Charles Robert Darwin (1809–1882). Origin of Species.
The Harvard Classics. 1909–14.
Special Instincts
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Instincts of the Cuckoo.—It is supposed by some naturalists that the more immediate cause of the instinct of the cuckoo 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 migrates 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 both asserted and denied that the American cuckoo occasionally lays her eggs in other birds’ nests; but I have lately heard from Dr. Merrell, of Iowa, that he once found in Illinois a young cuckoo together with a young jay in the nest of a Blue
It has been objected that I have not noticed other related instincts and adaptations of structure in the cuckoo, which are spoken of as necessarily co-ordinated. But in all cases, speculation on an instinct known to us only in a single species, is useless, for we have hitherto had no facts to guide us. Until recently the instincts of the European and of the nonparasitic American cuckoo alone were known. now, owing to Mr. Ramsay’s observations, we have learnt something about three Australian species, which lay their eggs in other birds’ nests. The chief points to be referred to are three: first, that the common cuckoo, with rare exceptions, lays only one egg in a nest, so that the large and voracious young bird receives ample food. Secondly, that the eggs are remarkably small, not exceeding those of the skylark,—a bird about one-fourth
Turning now to the Australian species; though these birds generally lay only one egg in a nest, it is not rare to find two or even three eggs in the same nest. In the bronze cuckoo the eggs vary greatly in size, from eight to ten times in length. Now if it had been of an advantage to this species to have laid eggs even smaller than those now laid, so as to have deceived certain foster-parents, or, as is more probable, to have been hatched within a shorter period (for it is asserted that there is a relation between the size of eggs and the period of their incubation), then there is no difficulty in believing that a race or species might have been formed which would have laid smaller and smaller eggs; for these would have been more safely hatched and reared. Mr. Ramsay remarks that two of the Australian cuckoos, when they lay their eggs in an open nest, manifest a decided preference for nests containing eggs similar in colour to their own. The European species apparently manifests some tendency towards a similar instinct, but not rarely departs from it, as is shown by her laying her dull and pale-coloured eggs in the nest of the Hedge-warbler with bright greenish-blue eggs. Had our cuckoo invariably displayed the above instinct, it would assuredly have been added to those which it is assumed must all have been acquired together. The eggs of the Australian bronze cuckoo vary, according to Mr. Ramsay, to an extraordinary degree in colour; so that in this respect, as well as in size, natural selection might have secured and fixed any advantageous variation.
In the case of the European cuckoo, the offspring of the foster-parents are commonly ejected from the nest within three days after the cuckoo is hatched; and as the latter at
Some species of Molothrus, a widely distinct genus of American birds, allied to our starlings, have parasitic habits like those of the cuckoo; and the species present an interesting gradation in the perfection of their instincts. The sexes of Molothrus badius are stated by an excellent observer, Mr. Hudson, sometimes to live promiscuously together in flocks, and sometimes to pair. They either build a nest of their own, or seize on one belonging to some other bird, occasionally throwing out the nestlings of the stranger. They either lay
Various birds, as has already been remarked, occasionally lay their eggs in the nest of other birds. This habit is not very uncommon with the Gallinaceæ, and throws some light on the singular instinct of the ostrich. In this family several hen-birds 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 with the cuckoo, at intervals of two or three days. The instinct, however, of the American ostrich, as in the case of the Molothrus bonariensis, has not
Many bees are parasitic, and regularly lay their eggs in the nests of other kinds of bees. This case is more remarkable than that of the cuckoo; for these bees have not only had their instincts but their structure modified in accordance with their parasitic habits; for they do not possess the pollen-collecting apparatus which would have been indispensable if they had stored up food for their own young. Some species of Sphegidea (wasp-like insects) are likewise parasitic; 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 larvæ, yet that, when this insect finds a burrow already made and stored by another species, it takes advantage of the prize and becomes for the occasion parasitic. In this case, as with that of the Molothrus or 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 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 female do no work of any kind, and 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 larvæ. 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 own larvæ and pupæ 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
Another species, 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 existence of so extraordinary an instinct as that of making slaves. Hence, I will give the observations which I 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 (F. fusca) 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 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 larvæ and pupæ are exposed, the slaves work energetically together 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 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.
