America as the Galapagos Islands do from South America, and which has a very peculiar soil, does not possess one endemic land bird; and we know from Mr. J. M. Jones’s admirable account of Bermuda, that very many North American birds, during their great annual migrations, visit either periodically or occasionally this island. Madeira does not possess one peculiar bird, and many European and African birds are almost every year blown there, as I am informed by Mr. E. V. Harcourt. So that these two islands of Bermuda and Madeira have been stocked by birds, which for long ages have struggled together in their former homes, and have become mutually adapted to each other; and when settled in their new homes, each kind will have been kept by the others to their proper places and habits, and will consequently have been little liable to modification.....

Oceanic islands are sometimes deficient in certain classes, and their places are apparently occupied by the other inhabitants; in the Galapagos Islands reptiles, and in New Zealand gigantic wingless birds, take the place of mammals. In the plants of the Galapagos Islands, Dr. Hooker has shown that the proportional numbers of the different orders are very different from what they are elsewhere. Such cases are generally accounted for by the physical conditions of the islands; but this explanation seems to me not a little doubtful. Facility of immigration, I believe, has been at least as important as the nature of the conditions.


With respect to the absence of whole orders on oceanic islands, Bory St. Vincent long ago remarked that Batrachians (frogs, toads, newts) have never been found on any of the many islands with which the great oceans are studded. I have taken pains to verify this assertion, and I have found it strictly true. I have, however, been assured that a frog exists on the mountains of the great island of New Zealand; but I suspect that this exception (if the information be correct) may be explained through glacial agency. This general absence of frogs, toads, and newts on so many oceanic islands cannot be accounted for by their physical conditions; indeed it seems that islands are peculiarly well fitted for these animals; for frogs have been introduced into Madeira, the Azores, and Mauritius, and have multiplied so as to become a nuisance. But as these animals and their spawn are known to be immediately killed by sea-water, on my view we can see that there would be great difficulty in their transportal across the sea, and therefore why they do not exist on any oceanic island. But why, on the theory of creation, they should not have been created there, it would be very difficult to explain.

Mammals offer another and similar case. I have carefully searched the oldest voyages, but have not finished my search; as yet I have not found a single instance, free from doubt, of a terrestrial mammal (excluding domesticated animals kept by the natives) inhabiting an island situated above 300 miles from a continent or great continental island; and many islands situated at a much less distance are equally barren. The Falkland Islands, which are inhabited by a wolf-like fox, come nearest to an exception; but this group cannot be considered as oceanic, as it lies on a bank connected with the mainland; moreover, icebergs formerly brought boulders to its western shores, and they may have formerly transported foxes, as so frequently now happens in the arctic regions. Yet it cannot be said that small islands will not support small mammals, for they occur in many parts of the world on very small islands, if close to a continent; and hardly an island can be named on which our smaller quadrupeds have not become naturalised and greatly multiplied....

Besides the absence of terrestrial mammals in relation to the remoteness of islands from continents, there is also a relation, to a certain extent independent of distance, between the depth of the sea separating an island from the neighbouring mainland, and the presence in both of the same mammiferous species or of allied species in a more or less modified condition. Mr. Windsor Earl has made some striking observations on this head in regard to the great Malay Archipelago, which is traversed near Celebes by a space of deep ocean; and this space separates two widely distinct mammalian faunas. On either side the islands are situated on moderately deep submarine banks, and they are inhabited by closely allied or identical quadrupeds. No doubt some few anomalies occur in this great archipelago, and there is much difficulty in forming a judgment in some cases owing to the probable naturalisation of certain mammals through man’s agency; but we shall soon have much light thrown on the natural history of this archipelago by the admirable zeal and researches of Mr. Wallace. I have not as yet had time to follow up this subject in all other quarters of the world; but as far as I have gone, the relation generally holds good. We see Britain separated by a shallow channel from Europe, and the mammals are the same on both sides; we meet with analogous facts on many islands separated by similar channels from Australia. The West Indian Islands stand on a deeply submerged bank, nearly 1000 fathoms in depth, and here we find American forms, but the species and even the genera are distinct. As the amount of modification in all cases depends to a certain degree on the lapse of time, and as during changes of level it is obvious that islands separated by shallow channels are more likely to have been continuously united within a recent period to the mainland than islands separated by deeper channels, we can understand the frequent relation between the depth of the sea and the degree of affinity of the mammalian inhabitants of islands with those of a neighbouring continent, — an inexplicable relation on the view of independent acts of creation.

All the foregoing remarks on the inhabitants of oceanic islands, — namely, the scarcity of kinds — the richness in endemic forms in particular classes or sections of classes, — the absence of whole groups, as of batrachians, and of terrestrial mammals notwithstanding the presence of aerial bats, — the singular proportions of certain orders of plants, — herbaceous forms having been developed into trees, etc., — seem to me to accord better with the view of occasional means of transport having been largely efficient in the long course of time, than with the view of all our oceanic islands having been formerly connected by continuous land with the nearest continent; for on this latter view the migration would probably have been more complete; and if modification be admitted, all the forms of life would have been more equally modified, in accordance with the paramount importance of the relation of organism to organism.

I do not deny that there are many and grave difficulties in understanding how several of the inhabitants of the more remote islands, whether still retaining the same specific form or modified since their arrival, could have reached their present homes. But the probability of many islands having existed as halting-places, of which not a wreck now remains, must not be overlooked.....

The most striking and important fact for us in regard to the inhabitants of islands, is their affinity to those of the nearest mainland, without being actually the same species. Numerous instances could be given of this fact. I will give only one, that of the Galapagos Archipelago, situated under the equator, between 500 and 600 miles from the shores of South America. Here almost every product of the land and water bears the unmistakeable stamp of the American continent. There are twenty-six land birds, and twenty-five of these are ranked by Mr. Gould as distinct species, supposed to have been created here; yet the close affinity of most of these birds to American species in every character, in their habits, gestures, and tones of voice, was manifest. So it is with the other animals, and with nearly all the plants, as shown by Dr. Hooker in his admirable memoir on the Flora of this archipelago. The naturalist, looking at the inhabitants of these volcanic islands in the Pacific, distant several hundred miles from the continent, yet feels that he is standing on American land. Why should this be so? why should the species which are supposed to have been created in the Galapagos Archipelago, and nowhere else, bear so plain a stamp of affinity to those created in America? There is nothing in the conditions of life, in the geological nature of the islands, in their height or climate, or in the proportions in which the several classes are associated together, which resembles closely the conditions of the South American coast: in fact there is a considerable dissimilarity in all these respects. On the other hand, there is a considerable degree of resemblance in the volcanic nature of the soil, in climate, height, and size of the islands, between the Galapagos and Cape de Verde Archipelagos: but what an entire and absolute difference in their inhabitants! The inhabitants of the Cape de Verde Islands are related to those of Africa, like those of the Galapagos to America. I believe this grand fact can receive no sort of explanation on the ordinary view of independent creation; whereas on the view here maintained, it is obvious that the Galapagos Islands would be likely to receive colonists, whether by occasional means of transport or by formerly continuous land, from America; and the Cape de Verde Islands from Africa; and that such colonists would be liable to modification; — the principle of inheritance still betraying their original birthplace.


The law which causes the inhabitants of an archipelago, though specifically distinct, to be closely allied to those of the nearest continent, we sometimes see displayed on a small scale, yet in a most interesting manner, within the limits of the same archipelago. Thus the several islands of the Galapagos Archipelago are tenanted, as I have elsewhere shown, in a quite marvellous manner, by very closely related species; so that the inhabitants of each separate island, though mostly distinct, are related in an incomparably closer degree to each other than to the inhabitants of any other part of the world. And this is just what might have been expected on my view, for the islands are situated so near each other that they would almost certainly receive immigrants from the same original source, or from each other. But this dissimilarity between the endemic inhabitants of the islands may be used as an argument against my views; for it may be asked, how has it happened in the several islands situated within sight of each other, having the same geological nature, the same height, climate, etc., that many of the immigrants should have been differently modified, though only in a small degree. This long appeared to me a great difficulty: but it arises in chief part from the deeply-seated error of considering the physical conditions of a country as the most important for its inhabitants; whereas it cannot, I think, be disputed that the nature of the other inhabitants, with which each has to compete, is at least as important, and generally a far more important element of success. Now if we look to those inhabitants of the Galapagos Archipelago which are found in other parts of the world (laying on one side for the moment the endemic species, which cannot be here fairly included, as we are considering how they have come to be modified since their arrival), we find a considerable amount of difference in the several islands. This difference might indeed have been expected on the view of the islands having been stocked by occasional means of transport — a seed, for instance, of one plant having been brought to one island, and that of another plant to another island. Hence when in former times an immigrant settled on any one or more of the islands, or when it subsequently spread from one island to another, it would undoubtedly be exposed to different conditions of life in the different islands, for it would have to compete with different sets of organisms: a plant, for instance, would find the best-fitted ground more perfectly occupied by distinct plants in one island than in another, and it would be exposed to the attacks of somewhat different enemies. If then it varied, natural selection would probably favour different varieties in the different islands. Some species, however, might spread and yet retain the same character throughout the group, just as we see on continents some species spreading widely and remaining the same.

