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07 - Instinct 07-05 - Domestic instincts, their origin 50 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.

pointer
pointer

Golden Retriever
Golden Retriever

German Shepherd
German Shepherd


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.

White Butterfly
White Butterfly

cabbage
cabbage



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.

wolf
wolf

deer
deer


Domestic instincts, as they may be called, are certify 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.

bulldog
bulldog

greyhound
greyhound

German Shepherd
German Shepherd

hare
hare


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.

dog
dog

wolf
wolf


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.

Tumbler Pigeon
Tumbler Pigeon


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.

pointer
pointer

terrier
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.

Wild Rabbit
Wild Rabbit

rabbit
rabbit


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.

fowl
fowl


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.

dog
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.

wolf
wolf

fox
fox

jackal
jackal

cat
cat

poultry
poultry

sheep
sheep

Tierra del Fuego
Tierra del Fuego

Australia
Australia


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.

chicken
chicken

dog
dog

cat
cat

pheasant
pheasant

hen
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.

Turkey Cock
Turkey Cock

grass
grass


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.
07 - Instinct 07-06 - Natural instincts of the cuckoo, ostrich, and parasitic bees 60 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.

cuckoo
cuckoo

ant
ant

Honey Comb
Honey Comb


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.

gallinacea
gallinacea

ostrich
ostrich


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.

cuckoo
cuckoo


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.

bee
bee


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.

sphex
sphex

Tachytes Nigra
Tachytes Nigra


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.

cuckoo
cuckoo
07 - Instinct 07-07 - Slave-making ants 70 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.

Polyerges Rufescens
Polyerges Rufescens


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.

F. Fusca
F. Fusca


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.

Formica Sanguinea
Formica Sanguinea


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.

England
England


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 made 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 their 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.

Sussex
Sussex


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.

Scotch Fir
Scotch Fir

aphid
aphid


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.

Switzerland
Switzerland


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.

England
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.

Formica Sanguinea
Formica Sanguinea

F. Fusca
F. Fusca


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.

heath
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.

Formica Sanguinea
Formica Sanguinea

Polyerges Rufescens
Polyerges 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.

aphid
aphid


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.

England
England

Switzerland
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.
07 - Instinct 07-08 - Hive-bee, its cell-making instinct 80 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.

Hive Bee
Hive Bee

Honey Comb
Honey Comb


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 previous wax in their construction.
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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.
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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.
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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 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.

Humble Bee
Humble Bee

cocoon
cocoon

wax
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 know, 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.

rhomb
rhomb


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.

Mexican Melipona Domestica
Mexican Melipona Domestica

Mexican Melipona Domestica Comb
Mexican Melipona Domestica Comb


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.

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 probably 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 spheres be described with their centres placed in two parallel layers; with the centre of each sphere at the distance of radius X /sqrt[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.
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wax
wax

honey
honey


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 the Melipona to make 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 in wood many insects can make, 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 largely; and then she unites the points of intersection by perfectly flat surfaces.

We have further to suppose, but this is no difficulty, that after hexagonal prisms have been formed by the intersection of adjoining spheres in the same layer, she can prolong the hexagon to any length requisite to hold the stock of honey; in the same way as the rude humble-bee adds cylinders of wax to the circular mouths of her old cocoons.

By such modifications of instincts in themselves 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.
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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, square 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.
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It was most interesting to me 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 about one sixth of the diameter of the sphere of which they formed a part, the rims of the basins intersected or broke into each other.

As soon as this occurred, the bees ceased to excavate, and began to build up flat walls of wax on the lines of intersection between the basins, so that each hexagonal prism was built upon the festooned edge of a smooth basin, instead of on the straight edges of a three-sided pyramid as in the case of ordinary cells.

I then put into the hive, instead of a thick, square piece of wax, a thin and narrow, knife-edged ridge, coloured with vermilion.

wax
wax

vermilion
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 bottoms; and these flat bottoms, formed by thin little plates of the vermilion wax having been 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 parts, only little bits, in other parts, large portions of a rhombic plate had been 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 on the opposite side of the ridge of vermilion wax, as they circularly gnawed away and deepened the basins on both sides, in order to have succeeded in thus leaving flat plates between the basins, by stopping work along the intermediate planes or 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 bees had excavated too quickly, and convex on the opposed side, where the bees had worked less quickly.

In one well-marked instance, I put the comb back into the hive and allowed the bees to go on working for a short time and again examined the cell, and I found that the rhombic plate had been completed, and had become perfectly flat: it was absolutely impossible, from the extreme thinness of the little rhombic plate, that they could have affected this by gnawing away the convex side; and I suspect that the bees in such cases stand in the opposed cells and push and bend the ductile and warm wax (which as I have tried is easily done) into its proper intermediate plane, and thus flatten it.