One day I fortunately witnessed a migration of F. sanguinea from one nest to another, and it was a most interesting spectacle to behold the masters carefully carrying their slaves in their jaws instead of being carried by them, as in the case of F. rufescens. 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 slavemaking 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 pupæ to rear as slaves. I then dug up a small parcel of the pupæ 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 pupæ of another species, F. flava, with a few of these little yellow ants still clinging to the fragments of their 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
One evening I visited another community of F. sanguinea, and found a number of these ants returning home and entering their nests, carrying the dead bodies of F. fusca (showing that it was not a migration) and numerous pupæ. I traced a long file of ants burthened with booty, for about forty yards back, 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, an image of despair 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 continental 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
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 the pupæ of other species, if scattered near their nests, it is possible that such pupæ originally stored as food might become developed; and the foreign 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 pupæ, 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, natural selection might increase and modify 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
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 this 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 an inverted pyramid, 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
Reflecting on this case, it occurred to me that if the Melipona had made its spheres at some given distance from each other, and had made them of equal sizes and had arranged them symmetrically in a double layer, the resulting structure would have been as perfect as the comb of the hive-bee. Accordingly I wrote to Professor Miller of Cambridge, and this geometer has kindly read over the following statement, drawn up from his information, and tells me that it is strictly correct:—
If a number of equal squares be described with their centres placed in two parallel layers; with the centre of each sphere at the distance of radius x square root of 2 , or radius x 1.41421 (or at some lesser distance), from the centres of the six surrounding spheres in the same layer; and at the same distance from the centres of the adjoining spheres in the other and parallel layer; then, if planes of intersection between the several spheres in both layers be formed, there will result a double layer of hexagonal prisms united together by pyramidal bases formed of three rhombs; and the rhombs and the sides of the hexagonal prisms will have every angle identically the same with the best measurements which have been made of the cells of the hive-bee. But I hear from
Hence we may safely conclude that, if we could slightly modify the instincts already possessed by the Melipona, and in themselves not very wonderful, this bee would make a structure as wonderfully perfect as that of the hive-bee. We must suppose that Melipona to have the power of forming her cells truly spherical, and of equal sizes, and this would not be very surprising, seeing that she already does so to a certain extent, and seeing what perfectly cylindrical burrows many insects make in wood, apparently by turning round on a fixed point. We must suppose the Melipona to arrange her cells in level layers, as she already does her cylindrical cells; and we must further suppose, and this is the greatest difficulty, that she can somehow judge accurately at what distance to stand from her fellow-labourers when several are making their spheres; but she is already so far enabled to judge of distance, that she always describes her spheres so as to intersect to a certain extent; and then she unites the points of intersection by perfectly flat surfaces. By such modifications of instincts which in themselves are not very wonderful,—hardly more wonderful than those which guide a bird to make its nest,—I believe that the hive-bee has acquired, through natural selection, her inimitable architectural powers.
But this theory can be tested by experiment. Following the example of Mr. Tegetmeier, I separated two combs, and put between them a long, thick, rectangular strip of wax: the bees instantly began to excavate minute circular pits in it; and as they deepened these little pits, they made them wider and wider until they were converted into shallow basins, appearing to the eye perfectly true or parts of a sphere, and of about the diameter of a cell. It was most interesting to observe that, wherever several bees had begun to excavate these basins near together, they had begun their work at such a distance from each other, that by the time the basins had acquired the above stated width (i. e. about the width of an ordinary cell), and were in depth
I then put into the hive, instead of a thick, rectangular piece of wax, a thin and narrow, knife-edged ridge, coloured with vermilion. The bees instantly began on both sides to excavate little basins near to each other, in the same way as before; but the ridge of wax was so thin, that the bottoms of the basins, if they had been excavated to the same depth as in the former experiment, would have broken into each other from the opposite sides. The bees, however, did not suffer this to happen, and they stopped their excavations in due time; so that the basins, as soon as they had been a little deepened, came to have flat bases; and these flat bases, formed by thin little plates of the vermilion wax left ungnawed, were situated, as far as the eye could judge, exactly along the planes of imaginary intersection between the basins on the opposite sides of the ridge of wax. In some parts, only small portions, in other parts, large portions of a rhombic plate were thus left between the opposed basins, but the work, from the unnatural state of things, had not been neatly performed. The bees must have worked at very nearly the same rate in circularly gnawing away and deepening the basins on both sides of the ridge of vermilion wax, in order to have thus succeeded in leaving flat plates between the basins, by stopping work at the planes of intersection.