The really surprising fact in this case of the Galapagos Archipelago, and in a lesser degree in some analogous instances, is that the new species formed in the separate islands have not quickly spread to the other islands. But the islands, though in sight of each other, are separated by deep arms of the sea, in most cases wider than the British Channel, and there is no reason to suppose that they have at any former period been continuously united. The currents of the sea are rapid and sweep across the archipelago, and gales of wind are extraordinarily rare; so that the islands are far more effectually separated from each other than they appear to be on a map. Nevertheless a good many species, both those found in other parts of the world and those confined to the archipelago, are common to the several islands, and we may infer from certain facts that these have probably spread from some one island to the others. But we often take, I think, an erroneous view of the probability of closely allied species invading each other’s territory, when put into free intercommunication. Undoubtedly if one species has any advantage whatever over another, it will in a very brief time wholly or in part supplant it; but if both are equally well fitted for their own places in nature, both probably will hold their own places and keep separate for almost any length of time. .... In the Galapagos Archipelago, many even of the birds, though so well adapted for flying from island to island, are distinct on each; thus there are three closely-allied species of mocking-thrush, each confined to its own island. Now let us suppose the mocking-thrush of Chatham Island to be blown to Charles Island, which has its own mocking-thrush: why should it succeed in establishing itself there? We may safely infer that Charles Island is well stocked with its own species, for annually more eggs are laid there than can possibly be reared; and we may infer that the mocking-thrush peculiar to Charles Island is at least as well fitted for its home as is the species peculiar to Chatham Island. Sir C. Lyell and Mr. Wollaston have communicated to me a remarkable fact bearing on this subject; namely, that Madeira and the adjoining islet of Porto Santo possess many distinct but representative land-shells, some of which live in crevices of stone; and although large quantities of stone are annually transported from Porto Santo to Madeira, yet this latter island has not become colonised by the Porto Santo species: nevertheless both islands have been colonised by some European land-shells, which no doubt had some advantage over the indigenous species. From these considerations I think we need not greatly marvel at the endemic and representative species, which inhabit the several islands of the Galapagos Archipelago, not having universally spread from island to island. In many other instances, as in the several districts of the same continent, pre-occupation has probably played an important part in checking the commingling of species under the same conditions of life. Thus, the south-east and south-west corners of Australia have nearly the same physical conditions, and are united by continuous land, yet they are inhabited by a vast number of distinct mammals, birds, and plants.

The principle which determines the general character of the fauna and flora of oceanic islands, namely, that the inhabitants, when not identically the same, yet are plainly related to the inhabitants of that region whence colonists could most readily have been derived, — the colonists having been subsequently modified and better fitted to their new homes, — is of the widest application throughout nature. We see this on every mountain, in every lake and marsh. For Alpine species, excepting in so far as the same forms, chiefly of plants, have spread widely throughout the world during the recent Glacial epoch, are related to those of the surrounding lowlands; — thus we have in South America, Alpine humming-birds, Alpine rodents, Alpine plants, etc., all of strictly American forms, and it is obvious that a mountain, as it became slowly upheaved, would naturally be colonised from the surrounding lowlands. So it is with the inhabitants of lakes and marshes, excepting in so far as great facility of transport has given the same general forms to the whole world. We see this same principle in the blind animals inhabiting the caves of America and of Europe. Other analogous facts could be given. And it will, I believe, be universally found to be true, that wherever in two regions, let them be ever so distant, many closely allied or representative species occur, there will likewise be found some identical species, showing, in accordance with the foregoing view, that at some former period there has been intercommunication or migration between the two regions. And wherever many closely-allied species occur, there will be found many forms which some naturalists rank as distinct species, and some as varieties; these doubtful forms showing us the steps in the process of modification.

This relation between the power and extent of migration of a species, either at the present time or at some former period under different physical conditions, and the existence at remote points of the world of other species allied to it, is shown in another and more general way. Mr. Gould remarked to me long ago, that in those genera of birds which range over the world, many of the species have very wide ranges. I can hardly doubt that this rule is generally true, though it would be difficult to prove it. Amongst mammals, we see it strikingly displayed in Bats, and in a lesser degree in the Felidae and Canidae. We see it, if we compare the distribution of butterflies and beetles. So it is with most fresh-water productions, in which so many genera range over the world, and many individual species have enormous ranges. It is not meant that in world-ranging genera all the species have a wide range, or even that they have on an AVERAGE a wide range; but only that some of the species range very widely; for the facility with which widely-ranging species vary and give rise to new forms will largely determine their average range. For instance, two varieties of the same species inhabit America and Europe, and the species thus has an immense range; but, if the variation had been a little greater, the two varieties would have been ranked as distinct species, and the common range would have been greatly reduced. Still less is it meant, that a species which apparently has the capacity of crossing barriers and ranging widely, as in the case of certain powerfully-winged birds, will necessarily range widely; for we should never forget that to range widely implies not only the power of crossing barriers, but the more important power of being victorious in distant lands in the struggle for life with foreign associates. But on the view of all the species of a genus having descended from a single parent, though now distributed to the most remote points of the world, we ought to find, and I believe as a general rule we do find, that some at least of the species range very widely; for it is necessary that the unmodified parent should range widely, undergoing modification during its diffusion, and should place itself under diverse conditions favourable for the conversion of its offspring, firstly into new varieties and ultimately into new species....


Classification, groups subordinate to groups. Natural system. Rules and difficulties in classification, explained on the theory of descent with modification. Classification of varieties. Descent always used in classification. Analogical or adaptive characters. Affinities, general, complex and radiating. Extinction separates and defines groups. Morphology, between members of the same class, between parts of the same individual. Embryology, laws of, explained by variations not supervening at an early age, and being inherited at a corresponding age. Rudimentary organs; their origin explained. Summary.

From the first dawn of life, all organic beings are found to resemble each other in descending degrees, so that they can be classed in groups under groups. This classification is evidently not arbitrary like the grouping of the stars in constellations. The existence of groups would have been of simple signification, if one group had been exclusively fitted to inhabit the land, and another the water; one to feed on flesh, another on vegetable matter, and so on; but the case is widely different in nature; for it is notorious how commonly members of even the same subgroup have different habits. ...  I have attempted to show that it is the widely ranging, the much diffused and common, that is the dominant species belonging to the larger genera, which vary most. The varieties, or incipient species, thus produced ultimately become converted, as I believe, into new and distinct species; and these, on the principle of inheritance, tend to produce other new and dominant species. Consequently the groups which are now large, and which generally include many dominant species, tend to go on increasing indefinitely in size. I further attempted to show that from the varying descendants of each species trying to occupy as many and as different places as possible in the economy of nature, there is a constant tendency in their characters to diverge. This conclusion was supported by looking at the great diversity of the forms of life which, in any small area, come into the closest competition, and by looking to certain facts in naturalisation.


Naturalists try to arrange the species, genera, and families in each class, on what is called the Natural System. But what is meant by this system? Some authors look at it merely as a scheme for arranging together those living objects which are most alike, and for separating those which are most unlike; or as an artificial means for enunciating, as briefly as possible, general propositions, — that is, by one sentence to give the characters common, for instance, to all mammals, by another those common to all carnivora, by another those common to the dog-genus, and then by adding a single sentence, a full description is given of each kind of dog. The ingenuity and utility of this system are indisputable. But many naturalists think that something more is meant by the Natural System; they believe that it reveals the plan of the Creator; but unless it be specified whether order in time or space, or what else is meant by the plan of the Creator, it seems to me that nothing is thus added to our knowledge. Such expressions as that famous one of Linnaeus, and which we often meet with in a more or less concealed form, that the characters do not make the genus, but that the genus gives the characters, seem to imply that something more is included in our classification, than mere resemblance. I believe that something more is included; and that propinquity of descent, — the only known cause of the similarity of organic beings, — is the bond, hidden as it is by various degrees of modification, which is partially revealed to us by our classifications.

