On the Origin of Species By Means of Natural Selection

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. A number of curious and authentic instances could be given of the inheritance of all shades of disposition and tastes, and likewise of the oddest tricks, associated with certain frames of mind or periods of time. But let 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.

How strongly these domestic instincts, habits, and dispositions are inherited, and how curiously they become mingled, is well shown when different breeds of dogs are crossed. Thus it is known that a cross with a bull-dog has affected for many generations the courage and obstinacy of greyhounds; and a cross with a greyhound has given to a whole family of shepherd-dogs a tendency to hunt hares. These domestic instincts, when thus tested by crossing, resemble natural instincts, which in a like manner become curiously blended together, and for a long period exhibit traces of the instincts of either parent: for example, Le Roy describes a dog, whose great-grandfather was a wolf, and this dog showed a trace of its wild parentage only in one way, by not coming in a straight line to his master when called.

Domestic instincts are sometimes spoken of as actions which have become inherited solely from long-continued and compulsory habit, but this, I think, is not true. No one would ever have thought of teaching, or probably could have taught, the tumbler-pigeon to tumble, – an action which, as I have witnessed, is performed by young birds, that have never seen a pigeon tumble. We may believe that some one pigeon showed a slight tendency to this strange habit, and that the long-continued selection of the best individuals in successive generations made tumblers what they now are; and near Glasgow there are house-tumblers, as I hear from Mr. Brent, which cannot fly eighteen inches high without going head over heels. It may be doubted whether any one would have thought of training a dog to point, had not some one dog naturally shown a tendency in this line; and this is known occasionally to happen, as I once saw in a pure terrier. When the first tendency was once displayed, methodical selection and the inherited effects of compulsory training in each successive generation would soon complete the work; and unconscious selection is still at work, as each man tries to procure, without intending to improve the breed, dogs which will stand and hunt best. On the other hand, habit alone in some cases has sufficed; no animal is more difficult to tame than the young of the wild rabbit; scarcely any animal is tamer than the young of the tame rabbit; but I do not suppose that domestic rabbits have ever been selected for tameness; and I presume that we must attribute the whole of the inherited change from extreme wildness to extreme tameness, simply to habit and long-continued close confinement.

Natural instincts are lost under domestication: a remarkable instance of this is seen in those breeds of fowls which very rarely or never become "broody," that is, never wish to sit on their eggs. Familiarity alone prevents our seeing how universally and largely the minds of our domestic animals have been modified by domestication. It is scarcely possible to doubt that the love of man has become instinctive in the dog. All wolves, foxes, jackals, and species of the cat genus, when kept tame, are most eager to attack poultry, sheep, and pigs; and this tendency has been found incurable in dogs which have been brought home as puppies from countries, such as Tierra del Fuego and Australia, where the savages do not keep these domestic animals. How rarely, on the other hand, do our civilised dogs, even when quite young, require to be taught not to attack poultry, sheep, and pigs! No doubt they occasionally do make an attack, and are then beaten; and if not cured, they are destroyed; so that habit, with some degree of selection, has probably concurred in civilising by inheritance our dogs. On the other hand, young chickens have lost, wholly by habit, that fear of the dog and cat which no doubt was originally instinctive in them, in the same way as it is so plainly instinctive in young pheasants, though reared under a hen. It is not that chickens have lost all fear, but fear only of dogs and cats, for if the hen gives the danger-chuckle, they will run (more especially young turkeys) from under her, and conceal themselves in the surrounding grass or thickets; and this is evidently done for the instinctive purpose of allowing, as we see in wild ground-birds, their mother to fly away. But this instinct retained by our chickens has become useless under domestication, for the mother-hen has almost lost by disuse the power of flight.

Hence, we may conclude, that domestic instincts have been acquired and natural instincts have been lost partly by habit, and partly by man selecting and accumulating during successive generations, peculiar mental habits and actions, which at first appeared from what we must in our ignorance call an accident. In some cases compulsory habit alone has sufficed to produce such inherited mental changes; in other cases compulsory habit has done nothing, and all has been the result of selection, pursued both methodically and unconsciously; but in most cases, probably, habit and selection have acted together.

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.