From the experiment of the ridge of vermilion wax, we can clearly 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 into this 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 the 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 could 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 these 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 between two adjoining spheres.

I have several specimens showing clearly that they can do this.

Even in the rude circumferential rim or wall of wax round a growing comb, flexures may sometimes be observed, corresponding in position to the planes of the rhombic basal plates of future cells.

But the rough wall of wax has in every case to be finished off, by being largely gnawed away on both sides. The manner in which the bees build is curious; they always make the first rough wall from ten to twenty times thicker than the excessively thin finished wall of the cell, which will ultimately be left.

We shall understand how they work, by supposing masons first to pile up a broad ridge of cement, and then to begin cutting it away equally on both sides near the ground, till a smooth, very thin wall is left in the middle; the masons always piling up the cut-away cement, and adding fresh cement, on the summit of the ridge.

We shall thus have a thin wall steadily growing upward; but always crowned by a gigantic coping.

From all the cells, both those just commenced and those completed, being thus crowned by a strong coping of wax, the bees can cluster and crawl over the comb without injuring the delicate hexagonal walls, which are only about one four-hundredth of an inch in thickness; the plates of the pyramidal basis being about twice as thick.

By this singular manner of building, strength is continually given to the comb, with the utmost ultimate economy of wax.

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 by the bees as delicately as a painter could have done with his brush by atoms of the coloured wax having been taken from the spot on which it had been placed, and worked into the growing edges of the cells all round.

wax
wax

cinnabar
cinnabar

atom
atom


The work of construction seems to be a sort of balance struck between many bees, all instinctively standing at the same relative distance from each other, all trying to sweep equal spheres, and then building up, or leaving ungnawed, the planes of intersection between these spheres. It was really curious to note in cases of difficulty, as when two pieces of comb met at an angle, how often the bees would entirely pull down and rebuild in different ways the same cell, sometimes recurring to a shape which they had at first rejected.

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 quite 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.

wasp
wasp

Wasp Comb
Wasp Comb


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 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.

Queen Wasp
Queen Wasp


It is even conceivable that an insect might, by fixing on a point at which to commence a cell, and then moving outside, first to one point, and then to five other points, at the proper relative distances from the central point and from each other, strike the planes of intersection, and so make an isolated hexagon: but I am not aware that any such case has been observed; nor would any good be derived from a single hexagon being built, as in its construction more materials would be required than for a cylinder.

As natural selection acts only by the accumulation of slight modifications of structure or instinct, each profitable to the individual under its conditions of life, it may reasonably be asked, how a long and graduated succession of modified architectural instincts, all tending towards the present perfect plan of construction, could have profited the progenitors of the hive-bee?

I think the answer is not difficult: it is known that bees are often hard pressed to get sufficient nectar; and I am informed by Mr. Tegetmeier that it has been experimentally found that no less than from twelve to fifteen pounds of dry sugar are consumed by a hive of bees for the secretion of each pound of wax; so that a prodigious quantity of fluid nectar must be collected and consumed by the bees in a hive for the secretion of the wax necessary for the construction of their combs.

Moreover, many bees have to remain idle for many days during the process of secretion.

A large store of honey is indispensable to support a large stock of bees during the winter; and the security of the hive is known mainly to depend on a large number of bees being supported. Hence the saving of wax by largely saving honey must be a most important element of success in any family of bees.

nectar
nectar

wax
wax

Honey
Honey


Of course the success of any species of bee may be dependent on the number of its parasites or other enemies, or on quite distinct causes, and so be altogether independent of the quantity of honey which the bees could collect.

But let us suppose that this latter circumstance determined, as it probably often does determine, the numbers of a humble-bee which could exist in a country; and let us further suppose that the community lived throughout the winter, and consequently required a store of honey: there can in this case be no doubt that it would be an advantage to our humble-bee, if a slight modification of her instinct led her to make her waxen cells near together, so as to intersect a little; for a wall in common even to two adjoining cells, would save some little wax.

Hence it would continually be more and more advantageous to our humble-bee, if she were to make her cells more and more regular, nearer together, and aggregated into a mass, like the cells of the Melipona; for in this case a large part of the bounding surface of each cell would serve to bound other cells, and much wax would be saved.

melipona
melipona


Again, from the same cause, it would be advantageous to the Melipona, if she were to make her cells closer together, and more regular in every way than at present; for then, as we have seen, the spherical surfaces would wholly disappear, and would all be replaced by plane surfaces; and the Melipona would make a comb as perfect as that of the hive-bee.

Beyond this stage of perfection in architecture, natural selection could not lead; for the comb of the hive-bee, as far as we can see, is absolutely perfect in economising wax.

architect
architect


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 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.