Let us now consider the rules followed in classification, and the difficulties which are encountered on the view that classification either gives some unknown plan of creation, or is simply a scheme for enunciating general propositions and of placing together the forms most like each other. It might have been thought (and was in ancient times thought) that those parts of the structure which determined the habits of life, and the general place of each being in the economy of nature, would be of very high importance in classification. Nothing can be more false. No one regards the external similarity of a mouse to a shrew, of a dugong to a whale, of a whale to a fish, as of any importance. These resemblances, though so intimately connected with the whole life of the being, are ranked as merely “adaptive or analogical characters;” but to the consideration of these resemblances we shall have to recur. It may even be given as a general rule, that the less any part of the organisation is concerned with special habits, the more important it becomes for classification. As an instance: Owen, in speaking of the dugong, says, “The generative organs being those which are most remotely related to the habits and food of an animal, I have always regarded as affording very clear indications of its true affinities. We are least likely in the modifications of these organs to mistake a merely adaptive for an essential character.” So with plants, how remarkable it is that the organs of vegetation, on which their whole life depends, are of little signification, excepting in the first main divisions; whereas the organs of reproduction, with their product the seed, are of paramount importance!


Again, no one will say that rudimentary or atrophied organs are of high physiological or vital importance; yet, undoubtedly, organs in this condition are often of high value in classification. No one will dispute that the rudimentary teeth in the upper jaws of young ruminants, and certain rudimentary bones of the leg, are highly serviceable in exhibiting the close affinity between Ruminants and Pachyderms....

Numerous instances could be given of characters derived from parts which must be considered of very trifling physiological importance, but which are universally admitted as highly serviceable in the definition of whole groups. For instance, whether or not there is an open passage from the nostrils to the mouth, the only character, according to Owen, which absolutely distinguishes fishes and reptiles — … — the manner in which the wings of insects are folded — mere colour in certain Algae — … — the nature of the dermal covering, as hair or feathers, in the Vertebrata. If the Ornithorhynchus [platypus] had been covered with feathers instead of hair, this external and trifling character would, I think, have been considered by naturalists as important an aid in determining the degree of affinity of this strange creature to birds and reptiles, as an approach in structure in any one internal and important organ.


We can see why characters derived from the embryo should be of equal importance with those derived from the adult, for our classifications of course include all ages of each species. But it is by no means obvious, on the ordinary view, why the structure of the embryo should be more important for this purpose than that of the adult, which alone plays its full part in the economy of nature. Yet it has been strongly urged by those great naturalists, Milne Edwards and Agassiz, that embryonic characters are the most important of any in the classification of animals; and this doctrine has very generally been admitted as true. The same fact holds good with flowering plants, of which the two main divisions have been founded on characters derived from the embryo, — on the number and position of the embryonic leaves or cotyledons, and on the mode of development of the plumule and radicle. In our discussion on embryology, we shall see why such characters are so valuable, on the view of classification tacitly including the idea of descent.

Our classifications are often plainly influenced by chains of affinities. Nothing can be easier than to define a number of characters common to all birds; but in the case of crustaceans, such definition has hitherto been found impossible. There are crustaceans at the opposite ends of the series, which have hardly a character in common; yet the species at both ends, from being plainly allied to others, and these to others, and so onwards, can be recognised as unequivocally belonging to this, and to no other class of the Articulata.


Finally, with respect to the comparative value of the various groups of species, such as orders, sub-orders, families, sub-families, and genera, they seem to be, at least at present, almost arbitrary. Several of the best botanists, such as Mr. Bentham and others, have strongly insisted on their arbitrary value. Instances could be given amongst plants and insects, of a group of forms, first ranked by practised naturalists as only a genus, and then raised to the rank of a sub-family or family; and this has been done, not because further research has detected important structural differences, at first overlooked, but because numerous allied species, with slightly different grades of difference, have been subsequently discovered.

All the foregoing rules and aids and difficulties in classification are explained, if I do not greatly deceive myself, on the view that the natural system is founded on descent with modification; that the characters which naturalists consider as showing true affinity between any two or more species, are those which have been inherited from a common parent, and, in so far, all true classification is genealogical; that community of descent is the hidden bond which naturalists have been unconsciously seeking, and not some unknown plan of creation, or the enunciation of general propositions, and the mere putting together and separating objects more or less alike.


It may be worth while to illustrate this view of classification, by taking the case of languages. If we possessed a perfect pedigree of mankind, a genealogical arrangement of the races of man would afford the best classification of the various languages now spoken throughout the world; and if all extinct languages, and all intermediate and slowly changing dialects, had to be included, such an arrangement would, I think, be the only possible one. Yet it might be that some very ancient language had altered little, and had given rise to few new languages, whilst others … had altered much, and had given rise to many new languages and dialects. The various degrees of difference in the languages from the same stock, would have to be expressed by groups subordinate to groups; but the proper or even only possible arrangement would still be genealogical; and this would be strictly natural, as it would connect together all languages, extinct and modern, by the closest affinities, and would give the filiation and origin of each tongue.


We can understand why a species or a group of species may depart, in several of its most important characteristics, from its allies, and yet be safely classed with them. This may be safely done, and is often done, as long as a sufficient number of characters, let them be ever so unimportant, betrays the hidden bond of community of descent. Let two forms have not a single character in common, yet if these extreme forms are connected together by a chain of intermediate groups, we may at once infer their community of descent, and we put them all into the same class. ….

We can understand, on these views, the very important distinction between real affinities and analogical or adaptive resemblances. Lamarck first called attention to this distinction, and he has been ably followed by Macleay and others. The resemblance, in the shape of the body and in the fin-like anterior limbs, between the dugong, which is a pachydermatous animal, and the whale, and between both these mammals and fishes, is analogical. … We see something of the same kind even in our domestic varieties, as in the thickened stems of the common and Swedish turnip. The resemblance of the greyhound and racehorse is hardly more fanciful than the analogies which have been drawn by some authors between very distinct animals. On my view of characters being of real importance for classification, only in so far as they reveal descent, we can clearly understand why analogical or adaptive character, although of the utmost importance to the welfare of the being, are almost valueless to the systematist. For animals, belonging to two most distinct lines of descent, may readily become adapted to similar conditions, and thus assume a close external resemblance; but such resemblances will not reveal — will rather tend to conceal their blood-relationship to their proper lines of descent. We can also understand the apparent paradox, that the very same characters are analogical when one class or order is compared with another, but give true affinities when the members of the same class or order are compared one with another: thus the shape of the body and fin-like limbs are only analogical when whales are compared with fishes, being adaptations in both classes for swimming through the water; but the shape of the body and fin-like limbs serve as characters exhibiting true affinity between the several members of the whale family; for these cetaceans agree in so many characters, great and small, that we cannot doubt that they have inherited their general shape of body and structure of limbs from a common ancestor. So it is with fishes.


On the principle of the multiplication and gradual divergence in character of the species descended from a common parent, together with their retention by inheritance of some characters in common, we can understand the excessively complex and radiating affinities by which all the members of the same family or higher group are connected together. For the common parent of a whole family of species, now broken up by extinction into distinct groups and sub-groups, will have transmitted some of its characters, modified in various ways and degrees, to all; and the several species will consequently be related to each other by circuitous lines of affinity of various lengths … mounting up through many predecessors. As it is difficult to show the blood-relationship between the numerous kindred of any ancient and noble family, even by the aid of a genealogical tree, and almost impossible to do this without this aid, we can understand the extraordinary difficulty which naturalists have experienced in describing, without the aid of a diagram, the various affinities which they perceive between the many living and extinct members of the same great natural class.

Extinction, as we have seen … has played an important part in defining and widening the intervals between the several groups in each class. We may thus account even for the distinctness of whole classes from each other — for instance, of birds from all other vertebrate animals — by the belief that many ancient forms of life have been utterly lost, through which the early progenitors of birds were formerly connected with the early progenitors of the other vertebrate classes. There has been less entire extinction of the forms of life which once connected fishes with batrachians. There has been still less in some other classes, as in that of the Crustacea, for here the most wonderfully diverse forms are still tied together by a long, but broken, chain of affinities. Extinction has only separated groups: it has by no means made them; for if every form which has ever lived on this earth were suddenly to reappear, though it would be quite impossible to give definitions by which each group could be distinguished from other groups, as all would blend together by steps as fine as those between the finest existing varieties, nevertheless a natural classification, or at least a natural arrangement, would be possible. ….