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. It has been asserted that the American cuckoo occasionally lays her eggs in other birds' nests; but I hear on the high authority of Dr. Brewer, that this is a mistake. Nevertheless, I could give several instances of various birds which have been known occasionally to lay their eggs in other birds' nests. 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.

The occasional habit of birds laying their eggs in other birds' nests, either of the same or of a distinct species, is not very uncommon with the Gallinaceae; and this perhaps explains the origin of a singular instinct in the allied group of ostriches. For several hen ostriches, at least in the case of the American species, unite and lay first a few eggs in one nest and then in another; and these are hatched by the males. This instinct may probably be accounted for by the fact of the hens laying a large number of eggs; but, as in the case of the cuckoo, at intervals of two or three days. This instinct, however, of the American ostrich has not as yet been perfected; for a surprising number of eggs lie strewed over the plains, so that in one day's hunting I picked up no less than twenty lost and wasted eggs.

Many bees are parasitic, and always lay their eggs in the nests of bees of other kinds. This case is more remarkable than that of the cuckoo; for these bees have not only their instincts but their structure modified in accordance with their parasitic habits; for they do not possess the pollen-collecting apparatus which would be necessary if they had to store food for their own young. Some species, likewise, of Sphegidae (wasp-like insects) are parasitic on other species; and M. Fabre has lately shown good reason for believing that although the Tachytes nigra generally makes its own burrow and stores it with paralysed prey for its own larvae to feed on, yet that when this insect finds a burrow already made and stored by another sphex, it takes advantage of the prize, and becomes for the occasion parasitic. In this case, as with the supposed case of the cuckoo, I can see no difficulty in natural selection making an occasional habit permanent, if of advantage to the species, and if the insect whose nest and stored food are thus feloniously appropriated, be not thus exterminated.

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, who had ample opportunities for observation, 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. 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.

I will not here enter on minute details on this subject, but will merely give an outline of the conclusions at which I have arrived. 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 shown, I think, to follow from a few very simple instincts.

I was led to investigate this subject by Mr. Waterhouse, who has shown that the form of the cell stands in close relation to the presence of adjoining cells; and the following view may, perhaps, be considered only as a modification of his theory. Let us look to the great principle of gradation, and see whether Nature does not reveal to us her method of work. At one end of a short series we have humble-bees, which use their old cocoons to hold honey, sometimes adding to them short tubes of wax, and likewise making separate and very irregular rounded cells of wax. At the other end of the series we have the cells of the hive-bee, placed in a double layer: each cell, as is well known, is an hexagonal prism, with the basal edges of its six sides bevelled so as to join on to a pyramid, formed of three rhombs. These rhombs have certain angles, and the three which form the pyramidal base of a single cell on one side of the comb, enter into the composition of the bases of three adjoining cells on the opposite side. In the series between the extreme perfection of the cells of the hive-bee and the simplicity of those of the humble-bee, we have the cells of the Mexican Melipona domestica, carefully described and figured by Pierre Huber. The Melipona itself is intermediate in structure between the hive and humble bee, but more nearly related to the latter: it forms a nearly regular waxen comb of cylindrical cells, in which the young are hatched, and, in addition, some large cells of wax for holding honey. These latter cells are nearly spherical and of nearly equal sizes, and are aggregated into an irregular mass. But the important point to notice, is that these cells are always made at that degree of nearness to each other, that they would have intersected or broken into each other, if the spheres had been completed; but this is never permitted, the bees building perfectly flat walls of wax between the spheres which thus tend to intersect. Hence each cell consists of an outer spherical portion and of two, three, or more perfectly flat surfaces, according as the cell adjoins two, three or more other cells. When one cell comes into contact with three other cells, which, from the spheres being nearly of the same size, is very frequently and necessarily the case, the three flat surfaces are united into a pyramid; and this pyramid, as Huber has remarked, is manifestly a gross imitation of the three-sided pyramidal basis of the cell of the hive-bee. As in the cells of the hive-bee, so here, the three plane surfaces in any one cell necessarily enter into the construction of three adjoining cells. It is obvious that the Melipona saves wax by this manner of building; for the flat walls between the adjoining cells are not double, but are of the same thickness as the outer spherical portions, and yet each flat portion forms a part of two cells.