Considering how flexible thin wax is, I do not see that there is any difficulty in the bees, whilst at work on the two sides of a strip of wax, perceiving when they have gnawed the wax away to the proper thinness, and then stopping their work. In ordinary combs it has appeared to me that the bees do not always succeed in working at exactly the same rate from the opposite sides; for I have noticed half-completed rhombs at the base of a just commenced cell, which were slightly concave on one side, where I suppose that the
From the experiment of the ridge of vermilion wax we can see that, if the bees were to build for themselves a thin wall of wax, they could make their cells of the proper shape, by standing at the proper distance from each other, by excavating at the same rate, and by endeavouring to make equal spherical hollows, but never allowing the spheres to break into each other. Now bees, as may be clearly seen by examining the edge of a growing comb, do make a rough, circumferential wall or rim all round the comb; and they gnaw this away from the opposite sides, always working circularly as they deepen each cell. They do not make the whole three-sided pyramidal base of any one cell at the same time, but only that one rhombic plate which stands on the extreme growing margin, or the two plates, as the case may be; and they never complete the upper edges of the rhombic plates, until the hexagonal walls are commenced. Some of these statements differ from those made by the justly celebrated elder Huber, but I am convinced of their accuracy; and if I had space, I would show that they are conformable with my theory.
Huber’s statement that the very first cell is excavated out of a little parallel-sided wall of wax, is not, as far as I have seen, strictly correct; the first commencement having always been a little hood of wax; but I will not here enter on details. We see how important a part excavation plays in the construction of the cells; but it would be a great error to suppose that the bees cannot build up a rough wall of wax in the proper position—that is, along the plane of intersection
It seems at first to add to the difficulty of understanding how the cells are made, that a multitude of bees all work together; one bee after working a short time at one cell going to another, so that, as Huber has stated, a score of individuals work even at the commencement of the first cell. I was able practically to show this fact, by covering the edges of the hexagonal walls of a single cell, or the extreme margin of the circumferential rim of a growing comb, with an extremely thin layer of melted vermilion wax; and I invariably found that the colour was most delicately diffused
When bees have a place on which they can stand in their proper positions for working,—for instance, on a slip of wood, placed directly under the middle of a comb growing downwards, so that the comb has to be built over one face of the slip—in this case the bees can lay the foundations of one wall of a new hexagon, in its strictly proper place, projecting beyond the other completed cells. It suffices that the bees should be enabled to stand at their proper relative distances from each other and from the walls of the last completed cells, and then, by striking imaginary spheres, they can build up a wall intermediate between two adjoining spheres; but, as far as I have seen, they never gnaw away and finish off the angles of a cell till a large part both of that cell and of the adjoining cells has been built. This capacity in bees of laying down under certain circumstances a rough wall in its proper place between two just-commenced cells, is important, as it bears on a fact, which seems at first subversive of the foregoing theory; namely, that the cells on the extreme margin of wasp-combs are sometimes strictly hexagonal; but I have not space here to enter on this subject. Nor does there seem to me any great difficulty in a single insect (as in the case of a queen-wasp) making hexagonal cells, if she were to work alternately on the inside and outside of two or three cells commenced at the same time, always standing at the proper relative distance from the parts of the cells just begun, sweeping spheres or cylinders, and building up intermediate planes.
Thus, as I believe, the most wonderful of all known instincts, that of the hive-bee, 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 the construction of cells of due strength and of the proper size and shape for the larvæ, this being effected with the greatest possible economy of labour and wax; that individual swarm which thus made the best cells with least labour, and least waste of honey in the secretion of wax, having succeeded best, and having transmitted their newly-acquired economical instincts to new swarms, which in their turn will have had the best chance of succeeding in the struggle for existence.