Finally, we have seen that natural selection, which results from the struggle for existence, and which almost inevitably induces extinction and divergence of character in the many descendants from one dominant parent-species, explains that great and universal feature in the affinities of all organic beings, namely, their subordination in group under group. … I believe this element of descent is the hidden bond of connexion which naturalists have sought under the term of the Natural System. On this idea of the natural system being, in so far as it has been perfected, genealogical in its arrangement, with the grades of difference between the descendants from a common parent, expressed by the terms genera, families, orders, etc., we can understand the rules which we are compelled to follow in our classification. We can understand why we value certain resemblances far more than others; why we are permitted to use rudimentary and useless organs, or others of trifling physiological importance; why, in comparing one group with a distinct group, we summarily reject analogical or adaptive characters, and yet use these same characters within the limits of the same group. We can clearly see how it is that all living and extinct forms can be grouped together in one great system; and how the several members of each class are connected together by the most complex and radiating lines of affinities. We shall never, probably, disentangle the inextricable web of affinities between the members of any one class; but when we have a distinct object in view, and do not look to some unknown plan of creation, we may hope to make sure but slow progress.

Morphology.— We have seen that the members of the same class, independently of their habits of life, resemble each other in the general plan of their organisation. This resemblance is often expressed by the term “unity of type;” or by saying that the several parts and organs in the different species of the class are homologous. The whole subject is included under the general name of Morphology. This is the most interesting department of natural history, and may be said to be its very soul. What can be more curious than that the hand of a man, formed for grasping, that of a mole for digging, the leg of the horse, the paddle of the porpoise, and the wing of the bat, should all be constructed on the same pattern, and should include the same bones, in the same relative positions? … [T]he parts may change to almost any extent in form and size, and yet they always remain connected together in the same order. …. Hence the same names can be given to the homologous bones in widely different animals. We see the same great law in the construction of the mouths of insects: what can be more different than the immensely long spiral proboscis of a sphinx-moth, the curious folded one of a bee or bug, and the great jaws of a beetle? — yet all these organs, serving for such different purposes, are formed by infinitely numerous modifications of an upper lip, mandibles, and two pairs of maxillae. Analogous laws govern the construction of the mouths and limbs of crustaceans. So it is with the flowers of plants.

Nothing can be more hopeless than to attempt to explain this similarity of pattern in members of the same class, by utility or by the doctrine of final causes. …. On the ordinary view of the independent creation of each being, we can only say that so it is; — that it has so pleased the Creator to construct each animal and plant.

The explanation is manifest on the theory of the natural selection of successive slight modifications, — each modification being profitable in some way to the modified form, but often affecting by correlation of growth other parts of the organisation. In changes of this nature, there will be little or no tendency to modify the original pattern, or to transpose parts. The bones of a limb might be shortened and widened to any extent, and become gradually enveloped in thick membrane, so as to serve as a fin; or a webbed foot might have all its bones, or certain bones, lengthened to any extent, and the membrane connecting them increased to any extent, so as to serve as a wing: yet in all this great amount of modification there will be no tendency to alter the framework of bones or the relative connexion of the several parts. If we suppose that the ancient progenitor, the archetype as it may be called, of all mammals, had its limbs constructed on the existing general pattern, for whatever purpose they served, we can at once perceive the plain signification of the homologous construction of the limbs throughout the whole class. So with the mouths of insects, we have only to suppose that their common progenitor had an upper lip, mandibles, and two pair of maxillae, these parts being perhaps very simple in form; and then natural selection will account for the infinite diversity in structure and function of the mouths of insects. Nevertheless, it is conceivable that the general pattern of an organ might become so much obscured as to be finally lost, by the atrophy and ultimately by the complete abortion of certain parts, by the soldering together of other parts, and by the doubling or multiplication of others, — variations which we know to be within the limits of possibility. In the paddles of the extinct gigantic sea-lizards, and in the mouths of certain suctorial crustaceans, the general pattern seems to have been thus to a certain extent obscured.

There is another and equally curious branch of the present subject; namely, the comparison not of the same part in different members of a class, but of the different parts or organs in the same individual. Most physiologists believe that the bones of the skull are homologous with — that is correspond in number and in relative connexion with — the elemental parts of a certain number of vertebrae. The anterior and posterior limbs in each member of the vertebrate and articulate classes are plainly homologous. We see the same law in comparing the wonderfully complex jaws and legs in crustaceans. It is familiar to almost every one, that in a flower the relative position of the sepals, petals, stamens, and pistils, as well as their intimate structure, are intelligible on the view that they consist of metamorphosed leaves, arranged in a spire. In monstrous plants, we often get direct evidence of the possibility of one organ being transformed into another; and we can actually see in embryonic crustaceans and in many other animals, and in flowers, that organs, which when mature become extremely different, are at an early stage of growth exactly alike.

How inexplicable are these facts on the ordinary view of creation! Why should the brain be enclosed in a box composed of such numerous and such extraordinarily shaped pieces of bone? As Owen has remarked, the benefit derived from the yielding of the separate pieces in the act of parturition of mammals, will by no means explain the same construction in the skulls of birds. Why should similar bones have been created in the formation of the wing and leg of a bat, used as they are for such totally different purposes? Why should one crustacean, which has an extremely complex mouth formed of many parts, consequently always have fewer legs; or conversely, those with many legs have simpler mouths? Why should the sepals, petals, stamens, and pistils in any individual flower, though fitted for such widely different purposes, be all constructed on the same pattern?

On the theory of natural selection, we can satisfactorily answer these questions. In the vertebrata, we see a series of internal vertebrae bearing certain processes and appendages; in the articulata, we see the body divided into a series of segments, bearing external appendages; and in flowering plants, we see a series of successive spiral whorls of leaves. An indefinite repetition of the same part or organ is the common characteristic (as Owen has observed) of all low or little-modified forms; therefore we may readily believe that the unknown progenitor of the vertebrata possessed many vertebrae; the unknown progenitor of the articulata, many segments; and the unknown progenitor of flowering plants, many spiral whorls of leaves. We have formerly seen that parts many times repeated are eminently liable to vary in number and structure; consequently it is quite probable that natural selection, during a long-continued course of modification, should have seized on a certain number of the primordially similar elements, many times repeated, and have adapted them to the most diverse purposes. And as the whole amount of modification will have been effected by slight successive steps, we need not wonder at discovering in such parts or organs, a certain degree of fundamental resemblance, retained by the strong principle of inheritance.


Naturalists frequently speak of the skull as formed of metamorphosed vertebrae: the jaws of crabs as metamorphosed legs; the stamens and pistils of flowers as metamorphosed leaves; but it would in these cases probably be more correct, as Professor Huxley has remarked, to speak of both skull and vertebrae, both jaws and legs, etc., — as having been metamorphosed, not one from the other, but from some common element. Naturalists, however, use such language only in a metaphorical sense: they are far from meaning that during a long course of descent, primordial organs of any kind — vertebrae in the one case and legs in the other — have actually been modified into skulls or jaws. Yet so strong is the appearance of a modification of this nature having occurred, that naturalists can hardly avoid employing language having this plain signification. On my view these terms may be used literally; and the wonderful fact of the jaws, for instance, of a crab retaining numerous characters, which they would probably have retained through inheritance, if they had really been metamorphosed during a long course of descent from true legs, or from some simple appendage, is explained....

Rudimentary, atrophied, or aborted organs.— Organs or parts in this strange condition, bearing the stamp of inutility, are extremely common throughout nature. For instance, rudimentary mammae are very general in the males of mammals: I presume that the “bastard-wing” in birds may be safely considered as a digit in a rudimentary state: in very many snakes one lobe of the lungs is rudimentary; in other snakes there are rudiments of the pelvis and hind limbs. Some of the cases of rudimentary organs are extremely curious; for instance, the presence of teeth in foetal whales, which when grown up have not a tooth in their heads; and the presence of teeth, which never cut through the gums, in the upper jaws of our unborn calves. It has even been stated on good authority that rudiments of teeth can be detected in the beaks of certain embryonic birds. Nothing can be plainer than that wings are formed for flight, yet in how many insects do we see wings so reduced in size as to be utterly incapable of flight, and not rarely lying under wing-cases, firmly soldered together!


...In reflecting on [rudimentary organs], every one must be struck with astonishment: for the same reasoning power which tells us plainly that most parts and organs are exquisitely adapted for certain purposes, tells us with equal plainness that these rudimentary or atrophied organs, are imperfect and useless. In works on natural history rudimentary organs are generally said to have been created “for the sake of symmetry,” or in order “to complete the scheme of nature;” but this seems to me no explanation, merely a restatement of the fact. Would it be thought sufficient to say that because planets revolve in elliptic courses round the sun, satellites follow the same course round the planets, for the sake of symmetry, and to complete the scheme of nature? An eminent physiologist accounts for the presence of rudimentary organs, by supposing that they serve to excrete matter in excess, or injurious to the system; but can we suppose that the minute papilla, which often represents the pistil in male flowers, and which is formed merely of cellular tissue, can thus act? Can we suppose that the formation of rudimentary teeth which are subsequently absorbed, can be of any service to the rapidly growing embryonic calf by the excretion of precious phosphate of lime? When a man’s fingers have been amputated, imperfect nails sometimes appear on the stumps: I could as soon believe that these vestiges of nails have appeared, not from unknown laws of growth, but in order to excrete horny matter, as that the rudimentary nails on the fin of the manatee were formed for this purpose.

On my view of descent with modification, the origin of rudimentary organs is simple. We have plenty of cases of rudimentary organs in our domestic productions, — as the stump of a tail in tailless breeds, — the vestige of an ear in earless breeds, — the reappearance of minute dangling horns in hornless breeds of cattle, … — and the state of the whole flower in the cauliflower. We often see rudiments of various parts in monsters. But I doubt whether any of these cases throw light on the origin of rudimentary organs in a state of nature, further than by showing that rudiments can be produced; for I doubt whether species under nature ever undergo abrupt changes. I believe that disuse has been the main agency; that it has led in successive generations to the gradual reduction of various organs, until they have become rudimentary, — as in the case of the eyes of animals inhabiting dark caverns, and of the wings of birds inhabiting oceanic islands, which have seldom been forced to take flight, and have ultimately lost the power of flying. Again, an organ useful under certain conditions, might become injurious under others, as with the wings of beetles living on small and exposed islands; and in this case natural selection would continue slowly to reduce the organ, until it was rendered harmless and rudimentary.

Any change in function, which can be effected by insensibly small steps, is within the power of natural selection; so that an organ rendered, during changed habits of life, useless or injurious for one purpose, might easily be modified and used for another purpose. Or an organ might be retained for one alone of its former functions. An organ, when rendered useless, may well be variable, for its variations cannot be checked by natural selection. At whatever period of life disuse or selection reduces an organ, and this will generally be when the being has come to maturity and to its full powers of action, the principle of inheritance at corresponding ages will reproduce the organ in its reduced state at the same age, and consequently will seldom affect or reduce it in the embryo. Thus we can understand the greater relative size of rudimentary organs in the embryo, and their lesser relative size in the adult. But if each step of the process of reduction were to be inherited, not at the corresponding age, but at an extremely early period of life (as we have good reason to believe to be possible) the rudimentary part would tend to be wholly lost, and we should have a case of complete abortion.….

As the presence of rudimentary organs is thus due to the tendency in every part of the organisation, which has long existed, to be inherited — we can understand, on the genealogical view of classification, how it is that systematists have found rudimentary parts as useful as, or even sometimes more useful than, parts of high physiological importance. Rudimentary organs may be compared with the letters in a word, still retained in the spelling, but become useless in the pronunciation, but which serve as a clue in seeking for its derivation. On the view of descent with modification, we may conclude that the existence of organs in a rudimentary, imperfect, and useless condition, or quite aborted, far from presenting a strange difficulty, as they assuredly do on the ordinary doctrine of creation, might even have been anticipated, and can be accounted for by the laws of inheritance.




Recapitulation of the difficulties on the theory of Natural Selection. Recapitulation of the general and special circumstances in its favour. Causes of the general belief in the immutability of species. How far the theory of natural selection may be extended. Effects of its adoption on the study of Natural history. Concluding remarks.

...That many and grave objections may be advanced against the theory of descent with modification through natural selection, I do not deny. I have endeavoured to give to them their full force. Nothing at first can appear more difficult to believe than that the more complex organs and instincts should have been perfected, not by means superior to, though analogous with, human reason, but by the accumulation of innumerable slight variations, each good for the individual possessor. Nevertheless, this difficulty, though appearing to our imagination insuperably great, cannot be considered real if we admit the following propositions, namely, — that gradations in the perfection of any organ or instinct, which we may consider, either do now exist or could have existed, each good of its kind, — that all organs and instincts are, in ever so slight a degree, variable, — and, lastly, that there is a struggle for existence leading to the preservation of each profitable deviation of structure or instinct. The truth of these propositions cannot, I think, be disputed.

It is, no doubt, extremely difficult even to conjecture by what gradations many structures have been perfected, more especially amongst broken and failing groups of organic beings; but we see so many strange gradations in nature, as is proclaimed by the canon, “Natura non facit saltum,” that we ought to be extremely cautious in saying that any organ or instinct, or any whole being, could not have arrived at its present state by many graduated steps. There are, it must be admitted, cases of special difficulty on the theory of natural selection; and one of the most curious of these is the existence of two or three defined castes of workers or sterile females in the same community of ants; but I have attempted to show how this difficulty can be mastered.


Turning to geographical distribution, the difficulties encountered on the theory of descent with modification are grave enough. All the individuals of the same species, and all the species of the same genus, or even higher group, must have descended from common parents; and therefore, in however distant and isolated parts of the world they are now found, they must in the course of successive generations have passed from some one part to the others. We are often wholly unable even to conjecture how this could have been effected. Yet, as we have reason to believe that some species have retained the same specific form for very long periods, enormously long as measured by years, too much stress ought not to be laid on the occasional wide diffusion of the same species; for during very long periods of time there will always be a good chance for wide migration by many means. A broken or interrupted range may often be accounted for by the extinction of the species in the intermediate regions. It cannot be denied that we are as yet very ignorant of the full extent of the various climatal and geographical changes which have affected the earth during modern periods; and such changes will obviously have greatly facilitated migration….

As on the theory of natural selection an interminable number of intermediate forms must have existed, linking together all the species in each group by gradations as fine as our present varieties, it may be asked, Why do we not see these linking forms all around us? Why are not all organic beings blended together in an inextricable chaos? With respect to existing forms, we should remember that we have no right to expect (excepting in rare cases) to discover directly connecting links between them, but only between each and some extinct and supplanted form. Even on a wide area, which has during a long period remained continuous, and of which the climate and other conditions of life change insensibly in going from a district occupied by one species into another district occupied by a closely allied species, we have no just right to expect often to find intermediate varieties in the intermediate zone. For we have reason to believe that only a few species are undergoing change at any one period; and all changes are slowly effected. I have also shown that the intermediate varieties which will at first probably exist in the intermediate zones, will be liable to be supplanted by the allied forms on either hand; and the latter, from existing in greater numbers, will generally be modified and improved at a quicker rate than the intermediate varieties, which exist in lesser numbers; so that the intermediate varieties will, in the long run, be supplanted and exterminated.

On this doctrine of the extermination of an infinitude of connecting links, between the living and extinct inhabitants of the world, and at each successive period between the extinct and still older species, why is not every geological formation charged with such links? Why does not every collection of fossil remains afford plain evidence of the gradation and mutation of the forms of life? We meet with no such evidence, and this is the most obvious and forcible of the many objections which may be urged against my theory. Why, again, do whole groups of allied species appear, though certainly they often falsely appear, to have come in suddenly on the several geological stages? Why do we not find great piles of strata beneath the Silurian system, stored with the remains of the progenitors of the Silurian groups of fossils? For certainly on my theory such strata must somewhere have been deposited at these ancient and utterly unknown epochs in the world’s history.

I can answer these questions and grave objections only on the supposition that the geological record is far more imperfect than most geologists believe. It cannot be objected that there has not been time sufficient for any amount of organic change; for the lapse of time has been so great as to be utterly inappreciable by the human intellect. The number of specimens in all our museums is absolutely as nothing compared with the countless generations of countless species which certainly have existed. We should not be able to recognise a species as the parent of any one or more species if we were to examine them ever so closely, unless we likewise possessed many of the intermediate links between their past or parent and present states; and these many links we could hardly ever expect to discover, owing to the imperfection of the geological record. Numerous existing doubtful forms could be named which are probably varieties; but who will pretend that in future ages so many fossil links will be discovered, that naturalists will be able to decide, on the common view, whether or not these doubtful forms are varieties? As long as most of the links between any two species are unknown, if any one link or intermediate variety be discovered, it will simply be classed as another and distinct species. Only a small portion of the world has been geologically explored. Only organic beings of certain classes can be preserved in a fossil condition, at least in any great number. Widely ranging species vary most, and varieties are often at first local, — both causes rendering the discovery of intermediate links less likely. Local varieties will not spread into other and distant regions until they are considerably modified and improved; and when they do spread, if discovered in a geological formation, they will appear as if suddenly created there, and will be simply classed as new species. Most formations have been intermittent in their accumulation; and their duration, I am inclined to believe, has been shorter than the average duration of specific forms. Successive formations are separated from each other by enormous blank intervals of time; for fossiliferous formations, thick enough to resist future degradation, can be accumulated only where much sediment is deposited on the subsiding bed of the sea. During the alternate periods of elevation and of stationary level the record will be blank. During these latter periods there will probably be more variability in the forms of life; during periods of subsidence, more extinction.


Such is the sum of the several chief objections and difficulties which may justly be urged against my theory; and I have now briefly recapitulated the answers and explanations which can be given to them. I have felt these difficulties far too heavily during many years to doubt their weight. But it deserves especial notice that the more important objections relate to questions on which we are confessedly ignorant; nor do we know how ignorant we are. We do not know all the possible transitional gradations between the simplest and the most perfect organs; it cannot be pretended that we know all the varied means of Distribution during the long lapse of years, or that we know how imperfect the Geological Record is. Grave as these several difficulties are, in my judgment they do not overthrow the theory of descent with modification.

Now let us turn to the other side of the argument. Under domestication we see much variability.... Man does not actually produce variability [b]ut man can and does select the variations given to him by nature, and thus accumulate them in any desired manner. He thus adapts animals and plants for his own benefit or pleasure. He may do this methodically, or he may do it unconsciously by preserving the individuals most useful to him at the time, without any thought of altering the breed. It is certain that he can largely influence the character of a breed by selecting, in each successive generation, individual differences so slight as to be quite inappreciable by an uneducated eye. This process of selection has been the great agency in the production of the most distinct and useful domestic breeds. That many of the breeds produced by man have to a large extent the character of natural species, is shown by the inextricable doubts whether very many of them are varieties or aboriginal species.

There is no obvious reason why the principles which have acted so efficiently under domestication should not have acted under nature. In the preservation of favoured individuals and races, during the constantly-recurrent Struggle for Existence, we see the most powerful and ever-acting means of selection. The struggle for existence inevitably follows from the high geometrical ratio of increase which is common to all organic beings. …. More individuals are born than can possibly survive. A grain in the balance will determine which individual shall live and which shall die, — which variety or species shall increase in number, and which shall decrease, or finally become extinct. As the individuals of the same species come in all respects into the closest competition with each other, the struggle will generally be most severe between them; it will be almost equally severe between the varieties of the same species, and next in severity between the species of the same genus. But the struggle will often be very severe between beings most remote in the scale of nature. The slightest advantage in one being, at any age or during any season, over those with which it comes into competition, or better adaptation in however slight a degree to the surrounding physical conditions, will turn the balance.

With animals having separated sexes there will in most cases be a struggle between the males for possession of the females. The most vigorous individuals, or those which have most successfully struggled with their conditions of life, will generally leave most progeny. But success will often depend on having special weapons or means of defence, or on the charms of the males; and the slightest advantage will lead to victory.

As geology plainly proclaims that each land has undergone great physical changes, we might have expected that organic beings would have varied under nature, in the same way as they generally have varied under the changed conditions of domestication. And if there be any variability under nature, it would be an unaccountable fact if natural selection had not come into play….

If then we have under nature variability and a powerful agent always ready to act and select, why should we doubt that variations in any way useful to beings, under their excessively complex relations of life, would be preserved, accumulated, and inherited? Why, if man can by patience select variations most useful to himself, should nature fail in selecting variations useful, under changing conditions of life, to her living products? What limit can be put to this power, acting during long ages and rigidly scrutinising the whole constitution, structure, and habits of each creature, — favouring the good and rejecting the bad? I can see no limit to this power, in slowly and beautifully adapting each form to the most complex relations of life. The theory of natural selection, even if we looked no further than this, seems to me to be in itself probable. I have already recapitulated, as fairly as I could, the opposed difficulties and objections: now let us turn to the special facts and arguments in favour of the theory.

On the view that species are only strongly marked and permanent varieties, and that each species first existed as a variety, we can see why it is that no line of demarcation can be drawn between species, commonly supposed to have been produced by special acts of creation, and varieties which are acknowledged to have been produced by secondary laws. On this same view we can understand how it is that in each region where many species of a genus have been produced, and where they now flourish, these same species should present many varieties; for where the manufactory of species has been active, we might expect, as a general rule, to find it still in action; and this is the case if varieties be incipient species. Moreover, the species of the larger genera, which afford the greater number of varieties or incipient species, retain to a certain degree the character of varieties; for they differ from each other by a less amount of difference than do the species of smaller genera. The closely allied species also of the larger genera apparently have restricted ranges, and they are clustered in little groups round other species — in which respects they resemble varieties. These are strange relations on the view of each species having been independently created, but are intelligible if all species first existed as varieties.

As each species tends by its geometrical ratio of reproduction to increase inordinately in number; and as the modified descendants of each species will be enabled to increase by so much the more as they become more diversified in habits and structure, so as to be enabled to seize on many and widely different places in the economy of nature, there will be a constant tendency in natural selection to preserve the most divergent offspring of any one species. Hence during a long-continued course of modification, the slight differences, characteristic of varieties of the same species, tend to be augmented into the greater differences characteristic of species of the same genus. New and improved varieties will inevitably supplant and exterminate the older, less improved and intermediate varieties; and thus species are rendered to a large extent defined and distinct objects. Dominant species belonging to the larger groups tend to give birth to new and dominant forms; so that each large group tends to become still larger, and at the same time more divergent in character. But as all groups cannot thus succeed in increasing in size, for the world would not hold them, the more dominant groups beat the less dominant. This tendency in the large groups to go on increasing in size and diverging in character, together with the almost inevitable contingency of much extinction, explains the arrangement of all the forms of life, in groups subordinate to groups, all within a few great classes, which we now see everywhere around us, and which has prevailed throughout all time. This grand fact of the grouping of all organic beings seems to me utterly inexplicable on the theory of creation.

As natural selection acts solely by accumulating slight, successive, favourable variations, it can produce no great or sudden modification; it can act only by very short and slow steps. Hence the canon of “Natura non facit saltum,” which every fresh addition to our knowledge tends to make more strictly correct, is on this theory simply intelligible. We can plainly see why nature is prodigal in variety, though niggard in innovation. But why this should be a law of nature if each species has been independently created, no man can explain.

Many other facts are, as it seems to me, explicable on this theory. How strange it is that a bird, under the form of woodpecker, should have been created to prey on insects on the ground; that upland geese, which never or rarely swim, should have been created with webbed feet; that a thrush should have been created to dive and feed on sub-aquatic insects; and that a petrel should have been created with habits and structure fitting it for the life of an auk or grebe! and so on in endless other cases. But on the view of each species constantly trying to increase in number, with natural selection always ready to adapt the slowly varying descendants of each to any unoccupied or ill-occupied place in nature, these facts cease to be strange, or perhaps might even have been anticipated.

As natural selection acts by competition, it adapts the inhabitants of each country only in relation to the degree of perfection of their associates; so that we need feel no surprise at the inhabitants of any one country, although on the ordinary view supposed to have been specially created and adapted for that country, being beaten and supplanted by the naturalised productions from another land. Nor ought we to marvel if all the contrivances in nature be not, as far as we can judge, absolutely perfect; and if some of them be abhorrent to our ideas of fitness. We need not marvel at the sting of the bee causing the bee’s own death; at drones being produced in such vast numbers for one single act, and being then slaughtered by their sterile sisters; at the astonishing waste of pollen by our fir-trees; at the instinctive hatred of the queen bee for her own fertile daughters; at ichneumonidae feeding within the live bodies of caterpillars; and at other such cases. The wonder indeed is, on the theory of natural selection, that more cases of the want of absolute perfection have not been observed.


Glancing at instincts, marvellous as some are, they offer no greater difficulty than does corporeal structure on the theory of the natural selection of successive, slight, but profitable modifications. We can thus understand why nature moves by graduated steps in endowing different animals of the same class with their several instincts. …. On the view of all the species of the same genus having descended from a common parent, and having inherited much in common, we can understand how it is that allied species, when placed under considerably different conditions of life, yet should follow nearly the same instincts; why the thrush of South America, for instance, lines her nest with mud like our British species. On the view of instincts having been slowly acquired through natural selection we need not marvel at some instincts being apparently not perfect and liable to mistakes, and at many instincts causing other animals to suffer.


If we admit that the geological record is imperfect in an extreme degree, then such facts as the record gives, support the theory of descent with modification. New species have come on the stage slowly and at successive intervals; and the amount of change, after equal intervals of time, is widely different in different groups. The extinction of species and of whole groups of species, which has played so conspicuous a part in the history of the organic world, almost inevitably follows on the principle of natural selection; for old forms will be supplanted by new and improved forms. Neither single species nor groups of species reappear when the chain of ordinary generation has once been broken. The gradual diffusion of dominant forms, with the slow modification of their descendants, causes the forms of life, after long intervals of time, to appear as if they had changed simultaneously throughout the world. The fact of the fossil remains of each formation being in some degree intermediate in character between the fossils in the formations above and below, is simply explained by their intermediate position in the chain of descent. The grand fact that all extinct organic beings belong to the same system with recent beings, falling either into the same or into intermediate groups, follows from the living and the extinct being the offspring of common parents. As the groups which have descended from an ancient progenitor have generally diverged in character, the progenitor with its early descendants will often be intermediate in character in comparison with its later descendants; and thus we can see why the more ancient a fossil is, the oftener it stands in some degree intermediate between existing and allied groups. Recent forms are generally looked at as being, in some vague sense, higher than ancient and extinct forms; and they are in so far higher as the later and more improved forms have conquered the older and less improved organic beings in the struggle for life. Lastly, the law of the long endurance of allied forms on the same continent, — of marsupials in Australia, of edentata in America, and other such cases, — is intelligible, for within a confined country, the recent and the extinct will naturally be allied by descent.

Looking to geographical distribution, if we admit that there has been during the long course of ages much migration from one part of the world to another, owing to former climatal and geographical changes and to the many occasional and unknown means of dispersal, then we can understand, on the theory of descent with modification, most of the great leading facts in Distribution. We can see why there should be so striking a parallelism in the distribution of organic beings throughout space, and in their geological succession throughout time; for in both cases the beings have been connected by the bond of ordinary generation, and the means of modification have been the same. We see the full meaning of the wonderful fact, which must have struck every traveller, namely, that on the same continent, under the most diverse conditions, under heat and cold, on mountain and lowland, on deserts and marshes, most of the inhabitants within each great class are plainly related; for they will generally be descendants of the same progenitors and early colonists. On this same principle of former migration, combined in most cases with modification, we can understand, by the aid of the Glacial period, the identity of some few plants, and the close alliance of many others, on the most distant mountains, under the most different climates; and likewise the close alliance of some of the inhabitants of the sea in the northern and southern temperate zones, though separated by the whole intertropical ocean. Although two areas may present the same physical conditions of life, we need feel no surprise at their inhabitants being widely different, if they have been for a long period completely separated from each other; for as the relation of organism to organism is the most important of all relations, and as the two areas will have received colonists from some third source or from each other, at various periods and in different proportions, the course of modification in the two areas will inevitably be different.

On this view of migration, with subsequent modification, we can see why oceanic islands should be inhabited by few species, but of these, that many should be peculiar. We can clearly see why those animals which cannot cross wide spaces of ocean, as frogs and terrestrial mammals, should not inhabit oceanic islands; and why, on the other hand, new and peculiar species of bats, which can traverse the ocean, should so often be found on islands far distant from any continent. Such facts as the presence of peculiar species of bats, and the absence of all other mammals, on oceanic islands, are utterly inexplicable on the theory of independent acts of creation.

The existence of closely allied or representative species in any two areas, implies, on the theory of descent with modification, that the same parents formerly inhabited both areas; and we almost invariably find that wherever many closely allied species inhabit two areas, some identical species common to both still exist. Wherever many closely allied yet distinct species occur, many doubtful forms and varieties of the same species likewise occur. It is a rule of high generality that the inhabitants of each area are related to the inhabitants of the nearest source whence immigrants might have been derived. We see this in nearly all the plants and animals of the Galapagos archipelago, of Juan Fernandez, and of the other American islands being related in the most striking manner to the plants and animals of the neighbouring American mainland; and those of the Cape de Verde archipelago and other African islands to the African mainland. It must be admitted that these facts receive no explanation on the theory of creation.

The fact, as we have seen, that all past and present organic beings constitute one grand natural system, with group subordinate to group, and with extinct groups often falling in between recent groups, is intelligible on the theory of natural selection with its contingencies of extinction and divergence of character. On these same principles we see how it is, that the mutual affinities of the species and genera within each class are so complex and circuitous. We see why certain characters are far more serviceable than others for classification; — why adaptive characters, though of paramount importance to the being, are of hardly any importance in classification; why characters derived from rudimentary parts, though of no service to the being, are often of high classificatory value; and why embryological characters are the most valuable of all. The real affinities of all organic beings are due to inheritance or community of descent. The natural system is a genealogical arrangement, in which we have to discover the lines of descent by the most permanent characters, however slight their vital importance may be.

The framework of bones being the same in the hand of a man, wing of a bat, fin of the porpoise, and leg of the horse, — the same number of vertebrae forming the neck of the giraffe and of the elephant, — and innumerable other such facts, at once explain themselves on the theory of descent with slow and slight successive modifications. The similarity of pattern in the wing and leg of a bat, though used for such different purpose, — in the jaws and legs of a crab, — in the petals, stamens, and pistils of a flower, is likewise intelligible on the view of the gradual modification of parts or organs, which were alike in the early progenitor of each class. On the principle of successive variations not always supervening at an early age, and being inherited at a corresponding not early period of life, we can clearly see why the embryos of mammals, birds, reptiles, and fishes should be so closely alike, and should be so unlike the adult forms. We may cease marvelling at the embryo of an air-breathing mammal or bird having branchial slits and arteries running in loops, like those in a fish which has to breathe the air dissolved in water, by the aid of well-developed branchiae.

Disuse, aided sometimes by natural selection, will often tend to reduce an organ, when it has become useless by changed habits or under changed conditions of life; and we can clearly understand on this view the meaning of rudimentary organs. But disuse and selection will generally act on each creature, when it has come to maturity and has to play its full part in the struggle for existence, and will thus have little power of acting on an organ during early life; hence the organ will not be much reduced or rendered rudimentary at this early age.


I have now recapitulated the chief facts and considerations which have thoroughly convinced me that species have changed, and are still slowly changing by the preservation and accumulation of successive slight favourable variations. Why, it may be asked, have all the most eminent living naturalists and geologists rejected this view of the mutability of species? It cannot be asserted that organic beings in a state of nature are subject to no variation; it cannot be proved that the amount of variation in the course of long ages is a limited quantity; no clear distinction has been, or can be, drawn between species and well-marked varieties. It cannot be maintained that species when intercrossed are invariably sterile, and varieties invariably fertile; or that sterility is a special endowment and sign of creation. The belief that species were immutable productions was almost unavoidable as long as the history of the world was thought to be of short duration; and now that we have acquired some idea of the lapse of time, we are too apt to assume, without proof, that the geological record is so perfect that it would have afforded us plain evidence of the mutation of species, if they had undergone mutation.

But the chief cause of our natural unwillingness to admit that one species has given birth to other and distinct species, is that we are always slow in admitting any great change of which we do not see the intermediate steps. The difficulty is the same as that felt by so many geologists, when Lyell first insisted that long lines of inland cliffs had been formed, and great valleys excavated, by the slow action of the coast-waves. The mind cannot possibly grasp the full meaning of the term of a hundred million years; it cannot add up and perceive the full effects of many slight variations, accumulated during an almost infinite number of generations.

Although I am fully convinced of the truth of the views given in this volume under the form of an abstract, I by no means expect to convince experienced naturalists whose minds are stocked with a multitude of facts all viewed, during a long course of years, from a point of view directly opposite to mine. It is so easy to hide our ignorance under such expressions as the “plan of creation,” “unity of design,” etc., and to think that we give an explanation when we only restate a fact. Any one whose disposition leads him to attach more weight to unexplained difficulties than to the explanation of a certain number of facts will certainly reject my theory. A few naturalists, endowed with much flexibility of mind, and who have already begun to doubt on the immutability of species, may be influenced by this volume; but I look with confidence to the future, to young and rising naturalists, who will be able to view both sides of the question with impartiality. Whoever is led to believe that species are mutable will do good service by conscientiously expressing his conviction; for only thus can the load of prejudice by which this subject is overwhelmed be removed.


It may be asked how far I extend the doctrine of the modification of species. The question is difficult to answer, because the more distinct the forms are which we may consider, by so much the arguments fall away in force. But some arguments of the greatest weight extend very far. All the members of whole classes can be connected together by chains of affinities, and all can be classified on the same principle, in groups subordinate to groups. Fossil remains sometimes tend to fill up very wide intervals between existing orders. Organs in a rudimentary condition plainly show that an early progenitor had the organ in a fully developed state; and this in some instances necessarily implies an enormous amount of modification in the descendants. Throughout whole classes various structures are formed on the same pattern, and at an embryonic age the species closely resemble each other. Therefore I cannot doubt that the theory of descent with modification embraces all the members of the same class. I believe that animals have descended from at most only four or five progenitors, and plants from an equal or lesser number.

[Editor’s note: It is now believed that all animals and plants and bacteria are descended from a single progenitor.]

Analogy would lead me one step further, namely, to the belief that all animals and plants have descended from some one prototype. But analogy may be a deceitful guide. Nevertheless all living things have much in common, in their chemical composition, their germinal vesicles, their cellular structure, and their laws of growth and reproduction. We see this even in so trifling a circumstance as that the same poison often similarly affects plants and animals; or that the poison secreted by the gall-fly produces monstrous growths on the wild rose or oak-tree. Therefore I should infer from analogy that probably all the organic beings which have ever lived on this earth have descended from some one primordial form, into which life was first breathed.

[Editor’s note: The universality of the DNA code shows that all life has indeed descended from one primordial form.]

When the views entertained in this volume on the origin of species, or when analogous views are generally admitted, we can dimly foresee that there will be a considerable revolution in natural history. Systematists will be able to pursue their labours as at present; but they will not be incessantly haunted by the shadowy doubt whether this or that form be in essence a species. This I feel sure, and I speak after experience, will be no slight relief. The endless disputes whether or not some fifty species of British brambles are true species will cease. Systematists will have only to decide (not that this will be easy) whether any form be sufficiently constant and distinct from other forms, to be capable of definition; and if definable, whether the differences be sufficiently important to deserve a specific name. This latter point will become a far more essential consideration than it is at present; for differences, however slight, between any two forms, if not blended by intermediate gradations, are looked at by most naturalists as sufficient to raise both forms to the rank of species. Hereafter we shall be compelled to acknowledge that the only distinction between species and well-marked varieties is, that the latter are known, or believed, to be connected at the present day by intermediate gradations, whereas species were formerly thus connected. Hence, without quite rejecting the consideration of the present existence of intermediate gradations between any two forms, we shall be led to weigh more carefully and to value higher the actual amount of difference between them. It is quite possible that forms now generally acknowledged to be merely varieties may hereafter be thought worthy of specific names, as with the primrose and cowslip; and in this case scientific and common language will come into accordance. In short, we shall have to treat species in the same manner as those naturalists treat genera, who admit that genera are merely artificial combinations made for convenience. This may not be a cheering prospect; but we shall at least be freed from the vain search for the undiscovered and undiscoverable essence of the term species.

The other and more general departments of natural history will rise greatly in interest. The terms used by naturalists of affinity, relationship, community of type, paternity, morphology, adaptive characters, rudimentary and aborted organs, etc., will cease to be metaphorical, and will have a plain signification. When we no longer look at an organic being as a savage looks at a ship, as at something wholly beyond his comprehension; when we regard every production of nature as one which has had a history; when we contemplate every complex structure and instinct as the summing up of many contrivances, each useful to the possessor, nearly in the same way as when we look at any great mechanical invention as the summing up of the labour, the experience, the reason, and even the blunders of numerous workmen; when we thus view each organic being, how far more interesting, I speak from experience, will the study of natural history become!

A grand and almost untrodden field of inquiry will be opened, on the causes and laws of variation, on correlation of growth, on the effects of use and disuse, on the direct action of external conditions, and so forth. The study of domestic productions will rise immensely in value. A new variety raised by man will be a far more important and interesting subject for study than one more species added to the infinitude of already recorded species. Our classifications will come to be, as far as they can be so made, genealogies; and will then truly give what may be called the plan of creation. The rules for classifying will no doubt become simpler when we have a definite object in view. We possess no pedigrees or armorial bearings; and we have to discover and trace the many diverging lines of descent in our natural genealogies, by characters of any kind which have long been inherited. Rudimentary organs will speak infallibly with respect to the nature of long-lost structures. Species and groups of species, which are called aberrant, and which may fancifully be called living fossils, will aid us in forming a picture of the ancient forms of life. Embryology will reveal to us the structure, in some degree obscured, of the prototypes of each great class.

When we can feel assured that all the individuals of the same species, and all the closely allied species of most genera, have within a not very remote period descended from one parent, and have migrated from some one birthplace; and when we better know the many means of migration, then, by the light which geology now throws, and will continue to throw, on former changes of climate and of the level of the land, we shall surely be enabled to trace in an admirable manner the former migrations of the inhabitants of the whole world. Even at present, by comparing the differences of the inhabitants of the sea on the opposite sides of a continent, and the nature of the various inhabitants of that continent in relation to their apparent means of immigration, some light can be thrown on ancient geography.

The noble science of Geology loses glory from the extreme imperfection of the record. The crust of the earth with its embedded remains must not be looked at as a well-filled museum, but as a poor collection made at hazard and at rare intervals. The accumulation of each great fossiliferous formation will be recognised as having depended on an unusual concurrence of circumstances, and the blank intervals between the successive stages as having been of vast duration. But we shall be able to gauge with some security the duration of these intervals by a comparison of the preceding and succeeding organic forms. We must be cautious in attempting to correlate as strictly contemporaneous two formations, which include few identical species, by the general succession of their forms of life. As species are produced and exterminated by slowly acting and still existing causes, and not by miraculous acts of creation and by catastrophes; and as the most important of all causes of organic change is one which is almost independent of altered and perhaps suddenly altered physical conditions, namely, the mutual relation of organism to organism, … it follows, that the amount of organic change in the fossils of consecutive formations probably serves as a fair measure of the lapse of actual time. A number of species, however, keeping in a body might remain for a long period unchanged, whilst within this same period, several of these species, by migrating into new countries and coming into competition with foreign associates, might become modified; so that we must not overrate the accuracy of organic change as a measure of time. During early periods of the earth’s history, when the forms of life were probably fewer and simpler, the rate of change was probably slower; and at the first dawn of life, when very few forms of the simplest structure existed, the rate of change may have been slow in an extreme degree. The whole history of the world, as at present known, although of a length quite incomprehensible by us, will hereafter be recognised as a mere fragment of time, compared with the ages which have elapsed since the first creature, the progenitor of innumerable extinct and living descendants, was created.

In the distant future I see open fields for far more important researches. Psychology will be based on a new foundation, that of the necessary acquirement of each mental power and capacity by gradation. Light will be thrown on the origin of man and his history.

Authors of the highest eminence seem to be fully satisfied with the view that each species has been independently created. To my mind it accords better with what we know of the laws impressed on matter by the Creator, that the production and extinction of the past and present inhabitants of the world should have been due to secondary causes, like those determining the birth and death of the individual. When I view all beings not as special creations, but as the lineal descendants of some few beings which lived long before the first bed of the Silurian system was deposited, they seem to me to become ennobled. Judging from the past, we may safely infer that not one living species will transmit its unaltered likeness to a distant futurity. And of the species now living very few will transmit progeny of any kind to a far distant futurity; for the manner in which all organic beings are grouped, shows that the greater number of species of each genus, and all the species of many genera, have left no descendants, but have become utterly extinct. We can so far take a prophetic glance into futurity as to foretell that it will be the common and widely-spread species, belonging to the larger and dominant groups, which will ultimately prevail and procreate new and dominant species. As all the living forms of life are the lineal descendants of those which lived long before the Silurian epoch, we may feel certain that the ordinary succession by generation has never once been broken, and that no cataclysm has desolated the whole world. Hence we may look with some confidence to a secure future of equally inappreciable length. And as natural selection works solely by and for the good of each being, all corporeal and mental endowments will tend to progress towards perfection.

It is interesting to contemplate an entangled bank, clothed with many plants of many kinds, with birds singing on the bushes, with various insects flitting about, and with worms crawling through the damp earth, and to reflect that these elaborately constructed forms, so different from each other, and dependent on each other in so complex a manner, have all been produced by laws acting around us. These laws, taken in the largest sense, being Growth with Reproduction; Inheritance which is almost implied by reproduction; Variability from the indirect and direct action of the external conditions of life … a Ratio of Increase so high as to lead to a Struggle for Life, and as a consequence to Natural Selection, entailing Divergence of Character and the Extinction of less-improved forms. Thus, from the war of nature, from famine and death, the most exalted object which we are capable of conceiving, namely, the production of the higher animals, directly follows. There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one; and that, whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved.