CHAPTER III.

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ANTS.
Within the last ten or twelve years our information on the habits and intelligence of these insects has been so considerably extended, that in here rendering a condensed epitome of our knowledge in this most interesting branch of comparative psychology, it will be found that the chapter is constituted principally of a statement of observations and experiments which have been conducted during the short period named. The observers to whom we are mainly indebted for this large increase of our knowledge are Messrs. Bates, Belt, MÜller, Moggridge, Lincecum, MacCook, and Sir John Lubbock. From the fact that these naturalists conducted their observations in different parts of the world and on widely different species of ants, it is not surprising that their results should present many points of difference; for this only shows, as we might have expected, that different species of ants differ considerably in habits and intelligence. Therefore, in now drawing all these numerous observations to a focus, I shall endeavour to show clearly their points of difference as well as their points of agreement; and in order that the facts to be considered may be arranged in some kind of order, I shall deal with them under the following heads:—Powers of special sense; Sense of direction; Powers of memory; Emotions; Powers of communication; Habits general in sundry species; Habits peculiar to certain species; General intelligence of various species.

Powers of Special Sense.

Taking first the sense of sight, Sir John Lubbock made a number of experiments on the influence of light coloured by passing through various tints of stained glass, with the following results. The ants which he observed greatly dislike the presence of light within their nests, hurrying about in search of the darkest corners when light is admitted. The experiments showed that the dislike is much greater in the case of some colours than in that of others. Thus under a slip of red glass there were congregated on one occasion 890 ants, under green 544, under yellow 495, and under violet only 5. To our eyes the violet is as opaque as the red, more so than the green, and much more so than the yellow. Yet, as the numbers show, the ants had scarcely any tendency to congregate under it: there were nearly as many under the same area of the uncovered portion of the nest as under that shaded by the violet glass. It is curious that the coloured glasses appear to act on the ants in a graduated series, which corresponds with the order of their influence on a photographic plate. Experiments were therefore made to test whether it might not be the actinic rays that were so particularly distasteful to the ants; but with negative results. Placing violet glass above red produces the same effect as red glass alone. Obviously, therefore, the ants avoid the violet glass because they dislike the rays which it transmits, and do not prefer the other colours because they like the rays which they transmit. Sodium, barium, strontium, and lithium flames were also tried, but not with so much effect as the coloured glass.

It has just been observed that the relative dislike which Sir John Lubbock's ants showed to lights of different colours seems to be determined by the position of the colour in the spectrum—there being a regular gradation of intolerance shown from the red to the violet end. As these ants dislike light, the question suggests itself that the reason of their graduated intolerance to light of different colours may be due to their eyes not being so much affected by the rays of low as by those of high refrangibility. In this connection it would be interesting to ascertain whether ants of the genus Atta show a similarly graduated intolerance to the light in different parts of the spectrum; for both Moggridge and MacCook record of this genus that it not only does not shun the light, but seeks it—coming to the glass sides of their artificial nests to enjoy the light of a lamp. Possibly, therefore, the scale of preference to lights of different colours would be found in this genus to be the reverse of that which Sir John Lubbock has found in the case of the British species.

As regards hearing, Sir John Lubbock found that sounds of various kinds do not produce any effect upon the insects. Tuning-forks and violin notes, shouting, whistling, &c., were all equally inefficient in producing the slightest influence upon the animals; and experiments with sensitive flames, microphone, telephone, &c., failed to yield any evidence of ants emitting sounds inaudible to human ears.

Lastly, as regards the sense of smell, Sir John Lubbock found that on bringing a camel's-hair brush steeped in various strong scents near where ants were passing, "some went on without taking any notice, but others stopped, and evidently perceiving the smell, turned back. Soon, however, they returned, and passed the scented pencil. After doing this two or three times, they generally took no further notice of the scent. This experiment left no doubt on my mind." In other cases the ants were observed to wave about and throw back their antennÆ when the scented pencil was brought near.

That ants track one another by scent was long ago mentioned by Huber, and also that they depend on this sense for their power of finding supplies which have been previously found by other ants. Huber proved their power of tracking a path previously pursued by their friends, by drawing his finger across the trail, so obliterating the scent at that point, and observing that when the ants arrived at that point they became confused and ran about in various directions till they again came upon the trail on the other side of the interrupted space, when they proceeded on their way as before. The more numerous and systematic experiments of Sir John Lubbock have fully corroborated Huber's observations, so far as these points are concerned. Thus, to give only one or two of these experiments; in the accompanying woodcut (Fig. 1) A is the nest, B a board, n f g slips of paper, h and m similar slides of glass, on one of which, h, there was placed pupÆ, while the other, m, was left empty. Sir John Lubbock watched two particular (marked) ants proceeding from A to h and back again, carrying the pupÆ on h to the nest A. Whenever an ant came out of A upon B he transposed the slips f and g. Therefore at the angle below n there was a choice presented to the ant of taking the unscented pathway leading to the full glass h, or the scented pathway leading to the empty glass m. The two marked ants, knowing their way, always took the right turn at the angle; but the stranger ants, being guided only by scent, for the most part took the wrong turn at the angle, so going to the empty glass m. For out of 150 stranger ants only 21 went to h, while the remaining 129 went to m. Still the fact that all the stranger ants did not follow the erroneous scent-trail to m, may be taken to indicate that they are also assisted in finding treasure by the sense of sight, though in a lesser degree. Therefore Sir John Lubbock concludes that in finding treasure 'they are guided in some cases by sight, while in others they track one another by scent.'

black drawing Fig. 2.

As further evidence showing how much more ants depend upon scent than upon sight in finding their way, the following experiment may be quoted. In the accompanying woodcut (Fig. 2) the line marked 1, 2, 3 represents the edge of a paper bridge leading to the nest; A the top of a pencil which is standing perpendicularly upon a board, represented by the general black surface; B the top of the same pencil when moved a distance of a few inches from its first position A. On the top of this pencil were placed some pupÆ. Sir John Lubbock, after contriving this arrangement, marked an ant and put it upon the pupÆ on the top of the pencil. After she had made two journeys carrying pupÆ from the pencil to the nest (the tracks she pursued being represented by the two thick white lines), while she was in the nest he moved the pencil to its position at B. The thin white line represents the course then pursued by the ant in its endeavours to find the pencil, which was shifted only a few inches from A to B. That is, 'the ants on their journey to the shifted object travelled very often backwards and forwards and round the spot where the coveted object first stood. Then they would retrace their steps towards the nest, wander hither and thither from side to side between the nest and the point A, and only after very repeated efforts around the original site of the larvÆ reach, as it were, accidentally the object desired at B.' Therefore the ants were clearly not guided by the sight of the pencil.

diagram: Heavy dash marks on either side of a dashed vertical line Fig. 3.
Diagram tilted to the left, dark parallel lines now vertical with diash lined diagonal Fig. 4.
Diagram straight again, dashed line vertical with dark parallel dashes forming a 45 degree angle on the left side, top down Fig. 5.

The same thing is well shown by another form of experiment. 'Some food was placed at the point a (Figs. 3 and 4) on a board measuring 20 inches by 12 inches, and so arranged that the ants in going straight from it to the nest would reach the board at the point b, and after passing under the paper tunnel c, would proceed between five pairs of wooden bricks, each 3 inches in length and 1¾ inches in height. When they got to know their way they went quite straight along the line d e to a. The board was then twisted as shown in Fig. 4. 'The bricks and tunnel being arranged exactly in the same direction as before, but the board having been moved, the line d e was now outside them. The change, however, did not at all discompose the ants; but instead of going, as before, through the tunnel and between the rows of bricks to a, they walked exactly along the old path to e.' Keeping the board steady, but moving the brick pathway to the left-hand corner of the board where the food was next placed (Fig. 5), had the effect of making the ant first go to the old position of the food at a, whence it veered to a new position, which we may call x. The bricks and food were then moved towards the right-hand corner of the board—i.e. over a distance of 8 inches (Fig. 6). The ant now first went to a, then to x, and not finding the food at either place, set to work to look for it at random, and was only successful after twenty-five minutes' wandering.

The same as the last diagram but with the dark parallel dashes forming angle on the right, top down Fig. 6.

And, as evidence how much more dependence they place upon scent in finding their way than upon any other of their faculties, it is desirable to quote yet one further experiment, which is of great interest as showing that when their sense of smell is made to contradict their sense of direction, they follow the former, notwithstanding, as we shall presently see, the wonderful accuracy of the information which is supplied to them by the latter. 'If, when F. niger were carrying off larvÆ placed in a cup on a piece of board, I turned the board round so that the side which had been turned towards the nest was away from it, and vice versÂ, the ants always returned over the same track on the board, and, in consequence, directly away from home. If I moved my board to the other side of my artificial nest, the result was the same. Evidently they followed the road, not the direction.'

There can be little doubt that ants have a sense of taste, as they are so well able to distinguish sugary substances; and it is unquestionable that in their antennÆ they possess highly elaborated organs of touch.

Sense of Direction.

As evidence of the accuracy and importance of the sense of direction in the Hymenoptera, we must here adduce Sir John Lubbock's highly interesting experiments on ants—leaving his experiments in this connection on bees and wasps to be considered in the next chapter. He first accustomed some ants (Lasius niger) to go to and fro to food over a wooden bridge. When they had got quite accustomed to the way, he watched when an ant was upon a bridge which could be rotated, and while she was passing along it, he turned it round, so that end b was at c, and c at b. 'In most cases the ant immediately turned round also; but even if she went on to b or c, as the case might be, as soon as she came to the end of the bridge she turned round.' Next, between the nest and the food he placed a hat-box twelve inches in diameter and seven inches high, cutting two small holes, so that the ants in passing from the nest to the food had to pass in at one hole and out at the other. The box was fixed upon a central pivot, so as to admit of being rotated easily without much friction or disturbance. When the ants had well learnt their way, the box was turned half round as soon as an ant had entered it, 'but in every case the ant turned too, thus retaining her direction.' Lastly, Sir John took a disk of white paper, which he placed in the stead of the hat-box between the nest and the food. When an ant was on the disk making towards the food, he gently drew the disk to the other side of the food, so that the ant was conveyed by the moving surface in the same direction as that in which she was going, but beyond the point to which she intended to go. Under these circumstances 'the ant did not turn round, but went on' to the further edge of the disk, when she seemed 'a good deal surprised at finding where she was.'

These experiments seem to show that the mysterious 'sense of direction,' and consequent faculty of 'homing,' are in ants, at all events, due to a process of registering, and, where desirable, immediately counteracting any change of direction, even when such change is gently made by a wholly closed chamber in which the animal is moving, and not by any muscular movements of the animal itself. And the fact that drawing the moving surface along in the same direction of advance as that which the insect is pursuing does not affect the movements of the latter, seems conclusively to show that the power of registration has reference only to lateral movements of the travelling surface; it has no reference to variations in the velocity of advance along the line in which the animal is progressing.[19]

Powers of Memory.

Little need here be said to prove that ants display some powers of memory; for many of the observations and experiments already detailed constitute a sufficient demonstration of the statement that they do. Thus, for instance, the general fact that whenever an ant finds her way to a store of food or larvÆ, she will return to it again and again in a more or less direct line from her nest, constitutes ample proof that the ant remembers the way to the store. It is of considerable interest, however, to note that the nature of this insect-memory appears to be, as far as it goes, precisely identical with that of memory in general. Thus, a new fact becomes impressed upon their memory by repetition, and the impression is liable to become effaced by lapse of time. More evidence on both these features of insect-memory will be adduced when we come to treat of the intelligence of bees; but meanwhile it is enough to refer to the fact that in his experiments on ants, Sir John Lubbock found it necessary to teach the insects by a repetition of several lessons their way to treasure, if that way was long or unusual.

With regard to the duration of memory, it does not appear that any experiments have been made; but the following observation by Mr. Belt on this point in the case of the leaf-cutting ant may here be stated. In June 1859 he found his garden invaded by these ants, and following up their paths he found their nest about a hundred yards distant. He poured down their burrows a pint of common brown carbolic acid, mixed with four buckets of water. The marauding parties were at once drawn off from the garden to meet the danger at home, and the whole formicarium was disorganised, the ants running up and down again in the utmost perplexity. Next day he found them busily employed bringing up the ant-food from the old burrows, and carrying it to newly formed ones a few yards distant. These, however, turned out to be only intended as temporary repositories; for in a few days both the old and the new burrows were entirely deserted, so that he supposed all the ants to have died. Subsequently, however, he found that they had migrated to a new site, about two hundred yards from the old one, and there established themselves in a new nest. Twelve months later the ants again invaded his garden, and again he treated them to a strong dose of carbolic acid. The ants, as on the previous occasion, were at once withdrawn from the garden, and two days afterwards he found 'all the survivors at work on one track that led directly to the old nest of the year before, where they were busily employed in making fresh excavations. Many were bringing along pieces of ant-food' from the nest most recently deluged with carbolic acid to that which had been similarly deluged a year before, and from which all the carbolic acid had long ago disappeared. 'Others carried the undeveloped white pupÆ and larvÆ. It was a wholesale and entire migration;' and the next day the nest down which he had last poured the carbolic acid was entirely deserted. Mr. Belt adds: 'I afterwards found that when much disturbed, and many of the ants destroyed, the survivors migrate to a new locality. I do not doubt that some of the leading minds in this formicarium recollected the nest of the year before, and directed the migration to it.'

Now, I do not insist that the facts necessarily point to this conclusion; for it may have been that the leaders of the migration simply stumbled upon the old and vacant nest by accident, and finding it already prepared as a nest, forthwith proceeded to transfer the food and pupÆ to it. Still, as the two nests were separated from one another by so considerable a distance, this hypothesis does not seem probable, and the only other one open to us is that the ants remembered the site of their former home for a period of twelve months. And this conclusion is rendered less improbable from a statement of Karl Vogt in his 'Thierstaaten,' to the effect that for several successive years ants from a certain nest used to go through certain inhabited streets to a chemist's shop 600 mÈtres distant, in order to obtain access to a vessel filled with syrup. As it cannot be supposed that this vessel was found in successive working seasons by as many successive accidents, it can only be concluded that the ants remembered the syrup store from season to season.

I shall now pass on to consider a class of highly remarkable facts, perhaps the most remarkable of the many remarkable facts connected with ant psychology.

It has been known since the observations of Huber that all the ants of the same nest or community recognise one another as friends, while an ant introduced from another nest, even though it be an ant of the same species, is known at once to be a foreigner, and is usually maltreated or put to death. Huber found that when he removed an ant from a nest and kept it away from its companions for a period of four months it was still recognised as a friend, and caressed by its previous fellow-citizens after the manner in which ants show friendship, viz., by stroking antennÆ. Sir John Lubbock, after repeating and fully confirming these observations, extended them as follows. He first tried keeping the separated ant away from the nest for a still longer period than four months, and found that even after a separation of more than a year the animal was recognised as before. He repeated this experiment a number of times, and always with the same invariable difference between the reception accorded to a foreigner and a native—no matter, apparently, how long the native had been absent.

Considering the enormous number of ants that go to make a nest, it seems astonishing enough that they should be all personally known to one another, and still more astonishing that they should be able to recognise members of their community after so prolonged an absence. Thinking that the facts could only be explained, either by all the ants in the same nest having a peculiar smell, or by all the members of the same community having a particular pass-word or gesture-sign, Sir John Lubbock, with the view of testing this theory, separated some ants from a nest while still in the condition of pupÆ, and, when they emerged from that state as perfect insects, transferred them back to the nest from which they had been taken as pupÆ. Of course in this case the ants in the nest could never have seen those which had been removed, for a larval ant is as unlike the mature insect as a grub is unlike a beetle; neither can it be supposed that a larva, hatched out away from the nest, should retain, when a perfect insect, any smell belonging to its parent nest—more especially as it had been hatched out by ants in another nest;[20] nor, lastly, is it reasonable to imagine that the animal, while still a larval grub, can have been taught any gesture-signal used as a pass-word by the matured animals. Yet, although all these possible hypotheses seem to be thus fully excluded by the conditions of the experiment, the result showed unequivocally that the ants recognised their transformed larvÆ as native-born members of their community.

Lastly, Sir John Lubbock tried the experiment of going still further back in the life-history of the ants before separating them from the nest. For in September he divided a nest into two halves, each having a queen. At this season there were neither larvÆ nor eggs. The following April both the queens began to lay eggs, and in August—i.e. nearly a year after the original partitioning of the nest—he took some of the ants newly hatched from the pupÆ in one division, and placed them in the other division, and vice versÂ. In all cases these ants were received by the members of the other half of the divided nest as friends, although if a stranger were introduced into either half it was invariably killed. Yet the ants which were thus so certainly recognised by their kindred ants as friends had never, even in the state of an egg, been present in that division of the nest before. On this highly remarkable fact Sir John Lubbock says:—

These observations seem to me conclusive as far as they go, and they are very surprising. In my experiments of last year, though the results were similar, still the ants experimented with had been brought up in the nest, and were only removed after they had become pupÆ. It might therefore be argued that the ants, having nursed them as larvÆ, recognised them when they came to maturity; and though this would certainly be in the highest degree improbable, it could not be said to be impossible. In the present case, however, the old ants had absolutely never seen the young ones until the moment when, some days after arriving at maturity, they were introduced into the nest; and yet in all ten cases they were undoubtedly recognised as belonging to the community.

It seems to me, therefore, to be established by these experiments that the recognition of ants is not personal and individual; that their harmony is not due to the fact that each ant is individually acquainted with every other member of the community.

At the same time, the fact that they recognise their friends even when intoxicated, and that they know the young born in their own nest even when they have been brought out of the chrysalis by strangers, seems to indicate that the recognition is not effected by means of any sign or pass-word.

We must, therefore, conclude with reference to this subject that the mode whereby recognition is undoubtedly effected is as yet wholly unintelligible; and I have introduced these facts under the heading of memory only because this heading is not more inappropriate than any other that could be devised for their reception.

It ought here to be added also that the power of thus recognising members of their community is not confined by the limits of blood-relationship, for in an experiment made by Forel it was shown that Amazon ants recognised their own slaves almost instantaneously after an absence of four months.

Under this heading I may also adduce the evidence as to enormous masses, or, as we might say, a whole nation of ants recognising each other as belonging to the same nationality. New nests often spring up as offshoots from the older ones, and thus a nation of towns gradually spreads to an immense circumference around the original centre. Forel describes a colony of F. exsecta which comprised more than two hundred nests, and covered a space of nearly two hundred square mÈtres. 'All the members of such a colony, even those from the furthermost nest, recognise each other and admit no stranger.'

Similarly, MacCook describes an 'ant town' in the Alleghany Mountains of North America ('Trans. Amer. Entom. Soc.,' Nov. 1877) which was inhabited by F. exsectoÏdes. It consists of 1,600 to 1,700 nests, which rise in cones to a height of from two to five feet. The ground below is riddled in every direction with subterranean passages of communication. The inhabitants are all on the most friendly terms, so that if any one nest is injured it is repaired by their united forces.

It remains to be added in connection with this subject that the recognition is not automatically invariable, but when 'ants are removed from a nest in the pupa state, tended by strangers, and then restored, some at least of their relatives are certainly puzzled, and in many cases doubt their claims to consanguinity. I say some, because while strangers under the circumstances would have been immediately attacked, these ants were in every case amicably received by the majority of the colony, and it was sometimes several hours before they came across one who did not recognise them.'

It may also be added that Lasius flavus behaves towards strangers quite differently and much more hospitably than is the case with L. niger. The stranger shows no alarm, but, on the contrary, will voluntarily enter the strange nest, and she is there received with kindness; although from the attention she excites, and the numerous communications which take place between her and her new friends, Sir John was 'satisfied that they knew she was not one of themselves..... Very different is the behaviour of L. niger under similar circumstances. I tried the same experiment with them. There was no communications with the antennÆ, there was no cleaning, but every ant which the stranger approached flew at her like a little tigress. I tried this experiment four times; each stranger was killed and borne off to the nest.'

Emotions.

The pugnacity, valour, and rapacity of ants are too well and generally known to require the narration of special instances of their display. With regard to the tenderer emotions, however, there is a difference of opinion among observers. Before the researches of Sir John Lubbock it was the prevalent view that these insects display marked signs of affection towards one another, both by caressing movements of their antennÆ, and by showing solicitude for friends in distress. Sir John, however, has found that the species of ants on which he has experimented are apparently deficient both in feelings of affection and of sympathy—or, at least, that such feelings are in these species much less strongly developed than the sterner passions.

He tried burying some specimens of Lasius niger beneath an ant-road; but none of the ants traversing the road made any attempt to release their imprisoned companions. He tried the same experiment with the same result on various other species. Even when the friends in difficulty are actually in sight, it by no means follows that their companions will assist them. Of this, he says, he could give almost any number of instances. Thus, when ants are entangled in honey, their companions devote themselves to the honey, and entirely neglect their friends in distress; and when partly drowned, their friends take no notice. When chloroformed or intoxicated their own companions either do not heed them, or else 'seem somewhat puzzled at finding their intoxicated fellow-creatures in such a condition, take them up, and carry them about for a time in a somewhat aimless manner.' Further experiments, however, on a larger scale, went to show that chloroformed ants were treated as dead, i.e. removed to the edge of the parade-board and dropped over into the surrounding moat of water; while intoxicated ants were generally carried into the nest, if they were ants belonging to that community; if not, they were thrown overboard. This care shown towards intoxicated friends appears to indicate a dim sense of sympathy towards afflicted individuals; but that this emotion or instinct does not in the case of these species extend to healthy individuals in distress seems to be proved, not only by the experiments of burying already described, but also by the following:—

On Sept. 2, therefore, I put two ants from one of my nests of F. fusca into a bottle, the end of which was tied up with muslin as described, and laid it down close to the nest. In a second bottle I put two ants from another nest of the same species. The ants which were at liberty took no notice of the bottle containing their imprisoned friends. The strangers in the other bottle, on the contrary, excited them considerably. The whole day one, two, or more ants stood sentry, as it were, over the bottle. In the evening no less than twelve were collected round it, a larger number than usually came out of the nest at any one time. The whole of the next two days, in the same way, there were more or less ants round the bottle containing the strangers; while, as far as we could see, no notice whatever was taken of the friends. On the 9th the ants had eaten through the muslin, and effected an entrance. We did not chance to be on the spot at the moment; but as I found two ants lying dead, one in the bottle and one just outside, I think there can be no doubt that the strangers were put to death. The friends throughout were quite neglected.

Sept. 21.—I then repeated the experiment, putting three ants from another nest in a bottle as before. The same scene was repeated. The friends were neglected. On the other hand, some of the ants were always watching over the bottle containing the strangers, and biting at the muslin which protected them. The next morning at 6 A.M. I found five ants thus occupied. One had caught hold of the leg of one of the strangers, which had unwarily been allowed to protrude through the meshes of the muslin. They worked and watched, though not, as far as I could see, with any system, till 7.30 in the evening, when they effected an entrance, and immediately attacked the strangers.

Sept. 24.—I repeated the same experiment with the same nest. Again the ants came and sat over the bottle containing the strangers, while no notice was taken of the friends.

The next morning again, when I got up, I found five ants round the bottle containing the strangers, none near the friends. As in the former case, one of the ants had seized a stranger by the leg, and was trying to drag her through the muslin. All day the ants clustered round the bottle, and bit perseveringly, though not systematically, at the muslin. The same thing happened all the following day.

On repeating these experiments with another species (viz., Formica rufescens) the ants took no notice of either bottle, and showed no sign either of affection or hatred. One is almost tempted to surmise that the spirit of these ants is broken by slavery [i.e. by the habit of keeping slaves]. But the experiments on F. fusca seem to show that in these curious insects hatred is a stronger passion than affection.

We must not, however, too readily assent to this general conclusion, that ants as a whole are deficient in the tenderer emotions; for although the case is doubtless so with the species which Sir John examined, it appears to be certainly otherwise with other species, as we shall presently see. But first it may be well to point out that even the hard-hearted species with which Sir John had to do seem not altogether devoid of sympathy with sick or mutilated friends, although they appear to be so towards healthy friends in distress. Thus the care shown to intoxicated friends seems to indicate, if not, as already observed, a dim sense of sympathy, at least an instinct to preserve the life of an ailing citizen for the future benefit of the community. Sir John also quotes some observations of Latreille showing that ants display sympathy with mutilated companions; and, lastly, mentions an instance which he has himself observed of the same thing. A specimen of F. fusca congenitally destitute of antennÆ was attacked and injured by an ant of another species. When separated by Sir John, another ant of her own species came by. 'She examined the poor sufferer carefully, then picked her up tenderly, and carried her away into the nest. It would have been difficult for any one who witnessed this scene to have denied to this ant the possession of humane feelings.' Moggridge is also of opinion that the habit of throwing sick and apparently dead ants into the water, is 'in part to be rid of them, and partly, perhaps, with a view to effecting a possible cure; for I have seen one ant carry another down the twig which formed their path to the surface of the water, and, after dipping it in for a minute, carry it laboriously up again, and lay it in the sun to dry and recover.'

But that some species of ants display marked signs of what we may call sympathy even towards healthy companions in distress, is proved by the following observation of Mr. Belt. He writes:[21]

One day, watching a small column of these ants (i.e. Eciton humata), I placed a little stone on one of them to secure it. The next that approached, as soon as it discovered its situation, ran backwards in an agitated manner, and soon communicated the intelligence to the others. They rushed to the rescue; some bit at the stone and tried to move it, others seized the prisoner by the legs and tugged with such force that I thought the legs would be pulled off, but they persevered until they got the captive free. I next covered one up with a piece of clay, leaving only the ends of its antennÆ projecting. It was soon discovered by its fellows, which set to work immediately, and by biting off pieces of the clay soon liberated it. Another time I found a very few of them passing along at intervals. I confined one of these under a piece of clay at a little distance from the line, with his head projecting. Several ants passed it, but at last one discovered it and tried to pull it out, but could not. It immediately set off at a great rate, and I thought it had deserted its comrade, but it had only gone for assistance, for in a short time about a dozen ants came hurrying up, evidently fully informed of the circumstances of the case, for they made directly for their imprisoned comrade and soon set him free. I do not see how this action could be instinctive. It was sympathetic help, such as man only among the higher mammalia shows. The excitement and ardour with which they carried on their unflagging exertions for the rescue of their comrade could not have been greater if they had been human beings.

This observation seems unequivocal as proving fellow-feeling and sympathy, so far as we can trace any analogy between the emotions of the higher animals and those of insects. That insects with such highly organised social habits, and depending so greatly on the principles of co-operation, should manifest emotions or instincts of an incipiently altruistic character, is no more than we should antecedently expect on the general principle of survival of the fittest. Our only surprise should be that these emotions, or instincts, should appear to be so feebly developed in some species of ants, and, as we shall subsequently see, also of bees. But it may be worth while in this connection to point out that the valuable observation of Mr. Belt above quoted refers to the species of ant which, as we shall subsequently find, presents the most highly organised instincts of co-operation that are to be met with among ants, and therefore the greatest dependence of the welfare of the individual on that of the community. And the same remark is applicable to our native species, F. sanguinea, which the Rev. W. W. F. White has repeatedly seen rescuing buried companions very much in the manner described by Mr. Belt; and he does not appear to be acquainted with Mr. Belt's observations. He figures one case in which he saw three ants co-operating to dig out a buried comrade.[22]

Powers of Communication.

Huber, Kirby and Spence, Dugardin, Burmeister, Franklin, and other observers have all expressed themselves as more or less strongly of the opinion that members of the same community of ants, and other social Hymenoptera, are able to communicate information to one another by some system of language or signs. The facts, however, on which their opinion rests have not been stated with that degree of caution and detail which the acceptance of the conclusion requires. Thus, Kirby and Spence give only one instance of supposed communication between ants,[23] and even this one is inconclusive, as the facts described admit of being explained by supposing that the ants simply tracked one another by scent; while Huber merely deals in general statements as to 'contact of antennÆ,' without narrating any particulars of his observations. Therefore, until within the last few years there was really no sufficient evidence to sustain the general opinion that ants are able to communicate with one another; but the observations which I shall now detail must be regarded as fully substantiating that general opinion by facts as abundant and conclusive as the most critical among us can desire. I shall first narrate in his own words the more important of Sir John Lubbock's experiments in this connection:—

I took three tapes, each about 2 feet 6 inches long, and arranged them parallel to one another and about 6 inches apart. An end of each I attached to one of the nests (F. niger), and at the other end I placed a glass. In the glass at the end of one tape I placed a considerable number (300 to 600) of larvÆ. In the second I put two or three larvÆ only, in the third none at all. The object of the last was to see whether many ants would come to the glasses under such circumstances by mere accident, and I may at once say that scarcely any did so. I then took two ants, and placed one of them to the glass with many larvÆ, the other to that with two or three. Each of them took a larva and carried it to the nest, returning for another, and so on. After each journey I put another larva in the glass with only two or three larvÆ, to replace that which had been removed. Now, if several ants came under the above circumstances as a mere matter of accident, or accompanying one another by chance, or if they simply saw the larvÆ which were being brought, and consequently concluded that they might themselves find a larva in the same place, then the numbers going to the two glasses ought to be approximately equal. In each case the number of journeys made by the ants would be nearly the same; consequently, if it was a matter of scent, the two glasses would be in the same position. It would be impossible for an ant, seeing another in the act of bringing a larva, to judge for itself whether there were few or many left behind. On the other hand, if the strangers were brought, then it would be curious to see whether more were brought to the glass with many larvÆ than to that which only contained two or three. I should also mention that every stranger was imprisoned until the end of the experiment.

The results of these experiments were that during 47½ hours the ants which had access to a glass containing numerous larvÆ brought 257 friends to their assistance; while during an interval 5½ hours longer those which visited the glass with only two or three larvÆ brought only 82 friends; and, as already mentioned, no single ant came to the glass which contained no larvÆ. Now, as all the glasses were exposed to similar conditions, and as the roads to the first two must, in the first instance at all events, have been equally scented by the passage of ants over them, these results look very conclusive as proving some power of definite communication, not only that larvÆ are to be found, but even where the largest store is to be met with.

To this interesting account Sir John Lubbock adds,—

One case of apparent communication struck me very much. I had had an ant (F. niger) under observation one day, during which she was occupied in carrying off larvÆ to her nest. At night I imprisoned her in a small bottle; in the morning I let her out at 6.15, when she immediately resumed her occupation. Having to go to London, I imprisoned her again at 9 o'clock. When I returned at 4.40 I put her again to the larvÆ. She examined them carefully, and went home without taking one. At this time no other ants were out of the nest. In less than a minute she came out again with eight friends, and the little heap made straight for the heap of larvÆ. When they had gone two-thirds of the way I again imprisoned the marked ant; the others hesitated a few minutes, and then with curious quickness returned home. At 5.15 I put her again to the larvÆ. She again went home without a larva, but after only a few seconds' stay in the nest, came out with no less than thirteen friends. They all went towards the larvÆ, but when they had got about two-thirds of the way, although the marked ant had on the previous day passed over the ground about 150 times, and though she had just gone straight from the larvÆ to the nest, she seemed to have forgotten her way, and considered; and after she had wandered about for half an hour, I put her to the larvÆ. Now, in this case, the twenty-one ants must have been brought out by my marked one, for they came exactly with her, and there were no other ants out. Moreover, it would seem that they must have been told, because (which is very curious in itself) she did not in either case bring a larva, and consequently it cannot have been the mere sight of a larva which had induced them to follow her.

Further experiments proved, as we might have expected, that although an ant is able to communicate to her friends in the nest that she has found treasure somewhere outside, she is not able to describe to them its precise locality. Thus, having exposed larvÆ and placed an ant upon them as before, Sir John watched every time she came out of the nest with friends to assist her, but instead of allowing her to pilot the way, he took her up and carried her to the larvÆ, allowing her to return with a larva upon her own feet. Under these circumstances the friends, although evidently coming out with the intention of finding some treasure, were never able to find it; but wandered about in various directions for a while, and then returned to the nest. Thus, during two hours she brought out in her successive journeys altogether no less than 120 ants, of which number only 5 in their unguided wanderings happened to find the sought-for treasure. This result seems to prove, as we might have expected, that the communication is of the nature of some sign amounting to no more than a 'follow me.' Other experiments confirmed this result, and also brought out the fact that 'some species act much more in association than others—Formica fusca, for instance, much less than Lasius niger.' Thus Sir John Lubbock placed some honey before a marked specimen of the former species; but although she visited and revisited the honey during an entire day, she brought out no friends to share it; and although in her journeys to and from the nest she happened to pass and repass many other individuals, they took no notice of each other.

The obvious objection to these experiments, that an ant observing a friend bringing home food or a pupa might infer, without being told, that by accompanying the friend on the return journey she 'might participate in the good things,' has been partly met by the fact already stated, viz., that there is so very marked a difference in the result if, on experimenting on two ants, one had access to a large treasure and the other only to a small one. But to put this matter beyond question, Sir John Lubbock tried the experiment of pinning down a dead fly, so that the ant which found it was unable, with all her tugging, to move it towards the nest. At length she went back to the nest for assistance, and returned accompanied by seven friends. So great was her excitement, however, that she outran these friends, 'who seemed to have come out reluctantly, as if they had been asleep, and were only half awake;' and they failed to find the fly, slowly meandering about for twenty minutes. After again tugging for a time at the fly, the first ant returned a second time to the nest for assistance, and in less than a minute came out with eight friends. They were even less energetic than the first party, and having lost sight of their guide in the same manner as happened before, they all returned to the nest. Meanwhile several of the first party, which had all the while been meandering about, found the fly, and proceeded to dismember it, carrying the trophy to the nest, and calling out more friends in the ordinary way. This experiment was repeated several times and on different species, always with the same result. Now, as Sir John remarks, 'the two cases (i.e. those in which the ant brought out friends to her assistance even when she had no booty to show) surely indicate a distinct power of communication..... It is impossible to doubt that the friends were brought out by the first ant; and as she returned empty-handed to the nest, the others cannot have been induced to follow her by merely observing her proceedings. I conclude, therefore, that they possess the power of requesting their friends to come and help them.'

In order to ascertain whether the signs which communicating ants make to one another are made by means of sound, Sir John Lubbock placed near a nest of Lasius flavus six small upright pillars of wood about 1½ inch high, and on one of these he put a drop of honey. 'I then put three ants to the honey, and when each had sufficiently fed, I imprisoned her, and put another; thus always keeping three ants at the honey, but not allowing them to go home. If, then, they could summon their friends by sound, there ought soon to be many ants at the honey.' The result showed that the ants were not able thus to call to one another from a distance.

As additional proof of the general fact that at all events some ants have the power of communicating information to one another, it will be enough here to quote an exceedingly interesting observation of the distinguished geologist Hague. The quotations are taken from his letters written to Mr. Darwin, and published in Nature:[24]

On the mantelshelf of our sitting-room my wife has the habit of keeping fresh flowers. A vase stands at each end, and near the middle a small tumbler, usually filled with violets. Some time ago I noticed a pile of very small red ants on the wall above the left-hand vase, passing upward and downward between the mantelshelf and a small hole near the ceiling, at a point where a picture nail had been driven. The ants, when first observed, were not very numerous, but gradually increased in number, until on some days the little creatures formed an almost unbroken procession, issuing from the hole at the nail, descending the wall, climbing the vase directly below the nail, satisfying their desire for water or perfume, and then returning. The other vase and tumbler were not visited at that time.

As I was just then recovering from a long illness it happened that I was confined to the house, and spent my days in the room where the operations of these insects attracted my attention. Their presence caused me some annoyance, but I knew of no effective means of getting rid of them. For several days in succession I frequently brushed the ants in great numbers from the wall down to the floor; but as they were not killed the result was that they soon formed a colony in the wall at the base of the mantel, ascending thence to the shelf, so that before long the vase was attacked from above and below.

One day I observed a number of ants, perhaps thirty or forty, on the shelf at the foot of the vase. Thinking to kill them, I struck them lightly with the end of my finger, killing some and disabling the rest. The effect of this was immediate and unexpected. As soon as those ants which were approaching arrived near to where their fellows lay dead and suffering, they turned and fled with all possible haste. In half an hour the wall above the mantelshelf was cleared of ants.

During the space of an hour or two the colony from below continued to ascend until reaching the lower bevelled edge of the shelf, at which point the more timid individuals, although unable to see the vase, somehow became aware of trouble, and turned about without further investigation, while the more daring advanced hesitatingly just to the upper edge of the shelf, when, extending their antennÆ and stretching their necks, they seemed to peep cautiously over the edge until beholding their suffering companions, when they too turned and followed the others, expressing by their behaviour great excitement and terror. An hour or two later, the path or trail leading from the lower colony to the vase was almost entirely free from ants.

I killed one or two ants on their path, striking them with my finger, but leaving no visible trace. The effect of this was that as soon as an ant ascending towards the shelf reached the spot where one had been killed, it gave signs immediately of great disturbance, and returned directly at the highest possible speed.

A curious and invariable feature of their behaviour was that when such an ant, returning in fright, met another approaching, the two would always communicate, but each would pursue its own way, the second ant continuing its journey to the spot where the first had turned about, and then following that example.

For some days after this there were no ants visible on the wall, either above or below the shelf.

Then a few ants from the lower colony began to reappear, but instead of visiting the vase which had been the scene of the disaster, they avoided it altogether, and following the lower front edge of the shelf to the tumbler standing near the middle, made their attack upon that. I repeated the same experiment here with precisely the same result. Killing or maiming a few of the ants and leaving their bodies about the base of the tumbler, the others on approaching, and even before arriving at the upper surface of the shelf where their mutilated companions were visible, gave signs of intense emotion, some running away immediately, and others advancing to where they could survey the field and then hastening away precipitately.

Occasionally an ant would advance towards the tumbler until it found itself among the dead and dying; then it seemed to lose all self-possession, running hither and thither, making wide circuits about the scene of the trouble, stopping at times and elevating the antennÆ with a movement suggestive of wringing them in despair, and finally taking flight. After this another interval of several days passed, during which no ants appeared. Now, three months later, the lower colony has been entirely abandoned. Occasionally, however, especially when fresh and fragrant violets have been placed on the shelf, a few 'prospectors' descend from the upper nail-hole, rarely, almost never, approaching the vase from which they were first driven away, but seeking to satisfy their desire at the tumbler. To turn back these stragglers and keep them out of sight for a number of days, sometimes for a fortnight, it is sufficient to kill one or two ants on the trail which they follow descending the wall. This I have recently done as high up as I can reach, three or four feet above the mantel. The moment this spot is reached, an ant turns abruptly and makes for home, and in a little while there is not an ant visible on the wall.

In a subsequent volume of 'Nature' (viii. p. 244), Mr. Darwin publishes another letter which he received from Mr. Hague upon the same subject. It seems that Mr. Moggridge suggested to Mr. Darwin that, as he and others had observed ants to be repelled by the mere scent of a finger drawn across their path, the observation of Mr. Hague might really resolve itself into a dislike on the part of the ants to cross a line over which a finger had been drawn, and have nothing to do with intelligent terror inspired by the sight of their slaughtered companions. The following is Mr. Hague's reply to Mr. Darwin's request for further experiments to test this point:—

Acting on Mr. M——'s suggestion, I first tried making simple finger-marks on their path (the mantel is of marble), and found just the results which he describes in his note as observed by himself at Mentone, that is, no marked symptoms of fear, but a dislike to the spot, and an effort to avoid it by going around it, or by turning back and only crossing it again after an interval of time. I then killed several ants on the path, using a smooth stone or piece of ivory, instead of my finger, to crush them. In this case the ants approaching all turned back as before, and with much greater exhibition of fear than when the simple finger-mark was made. This I did repeatedly. The final result was the same as obtained last winter. They persisted in coming for a week or two, during which I continued to kill them, and then they disappeared, and we have seen none since. It would appear from this that while the taint of the hand is sufficient to turn them back, the killing of their fellows with a stone or other material produces the effect described in my first note. This was made clear to me at that time, from the behaviour of the ants the first day I killed any, for on that occasion some of them approaching the vase from below, on reaching the upper edge of the mantel, peeped over, and drew back on seeing what had happened about the vase, then turned away a little, and after a moment tried again at another and another point along the edge, with the same result in the end. Moreover, those that found themselves among the dead and dying went from one writhing ant to another in great haste and excitement, exhibiting the signs of fright which I described.

I hardly hope that any will return again, but if they do, and give me an opportunity, I shall endeavour to act further on Mr. M——'s suggestion.

With this quotation I shall conclude the present division of the chapter; for, looking to all the other observations previously mentioned, there can be no question concerning the general fact that ants have the power of communicating with one another. And under subsequent headings abundant additional evidence on this point will be found implicated with the other facts detailed.

Habits General in Sundry Species.

Swarming.—The precise facts with regard to the swarming of ants are not yet certainly established. As regards some of the facts, however, there is no doubt. The winged males and females first quit the nest in enormous numbers, and choose some fine afternoon in July or August for their wedding flight. The entrances to the nest are widened by the workers and increased in number, and there is a great commotion on the surface of the nest. The swarm takes place as a thick cloud of all the male and female insects, rising together to a considerable height. The flight continues for several hours, usually circling round some tree or tower, and it is during the flight that fertilisation is effected. After it is effected, the swarm returns to the ground, when the males perish, either from falling a prey, in their shelterless condition, to birds or spiders, or, on account of not being able to feed themselves, from starvation. 'The workers, or neuter ants, of their own colony have lost all interest in them from the moment of their return, and trouble themselves no more about them, for they well know that the males have now fulfilled their vocation.' The great majority of the fertilised females share the same fate as the males. But a small proportion find concealment in holes, which they either dig for themselves, or happen to find ready made, and there found a new colony. The first thing they do is to pull off their now useless wings, by scratching and twisting them, one after the other, with the clawed ends of their feet. They then lay their eggs, and become the queens of new colonies.

Forel says that no fertilised female ever returns to her original home; but that the workers keep back a certain number of females which are fertilised before the swarming takes place; in this case the workers pull off the wings of the fertilised females. The majority of observers, however, maintain that some of the females composing the swarm return to their native home to become mothers where they had been children. Probably both statements are correct. A writer in the 'Groniger Deekblad' for June 16, 1877, observes that, looking to the injurious effects of in-breeding, the facts as related by Forel are less probable than those related by other observers, and that, if they actually occur, the females fertilised before flight are probably kept by the ants as a sort of 'reserve corps to which the workers resort only in case of need, and if they fail to secure any returning queens.'

Nursing.—The eggs will not develop into larvÆ unless nursed. The nursing is effected by licking the surface of the eggs, which under the influence of this process increase in size, or grow. In about a fortnight, during which time the workers carry the eggs from higher to lower levels of the nest, and vice versÂ, according to the circumstances of heat, moisture, &c., the larvÆ are hatched out, and require no less careful nursing than the eggs. The workers feed them by placing mouths together and regurgitating food stored up in the crop or proventriculus into the intestinal tract of the young. The latter show their hunger by 'stretching out their little brown heads.' Great care is also taken by the workers in cleaning the larvÆ, as well as in carrying them up and down the chambers of the nest for warmth or shelter.

When fully grown the larvÆ spin cocoons, and are then pupÆ, or the 'ants' eggs' of bird-fanciers. These require no food, but still need incessant attention with reference to warmth, moisture, and cleanliness. When the time arrives for their emergence as perfect insects, the workers assist them to get out of their larval cases by biting through the walls of the latter. It is noticeable that in doing this the workers do not keep to any exact time, but free them sometimes earlier and sometimes later, in accordance with their rate of development. 'The little animal when freed from its chrysalis is still covered with a thin skin, like a little shirt, which has to be pulled off. When we see how neatly and gently this is done, and how the young creature is then washed, brushed, and fed, we are involuntarily reminded of the nursing of human babies. The empty cases, or cocoons, are carried outside the nest, and may be seen heaped together there for a long time. Some species carry them far away from the nest, or turn them into building materials for the dwelling.'[25]

Education.—The young ant does not appear to come into the world with a full instinctive knowledge of all its duties as a member of a social community. It is led about the nest, and 'trained to a knowledge of domestic duties, especially in the case of the larvÆ.' Later on the young ants are taught to distinguish between friends and foes. When an ants' nest is attacked by foreign ants, the young ones never join in the fight, but confine themselves to removing the pupÆ; and that the knowledge of hereditary enemies is not wholly instinctive in ants is proved by the following experiment, which we owe to Forel. He put young ants belonging to three different species into a glass case with pupÆ of six other species—all the species being naturally hostile to one another. The young ants did not quarrel, but worked together to tend the pupÆ. When the latter hatched out, an artificial colony was formed of a number of naturally hostile species all living together after the manner of the 'happy families' of the showmen.

Habit of keeping Aphides.—It is well known that various species of ants keep aphides, as men keep milch cows, to supply a nutritious secretion. Huber first observed this fact, and noticed that the ants collected the eggs of the aphides and treated them exactly as they treated their own, guarding and tending them with the utmost care. When these eggs hatch out the aphides are usually kept and fed by the ants, to whom they yield a sweet honey-like fluid, which they eject from the abdomen upon being stroked on this region by the antennÆ of the ants. Mr. Darwin, who has watched the latter process, observes with regard to it,—

I removed all the ants from a group of about a dozen aphides on a dock plant, and prevented their attendance during several hours. After this interval, I felt sure that the aphides would want to excrete. I watched them for some time through a lens, but not one excreted; I then tickled them with a hair in the same manner, as well as I could, as the ants do with their antennÆ; but not one excreted. Afterwards I allowed an ant to visit them, and it immediately seemed, by its eager way of running about, to be well aware what a rich flock it had discovered; it then began to play with its antennÆ on the abdomen, first of one aphis and then of another; and each, as soon as it felt the antennÆ, immediately lifted up its abdomen and excreted a limpid drop of sweet juice, which was eagerly devoured by the ant. Even quite young aphides behaved in this manner, showing that the action was instinctive, and not the result of experience.

The facts also show that the yielding of the secretion to the ants is, as it were, a voluntary act on the part of the aphides, or, perhaps more correctly, that the instinct to yield it has been developed in such a relation to the requirements of the ants, that the peculiar stimulation supplied by the antennÆ of the latter is necessary to start the act of secretion; for in the absence of this particular stimulation the aphides will never excrete until compelled to do so by the superabundance of the accumulating secretion. The question, therefore, directly arises how, on evolutionary principles, such a class of facts is to be met; for it is certainly difficult to understand the manner in which this instinct, so beneficial to the ants, can have arisen in the aphides, to which it does not appear, at first sight, to offer any advantages. Mr. Darwin meets the difficulty thus: 'Although there is no evidence that any animal performs an action for the exclusive good of another species, yet each tries to take advantage of the instincts of others;' and 'as the secretion is extremely viscid, it is no doubt a convenience to the aphides to have it removed; therefore probably they do not excrete solely for the good of the ants.'[26]

Some ants which keep aphides build covered ways, or tunnels, to the trees or shrubs where the aphides live. Forel saw a tunnel of this kind which was taken up a wall and down again on the other side, in order to secure a safe covered way from the nest to the aphides. Occasionally such covered ways, or tubes, are continued so as to enclose the stems of the plants on which the aphides live. The latter are thus imprisoned by the walls of the tube, which, however, expand where they take on this additional function of stabling the aphides, so that these insects are really confined in tolerably large chambers. The doors of these chambers are too small to allow the aphides to escape, while large enough for the ants to pass in and out. Forel saw such a prison or stable shaped like a cocoon, and about a centimÈtre long, which was hanging on the branch of a tree, and contained aphides carefully tended by the ants. Huber records similar observations.

Sir John Lubbock has made an interesting addition to our knowledge respecting this habit as practised by a certain species of ant (Lasius flavus), which departs in a very remarkable manner from the habit as practised by other species. He says: 'The ants took the greatest care of these eggs, carrying them off to the lower chambers with the utmost haste when the nest was disturbed.' But the most interesting of Sir John Lubbock's observations in this connection is new, and reveals an astonishing amount of method shown by the ants in farming their aphides. He says:—

When my eggs hatched I naturally thought that the aphides belonged to one of the species usually found on the roots of plants in the nests of Lasius flavus. To my surprise, however, the young creatures made the best of their way out of the nest, and, indeed, were sometimes brought out by the ants themselves. In vain I tried them with roots of grass, &c.; they wandered uneasily about, and eventually died. Moreover, they did not in any way resemble the subterranean species. In 1878 I again attempted to rear these young aphides; but though I hatched a great many eggs, I did not succeed. This year, however, I have been more fortunate. The eggs commenced to hatch the first week in March. Near one of my nests of Lasius flavus, in which I had placed some of the eggs in question, was a glass containing living specimens of several species of plants commonly found on or around ants' nests. To this some of the young aphides were brought by the ants. Shortly afterwards I observed on a plant of daisy, in the axils of the leaves, some small aphides, very much resembling those from my nest, though we had not actually traced them continuously. They seemed thriving, and remained stationary on the daisy. Moreover, whether they had sprung from the black eggs or not, the ants evidently valued them, for they built up a wall of earth round and over them. So things remained throughout the summer, but on October 9 I found that the aphides had laid some eggs exactly resembling those found in the ants' nests; and on examining daisy plants from outside, I found on many of them similar aphides, and more or less of the same eggs.

I confess these observations surprised me very much. The statements of Huber have not, indeed, attracted so much notice as many of the other interesting facts which he has recorded, because if aphides are kept by ants in their nests, it seems only natural that their eggs should also occur. The above case, however, is much more remarkable. Here are aphides, not living in the ants' nests, but outside, on the leaf-stalks of plants. The eggs are laid early in October on the food-plant of the insect. They are of no direct use to the ants, yet they are not left where they are laid, where they would be exposed to the severity of the weather and to innumerable dangers, but brought into their nests by the ants, and tended by them with the utmost care through the long winter months until the following March, when the young ones are brought out and again placed on the young shoots of the daisy. This seems to me a most remarkable case of prudence. Our ants may not perhaps lay up food for the winter, but they do more, for they keep during six months the eggs which will enable them to procure food during the following summer.

The following, which is taken from BÜchner's 'Geistesleben der Thiere' is perhaps a still more striking performance of the same kind as that which Sir John Lubbock observed:—

The author is debtor to Herr Nottebohm, Inspector of Buildings at Karlsruhe, who related the following on May 24, 1876, under the title, 'Ants as Founders of Aphides' Colonies:'—'Of two equally strong young weeping ashes, which I planted in my garden at Kattowitz, in Upper Silesia, one succeeded well, and in about five or six years showed full foliage, while the other regularly every year was covered, when it began to bud, with millions of aphides, which destroyed the young leaves and sprouts, and thus completely delayed the development of the tree. As I perceived that the only reason for this was the action of the aphides, I determined to destroy them utterly. So in the March of the following year I took the trouble to clean and wash every bough, sprig, and bud before the bursting of the latter, with the greatest care, by means of a syringe. The result was that the tree developed perfectly healthy and vigorous leaves and young shoots, and remained quite free from the aphides until the end of May or the beginning of June. My joy was of short duration. One fine sunny morning I saw a surprising number of ants running quickly up and down the trunk of the tree; this aroused my attention, and led me to look more closely. To my great astonishment I then saw that many troops of ants were busied in carrying single aphides up the stem to the top, and that in this way many of the lower leaves had been planted with colonies of aphides. After some weeks the evil was as great as ever. The tree stood alone on the grass plot, and offered the only situation for an aphides' colony for the countless ants there present. I had destroyed this colony; but the ants replanted it by bringing new colonists from distant branches, and setting them on the young leaves.[27]

Again—

MacCook noticed, of the mound-making ants, that of the workers returning to the nest from the tree on which the milking was going on, a far smaller number had distended abdomens than among those descending the tree itself. A closer investigation showed that at the roots of the trees, at the outlets of the subterranean galleries, a number of ants were assembled, which were fed by the returning ants after the fashion already described in feeding the larvÆ, and which were distinguished by the observer as 'pensioners.' MacCook often observed the same fact later, among, with others, the already described Pennsylvanian wood-ant. Distinguished individuals in the body-guard of the queen were fed in like fashion. MacCook is inclined to think that the reason of this proceeding is to be found in the 'division of labour' so general in the ant republic, and that the members of the community which are employed in building and working within the nest, leave to the others the care of providing food for themselves as well as for the younger and helpless members; they thus have a claim to receive from time to time a reciprocal toll of gratitude, and take it, as is shown very clearly, in a way demanded by the welfare of the community.[28]

Aphides are not the only insects which ants employ as cows, several other insects which yield sweet secretions being similarly utilised in various parts of the world. Thus, gall insects and cocci are kept in just the same way as aphides; but MacCook observed that where aphides and cocci are kept by the same ants, they are kept in separate chambers, or stalls. The same observer saw caterpillars of the genus Lycoena kept by ants for the sake of a sweet secretion which they supply.

Habit of making Slaves.—This habit, or instinct, obtains among at least three species of ant, viz., Formica rufescens, F. sanguinea, and strongylognathus. It was originally observed by P. Huber in the first-named species. Here the species enslaved is F. fusca, which is appropriately coloured black. The slave-making ants attack a nest of F. fusca in a body; there is a great fight with much slaughter, and, if victorious, the slave-makers carry off the pupÆ of the vanquished nest in order to hatch them out as slaves. Mr. Darwin gives an account of a battle which he himself observed.[29]

When the pupÆ hatch out in the nest of their captors, the young slaves begin their life of work, and seem to regard their master's home as their own; for they never attempt to escape, and they fight no less keenly than their masters in defence of the nest. F. sanguinea content themselves with fewer slaves than do F. rufescens; and the work that devolves upon the slaves differs according to the species which has enslaved them. In the nests of F. sanguinea the comparatively few captives are kept as household slaves; they never either enter or leave the nest, and so are never seen unless the nest is opened. They are then very conspicuous from the contrast which their black colour and small size present to the red colour and much larger size of F. rufescens. As the slaves are by this species kept strictly indoors, all the outdoor work of foraging, slave-capturing, &c., is performed by the masters; and when for any reason a nest has to migrate, the masters carry their slaves in their jaws. F. rufescens, on the other hand, assigns a much larger share of labour to the slaves, which, as we have already seen, are present in much larger numbers to take it. In this species the males and fertile females do no work of any kind; and the workers, or sterile females, though most energetic in capturing slaves, do no other kind of work. Therefore the whole community is absolutely dependent upon its slaves. The masters are not able to make their own nests or to feed their own larvÆ. When they migrate, it is the slaves that determine the migration, and, reversing the order of things that obtains in F. sanguinea, carry their masters in their jaws. Huber shut up thirty masters without a slave and with abundance of their favourite food, and also with their own larvÆ and pupÆ as a stimulus to work; but they could not feed even themselves, and many died of hunger. He then introduced a single slave, and she at once set to work, fed the surviving masters, attended to the larvÆ, and made some cells.

In order to confirm this observation, LespÈs placed a piece of sugar near a nest of slave-makers. It was soon found by one of the slaves, which gorged itself and returned to the nest. Other slaves then came out and did likewise. Then some of the masters came out, and, by pulling the legs of the feeding slaves, reminded them that they were neglecting their duty. The slaves then immediately began to serve their masters with the sugar. Forel also has confirmed all these observations of Huber. Indeed, in the case of F. rufescens, the structure of the animal is such as to render self-feeding physically impossible. Its long and narrow jaws, adapted to pierce the head of an enemy, do not admit of being used for feeding, unless liquid food is poured into them by the mouth of a slave. This fact shows of how ancient an origin the instinct of slave-making must be; it has altered in an important manner a structure which could not have been so altered prior to the establishment of the instinct in question.

Mr. Darwin thus sums up the differences in the offices of the slaves in the nests of F. sanguinea and F. rufescens respectively:—

The latter does not build its own nest, does not determine its own migrations, does not collect food for itself or for its fellows, 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 larvÆ, 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 larvÆ. So that the masters in this country receive much less service from their slaves than they do in Switzerland.

Mr. Darwin further observes that '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;' and records that he has observed in a community of the English species having an unusually large stock of slaves that 'a few slaves mingled with their masters leaving the nest, and marched 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.' And, according to Huber, the principal office of the slaves in Switzerland is to search for aphides.

Mr. Darwin also made the following observation:—'Desiring to ascertain whether F. sanguinea could distinguish the pupÆ of F. fusca, which they habitually make into slaves, and which are an unwarlike species, from F. flava, which they rarely capture, and never without a severe fight,' he found 'it was evident that they did at once distinguish them;' for while 'they eagerly and instantly seized the pupÆ of F. fusca, they were much terrified when they came across the pupÆ, or even the earth from the nest, of F. flava, and quickly ran away; but in about a quarter of an hour, shortly after the little yellow ants had crawled away (from their nest having been disturbed by Mr. Darwin), they took heart and carried off the pupÆ.'

Concerning the origin of this remarkable instinct, Mr. Darwin writes:—

As ants which are not slave-makers will, as I have seen, carry off 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 the 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 such modification to be of use to the species, until an ant was found as abjectly dependent on its slave as is the Formica rufescens.

Ants do not appear to be the only animals of which ants make slaves; for there seems to be at least one case in which these wonderful insects enslave insects of another species, which therefore may be said to stand to the ants in the relation of beasts of burden. The case to which I allude is one that is recorded in Perty's 'Intellectual Life of Animals' (2nd ed. p. 329), and is as follows:—

According to Audubon certain leaf-bugs are used as slaves by the ants in the Brazilian forests. When these ants want to bring home the leaves which they have bitten off the trees, they do it by means of a column of these bugs, which go in pairs, kept in order on either side by accompanying ants. They compel stragglers to re-enter the ranks, and laggards to keep up by biting them. After the work is done the bugs are shut up within the colony and scantily fed.

Wars.—On the wars of ants a great deal might be said, as the facts of interest in this connection are very numerous; but for the sake of brevity I shall confine myself to giving only a somewhat meagre account.

One great cause of war is the plundering of ants' nests by the slave-making species. Observers all agree that this plundering is effected by a united march of the whole army composing a nest of the slave-making species, directed against some particular nest of the species which they enslave. According to LespÈs and Forel, single scouts or small companies are first sent out from the nest to explore in various directions for a suitable nest to attack. These scouts afterwards serve as guides to the marauding excursion. Forel saw several of these scouts of the species F. rufescens or Amazon carefully inspecting a nest of F. fusca which they had found, investigating especially the entrances. These are purposely made difficult to find by their architects, and it not unfrequently happens that after all precautions and inspections on the part of the invaders, an expedition fails on account of not finding the city gates.

When the scouts have been successful in discovering a suitable nest to plunder, and have completed their strategical investigations of the locality to their satisfaction, they return straight to their own nest or fortress. Forel has then seen them walking about on the surface of their nest for a long time, as if in consultation, or making up their minds. Then some of them entered the nest, soon after which hosts of warriors streamed out of the entrances, and ran about tapping each other with their heads and antennÆ. They then formed into column and set out to pillage the nest of the slave ants. The following is the account which LespÈs gives of such expeditions:—

They only take place towards the end of the summer and in autumn. At this time the winged members of the slave species (F. fusca and F. cunicularia) have left the nest, and the Amazons will not take the trouble to bring back useless consumers. When the sky is clear our robbers leave their town in the afternoon at about three or four o'clock. At first no order is perceptible in their movements, but when they are all gathered together they form a regular column, which then moves forward quickly, and each day in a different direction. They march closely pressed together, and the foremost always appear to be seeking for something on the ground. They are each moment overtaken by others, so that the head of the column is continually growing. They are in fact seeking the traces of the ants which they propose to plunder, and it is scent that guides them. They snuff over the ground like hounds following the track of a wild animal, and when they have found it they plunge headlong forward, and the whole column rushes on behind. The smallest armies I saw consisted of several hundred individuals, but I have also seen some four times as large. They then form columns which may be five mÈtres long, and as much as fifty centimÈtres wide. After a march, which often lasts a full hour, the column arrives at the nest of the slave species. The F. cuniculariÆ, which are the strongest, offer keen opposition, but without much result. The Amazons soon penetrate within the nest, to come out again a moment later, while the assailed ants at the same time rush out in masses. During the whole time attention is directed solely to the larvÆ and pupÆ, which the Amazons steal while the others try to save as many as possible. They know very well that the Amazons cannot climb, so they fly with their precious burdens to the surrounding bushes or plants, whereto their enemies cannot follow them. They then pursue the retreating robbers and try to take away from them as much of their booty as possible. But the latter do not trouble themselves much about them, and hasten on home. On their return they do not follow the shortest road, but exactly the one by which they came, finding their way back by smell. Arrived at their nest, they immediately hand over their booty to the slaves, and trouble themselves no more about it. A few days afterwards the stolen pupÆ or nymphÆ emerge, without memory of their childhood, and immediately and without compulsion take part in all tasks.

According to BÜchner's account,[30]

From time to time the army makes a short halt, partly to let the rearguard close up, partly because different opinions arise as to the direction of the host, or because the place at which they are is unknown to them. Forel several times saw the army completely lose its way—an incident only once observed by Huber. Forel puts the number of warriors in such an army at from one hundred to more than two thousand. Its speed is on an average a mÈtre per minute, but varies much according to circumstances, and is naturally least when returning laden with booty. If the distance be very great, such bodily fatigue may at last be felt that the whole attack on the hostile nest is given up, and a retreat is begun; Forel once saw this happen after they had passed over a distance of two hundred and forty yards. Sometimes it seems as though, on coming within sight of the hostile nest, a kind of discouragement took possession of them, and prevented their making the attack. If the nest cannot at once be found, the whole army halts, and some divisions are sent forward to search for it, and these are gradually seen returning towards the centre. Forel also saw such an army only searching the first day, advancing zigzag, and with frequent halts, whereas on the following day it went forward to its aim swiftly and without delay, having found out the road. It seems that a single ant, even if it knows the way and the place, is not able alone to lead a large army, but that a considerable number must be employed in this duty. Mistakes as to the road occur with special ease during the return journey, because the several ants are laden with booty and cannot readily understand each other. Individual ants are then seen to wander about in every direction often for a long time, until they at last reach a spot known to them, and then advance swiftly to their goal. Many never come back at all. These mistakes easily occur when the robbers which have passed into a hostile nest do not come out again at the same holes whereby they entered, but by others at some distance—for instance, by a subterranean canal. Coming out thus in a strange neighbourhood, they do not know which way to take, and only some chance to find the right road during their aimless wanderings about, and recognise and follow it by smell. On the other hand, such mistakes scarcely ever happen to individuals in an unladen train, kept in good array. Other species of ants (F. fusca, rufa, sanguinea) know better how to manage under such circumstances than do the Amazons. The laden ones lay down their loads, first find where they are, and only take them up again after they have found their way. If the booty seized in the nest first attacked is too large to be all taken at once, the robbers return once, or oftener, so as to complete their work..... The ants, as already said, have no regular leaders nor chiefs, yet it is certain that in each expedition, alteration of road, or other change, the decision during that event comes from a small knot of individuals, which have previously come to an understanding, and carry the rest and the undecided along with them. These do not always follow immediately, but only after they have received several taps on the head from the members of the 'ring.' The procession does not advance until the leaders have convinced themselves by their own eyesight that the main part of the army is following.

One day Forel saw some Amazons on the surface of a nest of the F. fusca seeking and sounding in all directions, without being able to find the entrance. At last one of them found a very little hole, hardly as large as a pin's head, through which the robbers penetrated. But since, owing to the smallness of the hole, the invasion went on slowly, the search was continued, and an entrance was found further off, through which the Amazon army gradually disappeared. All was quiet. About five minutes later Forel saw a booty-laden column emerge from each hole. Not a single ant was without a load. The two columns united outside and retreated together.

A marauding excursion of the Amazons against the F. rufibarbis, a sub-species of the F. fusca, or small black ants, took place as follows:—The vanguard of the robber army found that it had reached the neighbourhood of the hostile nest more quickly than it had expected; for it halted suddenly and decidedly, and sent a number of messengers which brought up the main body and the rearguard with incredible speed. In less than thirty seconds the whole army had closed up, and hurled itself in a mass on the dome of the hostile nest. This was the more necessary as the rufibarbes during the short halt had discovered the approach of the enemy, and had utilised the time to cover the dome with defenders. An indescribable struggle followed, but the superior numbers of the Amazons overcame, and they penetrated into the nest, while the defenders poured by thousands out of the same holes, with their larvÆ and pupÆ in their jaws, and escaped to the nearest plants and bushes, running over the heaps of their assailants. These looked on the matter as hopeless, and began to retreat. But the rufibarbes, furious at their proceedings, pursued them, and endeavoured to get away from them the few pupÆ they had obtained, by trying to seize the Amazons' legs and to snatch away the pupÆ. The Amazon lets its jaws slip slowly along the captive pupa, as far as the head of its opponent, and pierces it, if it does not, as generally happens, draw back. But it often manages to seize the pupa at the instant at which the Amazon lets it go and flies with it. This is managed yet more easily when a comrade holds the robber by the legs, and compels it to loose its prey in order to guard itself against its assailant. Sometimes the robbers seize empty cocoons and carry them away, but they leave them on the road when they have discovered their mistake. In the above case the strength of the rufibarbes proved at last so great that the rearguard of the retreating army was seriously pressed, and was obliged to give up its booty. A number of the Amazons also were overpowered and killed, but not without the rufibarbes also losing many people. None the less did some individuals, as though desperate, rush into the thickest hosts of the enemy, penetrated again into the nest, and carried off several pupÆ by sheer audacity and skill. Most of them left their prey to go to the help of their comrades when assailed by the rufibarbes. Ten minutes after the commencement of the retreat all the Amazons had left the nest, and, being swifter than their opponents, they were only pursued for about halfway back. Their attack had failed on account of a short delay!

On another occasion observed by Forel, in which several fertile Amazons also took part and killed many enemies, the nest was thoroughly ravished, but the retreat was also in this case very much disturbed and harassed by the superior numbers of the enemy. There were many slain on both sides. That in spite of the above-mentioned unanimity different opinions among the members of an expedition sometimes hinder its conduct, the following observation seems to show:—An advancing column divided after it had gone about ten yards from the nest. Half turned back, while the other half went on, but after some time hesitated and also turned back. Arrived at home, it found those which had formerly turned back putting themselves in motion in a new direction. The newly returned followed them, and the reunited army, after various wheelings, halts, &c., at last turned home again by a long way round. The whole business looked like a promenade. But apparently different parties had different nests in view, while others were entirely against the expedition. Yet perhaps it was only a march for exercise.

Outer obstacles do not, as a rule, hinder the Amazons when they are once on the march. Forel saw them wade through some shallow water, although many were drowned in it, and then march over a dusty high road, although the wind blew half of them away. As they returned, booty-laden, neither wind, nor dust, nor water could make them lay down their prey. They only got back with great trouble, and turned back again to bring fresh booty, although many lost their lives.

The following is also quoted from BÜchner's excellent epitome of Forel's observations in this connection:—

The most terrible enemy of the Amazons is the sanguine ant (F. sanguinea), which also keeps slaves, and thereby often comes into collision with the Amazons on their marauding excursions. It is not equal to it in bodily strength or fighting capacity, but surpasses it in intelligence; according to Forel it is the most intelligent of all the species of ants. If Forel, for instance, poured out the contents of a sack filled with a nest of the slave species near an Amazon nest, the Amazons apparently generally regarded the tumbled together heap of ants, larvÆ, pupÆ, earth, building materials, &c., as the dome of a hostile nest, and took all imaginable but useless pains to find out the entrances thereinto, leaving on one side for this investigation their only object, the carrying off the pupÆ; but the sanguine ants under similar circumstances did not allow themselves to be deceived, but at once ransacked the whole heap.

On another occasion, while a procession of Amazon ants was on its way to plunder a nest of F. fusca, before it arrived Forel poured out a sack-full of sanguine ants, and made a break in the nest:—

The sanguine ants pressed in, while the fusca came out to defend themselves. At this moment the first Amazons arrived. When they saw the sanguine ants they drew back and awaited the main army, which appeared much disturbed at the news. But once united, the bold robbers rushed at their foes. The latter gathered together and beat back the first attack, but the Amazons closed up their ranks and made a second assault, which carried them on to the dome and into the midst of the enemy. These were overthrown, as well as a number of F. pratensis, which Forel at this moment poured out on the nest. The conquerors delayed for a moment on the dome after their victory, and then entered the nest to bring out a little of the valuable booty. A few Amazons which were mad with anger did not return with the main army, but went on slaughtering blindly among the conquered and the fugitives of the three species, fusca, pratensis, and sanguinea.

The ravished rufibarbes once became so desperate at their overthrow that they followed the robbers to their own nest, and the latter had some trouble in defending it. The rufibarbes let themselves be killed in hundreds, and really seemed as though they courted death. A small number of the Amazons also sank under the bites of their enemies. The nest contained slaves of the rufibarbis species, which on this emergency fought actively against their own race. There were also slaves of the species fusca, so that the nest included three different species of ants.

The same nest is often revisited many times on the same day or at different periods, until either there is no more to steal, or the plundered folk have hit upon better mode of defence. A column which was in the act of going back to such a plundered nest turned when halfway there, and halted, apparently on no other ground than because it had met the rearguard of the army, and had learned that the nest was exhausted, and that there was nothing more to be had there. The robbers then went off to a rufibarbis nest which was in the neighbourhood, and killed half the inhabitants while plundering the nest. The surviving rufibarbes returned after the robbery and brought up new progeny; but thirteen days later the Amazons again reaped a rich harvest from the same nest. The Amazon army often severs itself into two separate divisions when there is not enough for both to do at the same spot. Sometimes one division finds something and the other nothing, and they then reunite. If any obstacle be placed in their way they try to overcome it, in doing which some leave the main army, lose themselves, and only find their way home again with difficulty. Forel has tried to establish the normal frequency of expeditions, and found that a colony watched by himself for a space of thirty days sent out no less than forty-four marauding excursions. Of these about eight-and-twenty were completely, nine partially, and the remainder not at all successful. He four times saw the army divide into two. Half the expeditions were levelled against the rufibarbes, half against the fuscÆ. On an average a successful expedition would bring back to the colony a thousand pupÆ or larvÆ. On the whole, the number of future slaves stolen by a strong colony during a favourable summer may be reckoned at forty thousand!

The internecine battles which occasionally break out among the Amazons themselves are naturally the most cruel. They tear each other to pieces with incredible fury, and knots of five or six individuals which have pierced each other may be seen rolling over each other on the ground, it being impossible to distinguish between friend and foe. Civil wars among men are also known to be the most embittered and the most bloody.

The mode of attack practised by the other best known species of slave-making ant, sanguinea, is somewhat different:—

They march in small troops which, in case of need, summon reinforcements, and therefore as a rule only reach their goal slowly. Between the individual troops messengers or scouts run continually backwards and forwards. The first troop which arrives at the hostile nest does not rush at it, as do the Amazons, but contents itself with making provisional reconnaissances, wherein some of the assailants are generally made prisoners by the enemy, which have time to bethink and to collect themselves. Reinforcements are now brought up, and a regular siege of the nest begins. A sudden invasion, like that of the Amazons, is never seen. The besieging army forms a complete ring round the hostile nest, and the besiegers hold this with mandibles open and antennÆ drawn back, without going nearer. In this position they beat off all assaults of the besieged, until they feel themselves strong enough to advance to the attack. This attack scarcely ever fails, and has for its chief object the mastering of the entrances and outlets of the nest. A special troop guards each opening, and only allows such of the besieged to pass out as carry no pupÆ. This manoeuvre gives rise to a number of comical and characteristic scenes. By this means the sanguine ants in a few minutes manage to have all the defenders out of the nests and the pupÆ left behind. This is the case at least with the rufibarbes, while the rather less timid fuscÆ try, even at the last moment when it is useless, to stop up or barricade the entrances. The sanguine ants do not indeed possess the terrible weapons and the warlike impetuosity of the Amazons, but they are stronger and larger. If a fusca or a rufibarbis fights with a sanguine ant for the possession of a pupa, it is generally very soon overcome. While the main part of the army is penetrating into the nest to steal the pupÆ, some divisions pursue the fugitives, to take away from them the few pupÆ which may chance to have been saved. They drive them even out of the cricket-holes in which they have meanwhile taken refuge. In short, it is a razzia, or sweeping burglary, as complete as can be imagined. In the retreat the robbers in no wise hurry themselves, for they know that they are threatened by no danger and no loss, and the complete emptying of a large and distant nest often takes several days in accomplishing. The ants which have been so thoroughly robbed scarcely ever return to their former abode.

It must be admitted that a human army, robbing a foreign town or fortress, could not behave better or more prudently.

Huber gives the following account of a battle waged by sanguine ants:—

At ten, in a July morning, he noticed a small band of them emerge from their nest, and march rapidly towards a nest of negroes, around which it dispersed. A number of the blacks rushed out, gave battle, and succeeded in defeating their invaders, and in making several of them prisoners. Upon this, the remainder of the attacking force waited for a reinforcement. When this came up, they still declined further proceedings, and sent more aides-de-camp to their own nest. The result of these messages was a much larger reinforcement; but even yet the pirates appeared to shun the combat. At last, the negroes marched out from their nest in a phalanx of about two feet square, and a number of skirmishes began, which soon ended in a general mÊlÉe. Long before the event seemed certain, the negroes carried off their pupÆ to the most distant part of the nest; and when, after a longer encounter, they appeared to think further resistance vain, they retreated, attempting to take with them their young. In this, however, they were prevented, and the invaders obtained possession of their nest and the booty. When they had done this, they put in a garrison, and occupied the night and the succeeding day in carrying off their spoil.

BÜchner says—

Battles between ants of the same species often end with a lasting alliance, especially when the number of the workers on both sides is comparatively small. The wise little animals under such circumstances discover, much more quickly and better than men, that they can only destroy each other by fighting, while union would benefit both parties. Sometimes they drive each other out of their nests in a quite friendly way. Forel laid on a table a piece of bark with a nest of the gentle Leptothorax acervorum, and then put on it the contents of another nest of the same species. The last comers were by far the more numerous, and soon possessed themselves of the nest, driving out the inmates. But the latter did not know whither to go, and turned back again. They were then seized by their opponents one after the other, carried away as far as possible from the nest, and there put down. The oftener they came back the further were they carried away. One of the carriers arrived in this fashion at the edge of the table, and after it had by means of its feelers convinced itself that it had reached the end of the world, mercilessly let its burden drop into the fathomless abyss. It waited a moment to see if it had attained its object, and then turned back to the nest. Forel picked up the ant which had fallen on the floor, and put it down right in front of the returning ant. The latter repeated the same manoeuvre as at first, only stretching its neck further over the edge of the table. He several times reiterated his experiment, and always with the same result. Later the two colonies were shut up together in a glass case, and gradually learned to agree.

At other times, however, warlike ants show great and needless cruelty to one another:—

They slowly pull from their victim, that is rendered defenceless by wounds, exhaustion, or terror, first one feeler and then the other, then the legs one after another, until they at last kill it, or pull it in a completely mutilated and helpless condition to some out-of-the-way spot where it perishes miserably. Yet some compassionate hearts are to be found among the victors, which only pull the conquered to a distant place in order to get rid of them, and there let them go without injuring them.

The following account is also taken from BÜchner's 'Mind in Animals,' p. 87:—

The doors are often guarded by special sentries, which fulfil their important duty in various ways. Forel saw a nest of the Colobopsis truncata, the two or three very small round openings of which were watched by soldiers, arranged so that their thick cylindrical heads stopped them up, just as a cork stops up the mouth of a bottle. The same observer saw the Myrmecina Latreillei defend themselves against the invasions of the slave-making Strongylognathus, by placing a worker at each of the little openings of the nest, which quite stops up the opening either with its head or abdomen. The Camponotus species also defend their nests by stretching their heads in front of the openings, drawing back the antennÆ. Each approaching enemy thus receives a sharp blow or bite delivered with the whole weight of the body. MacCook noticed in the nests of the soon to be described Pennsylvanian mound-building ants, the employment of special sentries, which lay watching within the nest entrances, and sprang out at the first sight of danger to attack the enemy; and it was wonderful to see with what swiftness the news of such an alarm spread through the nest, and how the inhabitants came out en masse to meet the enemy. The Lasius species defend their large, strong, and very extensive nests against hostile attack or sieges with equal courage and skill, while other timid species seek to fly as speedily as possible with their larvÆ, pupÆ, and fruitful queens. There is, as Forel tells us, a regular barricade fight. Passage after passage is stopped and defended to the uttermost, so that the assailants can only advance step and step. Unless the latter are in an enormous majority, the struggle may last a very long time with these tactics. During this time, other workers are busy preparing subterranean passages backwards for eventual flight. Generally such passages are already made, and during a fight a new dome of the Lasius may be seen rising at a distance, it not being difficult for them to make this with the help of their extended subterranean passages and communications.

The F. exsecta or pressilabris fights in a peculiar way, which is due to care of their small and very tender bodies. It avoids all single combats, and always fights in closed ranks. Only when it thinks victory secure does it spring on its enemy's back. But its chief strength lies in the fact that many together always attack a foe. They nail down their opponent by seizing its legs and holding them firmly to the ground, while a comrade springs on the back of the defenceless creature and tries to bite through its neck. But if threatened the holders sometimes take flight, and so it happens that in battles between the exsectÆ and the much stronger pratenses not a few of the latter are seen running about with a small enemy clutching their shoulders, and making violent efforts to tear the neck of its foe. If the bearer is then seized with cramp, the nervous cord has been injured. On the other hand, if an exsecta is seized by the back by a pratensis it is at once lost.

The tactics of the turf ants resemble those of the exsectÆ, three or four of them seizing an opponent and pulling off his legs. In similar fashion the attack of the Lasius species is chiefly directed against the legs of its enemies, three, four, or five uniting in the effort. They understand barricade fighting particularly well in their large well-built dwellings, and if it comes to the worst fly by subterranean passages. They are feared by most ants on account of their numerical superiority. Forel one day poured the contents of ten nests of pratenses in front of a tree trunk inhabited by Lasius fuliginosus (jet ant). The siege at once began; but the jet ants called in help from the nests connected with their colony, and thick black columns were at once seen coming out from the surrounding trees. The pratenses were obliged to fly, and left behind them a mass of dead as well as their pupÆ, which last were carried off by the victors to their nests to be eaten.

Battles, however, are not confined to species of ants having warlike and slave-making habits. The agricultural ants likewise at times wage fierce wars with one another. The importance of seeds to these ants, and the consequent value which they set upon them, induce the animals, when supplies are scarce, to plunder each other's nests. Thus Moggridge says,—

By far the most savage and prolonged contests which I have witnessed were those in which the combatants belong to two different colonies of the same species..... The most singular contests are those which are waged for seeds by A. barbara, when one colony plunders the stores of an adjacent nest belonging to the same species, the weaker nest making prolonged though, for the most part, inefficient attempts to recover their property.

In the case of the other species of ant which I have watched fighting, the strife would last but a short time—a few hours or a day—but A. barbara will carry on the battle day after day and week after week. I was able to devote a good deal of time to watching the progress of a predatory war of this kind, waged by one nest of barbara against another, and which lasted for forty-six days, from January 18 to March 4!

I cannot of course declare positively that no cessation of hostilities may have taken place during the time, but I can affirm that whenever I visited the spot—and I did so on twelve days, or as nearly as possible twice a week—the scene was one of war and spoliation such as that which I shall now describe.

An active train of ants, nearly resembling an ordinary harvesting train, led from the entrance of one nest to that of another lower down the slope, and fifteen feet distant; but on closer examination it appeared that though the great mass of seed-bearers were travelling towards the upper nest, some few were going in the opposite direction and making for the lower. Besides this, at intervals, combats might be seen taking place, one ant seizing the free end of a seed carried by another, and endeavouring to wrench it away, and then frequently, as neither would let go, the stronger ant would drag seed and opponent towards its nest. At times other ants would interfere and seize one of the combatants and endeavour to drag it away, this often resulting in terrible mutilations, and especially in the loss of the abdomen, which would be torn off while the jaws of the victim retained their indomitable bull-dog grip upon the seed. Then the victor might be seen dragging away his prize, while its adversary, though now little more than a head and legs, offered a vigorous though of course ineffectual resistance. I frequently observed that the ants during these conflicts would endeavour to seize one another's antennÆ, and that if this were effected, the ant thus assaulted would instantly release his hold, whether of seed or adversary, and appear utterly discomfited. No doubt the antennÆ are their most sensitive parts, and injuries inflicted on these organs cause the greatest pain.

It was not until I had watched this scene for some days that I apprehended its true meaning, and discovered that the ants of the upper nest were robbing the granaries of the lower, while the latter tried to recover the stolen seeds both by fighting for them and by stealing seeds in their turn from the nest of their oppressors. The thieves, however, were evidently the stronger, and streams of ants laden with seeds arrived safely at the upper nest, while close observation showed that very few seeds were successfully carried on the reverse journey into the lower and plundered nest.

Thus when I fixed my attention on one of these robbed ants surreptitiously making its exit with the seed from the thieves' nest, and having overcome the opposition and dangers met with on its way, reaching, after a journey which took six minutes to accomplish, the entrance to its own home, I saw that it was violently deprived of its burden by a guard of ants stationed there apparently for the purpose, one of whom instantly started off and carried the seed all the way back again to the upper nest.

This I saw repeated several times.

After March 4 I never saw any acts of hostility between these nests, though the robbed nest was not abandoned. In another case of the same kind, however, where the struggle lasted thirty-one days, the robbed nest was at length completely abandoned, and on opening it I found all the granaries empty with one single exception, and this one was pierced by the matted roots of grasses and other plants, and must therefore have been long neglected by the ants. Strangely enough, not one of the seeds in this deserted granary showed traces of germination.

No doubt some very pressing need is the cause of these systematic raids in search of accumulations of seeds, and there can be little doubt that the requirements of distinct colonies of ants of the same species are often different even at the same season and date. Thus these warring colonies of ants were active on many days when the majority of the nests were completely closed; and I have even seen these robbers staggering along, enfeebled by the cold, and in wind and rain, when all other ants were safe below ground.

The agricultural ants of Texas do not appear to be less pugnacious than their European congeners. Thus MacCook says:—

A young community has sometimes to struggle into permanent prosperity through many perils. The following example is found in the unpublished Lincecum manuscripts. One day a new ant-city was observed to be located within ten or twelve yards of a long-established nest, a distance that the doctor thought would prove too near for peaceable possession—for the agriculturals seem to pre-empt a certain range of territory around their formicary as their own, within which no intrusion is allowed. He therefore concluded to keep these nests under close observation, and visited them frequently. Only a day or two had elapsed before he found that the inhabitants of the old city had made war upon the new. They had surrounded it in great numbers, and were entering, dragging out and killing the citizens. The young colonists, who seemed to be of less size than their adversaries, fought bravely, and, notwithstanding they were overwhelmed by superior numbers, killed and maimed many of their assailants. The parties were scattered in struggling pairs over a space ten or fifteen feet around the city gate, and the ground was strewed with many dead bodies. The new colonists aimed altogether at cutting off the legs of their larger foes, which they accomplished with much success. The old-city warriors, on the contrary, gnawed and clipped off the heads and abdomens of their enemies. Two days afterward the battlefield was revisited, and many ants were found lying dead tightly locked together by legs and mandibles, while hundreds of decapitated bodies and severed heads were strewed over the ground.

Another example, which is given in the published paper, is quite similar, and had like result. In forty-eight hours the old settlers had exterminated the new. The distance between the nests was about 20 feet. While the young colonists remained in concealment they were not disturbed, but as soon as they began to clear away their open disk war was declared.

MacCook, however, says that 'these ants are not always so jealous of territorial encroachment, or at least must have different standards of rights.' For he observed many cases of nests situated within twenty, and even ten feet of one another, without a battle ever occurring between members of the two communities. Therefore, without questioning the accuracy of Lincecum's observations—which, indeed, present no scope for inaccuracy—he adds, 'That neighbouring ants, like neighbouring nations of civilised men, will fall out and wage war Lincecum's examples show. Perhaps we should be quite as unsuccessful in case of these ants as of our human congeners, should we seek a sufficient reason for these wars, or satisfactory cause for these differences in dealing with neighbours which appear from the comparison of Lincecum's observations with mine.'

In connection with the wars of these ants, the following quotations may also be made from the same author:—

The erratic ants do not appear to be held as common enemies by the agriculturals, and they are even permitted to establish their formicaries within the limits of the open disk. Sometimes, however, the diminutive hillocks which mark the entrance to an erratic ant-nest multiply beyond the limit of the agriculturals' forbearance. But they do not declare war, nor resort to any personal violence. Nevertheless, they get rid of them, oddly enough, by a regular system of vexatious obstructions. They suddenly conclude that there is urgent demand for improving their public domain. Forthwith they sally forth in large numbers, fall eagerly to work gathering the little black balls which are thrown up by the earth-worms in great quantities everywhere in the prairie soil, which they bring and heap upon the paved disk until all the erratic ant-nests are covered! The entire pavement is thus raised an inch or so, and pains are taken to deposit more balls upon and around the domiciles of their tiny neighbours than elsewhere. The erratics struggle vigorously against this Pompeian treatment; they bore through the avalanche of balls, only to find barriers laid in their way. The obstructions at length become so serious that it is impossible to keep the galleries open. The dwarfs cease to contend against destiny, and, gathering together their household stores, quietly evacuate the premises of the inhospitable giants. It is the triumph of the policy of obstruction, a bloodless but effectual opposition.

Lastly, MacCook records the history of an interesting engagement which he witnessed between two nests of Tetramorium cÆspitum. It took place between Broad Street and Penn Square in Philadelphia, and lasted for nearly three weeks. Although all the combatants belonged to the same species, however great the confusion of the fight, friends were always distinguished from foes—apparently by contact of antennÆ.

Habit of keeping Domestic Pets.—Many species of ants display the curious habit of keeping in their nests sundry kinds of other insects, which, so far as observation extends, are of no benefit to the ants, and which therefore have been regarded by observers as mere domestic pets. These 'pets' are for the most part species which occur nowhere else except in ants' nests, and each species of 'pet' is peculiar to certain species of ants. Thus Moggridge found 'a large number of a minute shining brown beetle moving about among the seeds' in the nests of the harvesting ant of the south of Europe, 'belonging to the scarce and very restricted genus Colnocera, called by Kraatz C. attÆ, on account of its inhabiting the nests of ants belonging to the genus Atta.' He also observed inhabiting the same nests a minute cricket 'scarcely larger than a grain of wheat' (Gryllus myrmecophilus), which had been previously observed by Paolo Savi in the nests of several species of ants in Tuscany, where it lived on the best terms with its hosts, playing round the nests in warm weather, and retiring into them in stormy weather, while allowing the ants to carry it from place to place during migrations. Again, Mr. Bates observes that 'some of the most anomalous forms of coleopterous insects are those which live solely in the nests of ants.' Sir John Lubbock also, and other observers whom we need not wait to cite, mention similar facts. The Rev. Mr. White says that altogether 40 distinct species of Coleoptera, most of which he has in his own collection, are known to inhabit the nests of various species of ants, and to occur nowhere else.

As in all these cases the ants live on amicable terms with their guests, and in some cases even bestow labour upon them (as in carrying them from one nest to another during migration), it is evident that these insects are not only tolerated, but fostered by the ants. Moreover, as it seems absurd to credit ants with any mere fancy or caprice such as that of keeping pets, we can only conclude that these insects, like the aphides, are of some use to their hosts, although we are not yet in a position to surmise what this use can be.

Habits of Sleep and Cleanliness.—It is probable that all species of ants enjoy periods of true sleep alternating with those of activity; but actual observations on this subject have only been made on two or three species. The following is MacCook's account of these habits in the harvesting ant of Texas:—

The observation upon the ants now before me began at 8 o'clock; at 11 P.M. the cluster had nearly dissolved, only a few being asleep. To illustrate the soundness of this sleep I take the quill pen with which I write, and apply the feather end of it to an ant who is sleeping upon the soil. She has chosen a little oval depression in the surface, and lies with abdomen upon the raised edge, and face toward the lamp. Her legs are drawn up close to the body. She is perfectly still. I gently draw the feather tip along the body, stroking 'with the fur,' if I may so say. There is no motion. Again and again this action is repeated, the stroke gradually being made heavier, although always quite gentle. Still there is no change. The strokes are now directed upon the head, with the same result. Now the tip is applied to the neck, the point at which the head is united to the pro-thorax, with a waving motion intended to produce a sensation of tickling. The ant remains motionless. After continuing these experiments for several minutes, I arouse the sleeper by a sharp touch of the quill. She stretches out her head, then her legs, which she also shakes, steps nearer to the light, and begins to cleanse herself in the manner already described. This act invariably follows the waking of ants from sleep. The above description applies to the general habit of somnolence as observed upon the two named species of harvesting ants for nearly four months. I have often applied the quill, and even the point of a lead pencil, to the sleeping Floridians without breaking their slumber. There are some other details which have not appeared in the behaviour of the individual just put under observation.

Thus, I have several times seen the ants (Crudelis) yawning after awaking. I use this word for lack of one which more accurately expresses the behaviour. The action is very like that of the human animal; the mandibles are thrown open with the peculiar muscular strain which is familiar to all readers; the tongue also is sometimes thrust out, and the limbs stretched with the appearance, at least, of that tension which accompanies the yawn in the genus homo. During sleep the antennÆ have a gentle, quivering, apparently involuntary motion, which seemed to me, at times, to have the regularity of breathing. I also often noted an occasional regular lifting up and setting down of the fore-feet, one leg after another, with almost a rhythmic motion.

The length of time during which sleep is prolonged appears to vary according to circumstances and, perhaps, organism. The large head-soldiers of the Floridian harvesters appear to have a more sluggish nature than the smaller workers. Their sleep is longer and heavier. The former fact the watch readily determined. The latter appeared from the greater stolidity of the creatures under disturbance. While the ants of one group are taking sleep others may be busy at work, and these stalk among and over the sleepers, jostling them quite vigorously at times. Again, new members occasionally join the group, and, in their desire to get close up to the heat and light, crowd their drowsy comrades aside. I have seen ants who had been at work in the galleries drop their pellets, push thus into the cluster, and presently be apparently sound asleep. This rough treatment is invariably received with perfect good humour, as are like jostlings when the ants are awake. I have never seen the slightest display of anger or attempt to resent disturbance even under these circumstances, so peculiarly calculated to excite the utmost irritation in men. But of course some of the sleepers are aroused. They change position a little, or give themselves a brief combing, and then resume their nap, unless, indeed, they are satisfied. In watching these movements it was quite evident that the Florida soldiers were far less easily disturbed than their smaller fellows. They slept on stolidly while all the others were in agitation around them. Moreover, their very appearance, particularly when awaking out of sleep, indicated the greater sluggishness of their temperament in this respect.

The ordinary duration of sleep MacCook takes to be about three hours.

Ants, like many other insects, are in the habit of cleaning themselves, being, like them, provided by nature with combs and brushes, &c., for the purpose. But, unlike other insects, several species of ants are also in the habit of assisting each other in the performance of their toilet. The author last quoted gives the following account of this process in the genus Atta:—

We take a couple; the cleanser has begun at the face, which is licked thoroughly, even the mandibles being cared for, they being held apart for convenient manipulation. From the face the cleanser passes to the thorax, thence to the haunch, and so along the first leg, along the second and third in the same manner, around to the abdomen, and thence up the other side of the ant to the head. A third ant approaches and joins in the friendly task, but soon abandons the field to the original cleanser. The attitude of the cleansed all this while is one of intense satisfaction, quite resembling that of a family dog when one is scratching the back of his neck. The insect stretches out her limbs, and, as her friend takes them successively into hand, yields them limp and supple to her manipulation; she rolls gently over upon her side, even quite over upon her back, and with all her limbs relaxed presents a perfect picture of muscular surrender and ease. The pleasure which the creatures take in being thus 'combed' and 'sponged' is really enjoyable to the observer. I have seen an ant kneel down before another and thrust forward the head, drooping, quite under the face, and lie there motionless, thus expressing, as plainly as sign-language could, her desire to be cleansed. I at once understood the gesture, and so did the supplicated ant, for she at once went to work. If analogies in nature-studies were not so apt to be misleading, one might venture to suggest that our insect friends are thus in possession of a modified sort of Emmetonian Turkish bath.

The acrobatic skill of these ants, which has often furnished me amusement, and which I shall yet further illustrate, was fully shown one morning in these offices of ablution. The formicary was taken from the study, where the air had become chilled, and placed in an adjoining chamber upon the hearth, before an open-grate fire. The genial warmth was soon diffused throughout the nest, and aroused its occupants to unusual activity. A tuft of grass in the centre of the box was presently covered with them. They climbed to the very top of the spires, turned round and round, hanging by their paws, not unlike gymnasts performing upon a turning-bar. They hung or clung in various positions, grasping the grass blade with the third and fourth pairs of legs, which were spread out at length, cleansing their heads with the fore-legs or bending underneath to comb and lick the abdomen. Among these ants were several pairs, in one case a triplet, engaged in the cleansing operation just described. The cleanser clung to the grass, having a fore-leg on one side and a hind leg on the other side of the stem, stretched out at full length, while the cleansed hung in a like position below, and reached over and up, submitting herself to the pleasant process. As the progress of the act required a change of posture on the part of both insects, it was made with the utmost agility.

Similarly, Bates thus describes the cleansing process in another genus of ants (Ecitons):—

Here and there an ant was seen stretching forth first one leg and then another, to be brushed and washed by one or more of its comrades, who performed the task by passing the limb between the jaws and tongue, finishing by giving the antennÆ a friendly wipe.

Habits of Play and Leisure.—The life of ants is not all work, or, at least, is not so in all species; for in some species, at any rate, periods of recreation are habitually indulged in.

BÜchner('Geistesleben der Thiere,' p. 163) gives the following abstract of Huber's celebrated observations in this connection:—

It was of the pratensis that Huber wrote the observations touching its gymnastic sports which became so famous. He saw these ants on a fine day assembled on the surface of their nest, and behaving in a way that he could only explain as simulating festival sports or other games. They raised themselves on their hind legs, embraced each other with their fore-legs, seized each other by the antennÆ, feet, or mandibles, and wrestled—but all in friendliest fashion. They then let go, ran after each other, and played hide-and-seek. When one was victorious, it seized all the others in the ring, and tumbled them over like ninepins.

This account of Huber's found its way into many popular books, but in spite of its clearness won little credence from the reading public. 'I found it hard to believe Huber's observation,' writes Forel, 'in spite of its exactness, until I myself had seen the same.' A colony of the pratensis several times gave him the opportunity when he approached it carefully. The players caught each other by the feet or jaws, rolled over each other on the ground like boys playing, pulled each other inside the entrances of their nest, only to come out again, and so on. All this was done without bad temper, or any spurting of poison, and it was clear that all the rivalry was friendly. The least breath from the side of the observer was enough to put an end to the games. 'I understand,' continues Forel, 'that the affair must seem marvellous to those who have not seen it, especially when we remember that sexual attraction can here play no part.'

MacCook also gives an account of habits of play as indulged in among ants of the other Hemisphere:—

At one formicary half a dozen or more young queens were out at the same time. They would climb up a large pebble near the gate, face the wind, and assume a rampant posture. Several having ascended the stone at one time, there ensued a little playful passage-at-arms as to position. They nipped each other gently with the mandibles, and chased one another from favourite spots. They, however, never nipped the workers. These latter evidently kept a watch upon the sportive princesses, occasionally saluted them with their antennÆ in the usual way, or touched them at the abdomen, but apparently allowed them full liberty of action.

As to leisure, Bates writes:—

The life of these Ecitons is not all work, for I frequently saw them very leisurely employed in a way that looked like recreation. When this happened the place was always a sunny nook in the forest. The main column of the army and the branch columns, at these times, were in their ordinary relative positions; but instead of pressing forward eagerly and plundering right and left, they seemed to have been all smitten with a sudden fit of laziness. Some were walking sternly about, others were brushing their antennÆ with their fore-feet; but the drollest sight was their cleaning each other. [Here follows the above-quoted passage.] The actions of these ants looked like simple indulgence in idle amusement..... It is probable that these hours of relaxation and cleaning may be indispensable to the effective performance of their harder burdens; but whilst looking at them, the conclusion that the ants were engaged merely in play was irresistible.[31]

Funereal Habits.—In another connection it has already been stated that Sir John Lubbock found his ants to be very careful in disposing of the dead bodies of their comrades. This habit seems to be pretty general among many species of ants, and is no doubt due to sanitary requirements, thus becoming developed as a beneficial instinct by natural selection. The funereal habits of the agricultural ant are thus related by MacCook:[32]

There is nothing which is apt to awaken deeper interest in the life-history of ants than what may properly be called their funereal habits. All species whose manners I have closely observed are quite alike in their mode of caring for their own dead, and for the dry carcasses of aliens. The former they appear to treat with some degree of reverence, at least to the extent of giving them a sort of sepulture without feeding upon them. The latter, after having exhausted the juices of the body, they usually deposit together in some spot removed from the nest. I did not see any of the 'cemeteries' of the agricultural ant upon the field, nor, indeed, observe any of their behaviour towards the dead, but my artificial nests gave me some insight of this. In the first colony had been placed eight agriculturals of another nest, which were literally cut to pieces. Very soon after the ants were comfortably established in their new home, a number of them laid hold upon these disjecta membra, and began carrying them back and forth around the formicarium. The next day this continued, and several of their own number who had died were being treated in like manner. Back and forth, up and down, into every corner of the box the bearers wandered, the very embodiment of restlessness. For four days this conduct continued without any intermission. No sooner would a body or fragment thereof be dropped by one bearer than another would take it up and begin the restless circuit. The difficulty, I easily understood, was that there was no point to be found far enough removed from the living-rooms of the insects in which to inter these dead. Their desire to have their dead buried out of their sight was strong enough to keep them on this ceaseless round, apparently under the continuous influence of the hope that something might turn up to give them a more satisfactory burial-ground. It does not appear greatly to the credit of their wisdom that they were so long discovering that they were limited to a space beyond their power to enlarge. When, however, this fact was finally recognised they gave their habit its utmost bent, and began to deposit the carcasses in the extreme corner of the flat, as distant as possible from the galleries on the terrace above. Here a little hollow was made in the earth, quite up against the glass, wherein a number of bodies were laid. Portions of bodies were thrust into the chinks formed in the dry sod. This flat became the permanent charnel-house of the colony, and here, in corners, crevices, and holes, for the most part out of sight, but not always so, the dead were deposited. But the living never seemed quite reconciled to their presence. Occasionally, restless resurrectionists would disentomb the dead, shift them to another spot, or start them once more upon their unquiet wanderings. Even after the establishment of this cemetery, the creatures did not seem able to lay away their newly deceased comrades—for there were occasional deaths in the formicary—without first indulging in this funereal promenade.

In the formicaries established in glass jars, both of barbatus and crudelis, the same behaviour appeared. So great was the desire to get the dead outside the nest, that the bearers would climb up the smooth surface of the glass to the very top of the jar, laboriously carrying with them a dead ant. This was severe work, which was rarely undertaken except under the influence of this funereal enthusiasm. The jar was very smooth and quite high. Falls were frequent, but patiently the little 'undertaker' would follow the impulse of her instinct, and try and try again. Finally, as in the large box, the fact of a necessity seemed to dawn upon the ants, and a portion of the surface opposite from the entrance to the galleries, and close up against the glass, was used as burial-ground and sort of kitchen-midden, where all the refuse of the nest was deposited. Mrs. Treat has informed me that her artificial nests of crudelis behaved in precisely the same way.

An interesting fact in the funereal habits of Formica sanguinea was related to me by this lady. A visit was paid to a large colony of these slave-makers, which is established on the grounds adjoining her residence at Vineland, New Jersey. I noticed that a number of carcasses of one of the slave species, Formica fusca, were deposited together quite near the gates of the nest. These were probably chiefly the dry bodies of ants brought in from recent raids. It was noticed that the dead ants were all of one species, and thereupon Mrs. Treat informed me that the red slave-makers never deposited their dead with those of their black servitors, but always laid them by themselves, not in groups, but separately, and were careful to take them a considerable distance from the nest. One can hardly resist pointing here another likeness between the customs of these social hymenopters and those of human beings, certain of whom carry their distinctions of race, condition, or religious caste, even to the gates of the cemetery in which the poor body moulders into its mother dust!

It will be observed that none of these accounts furnish evidence of ants burying their dead, as Pliny asserts to have been the case with ants in the south of Europe. In the Proceedings of the LinnÆan Society, however (1861), there is a very definite account of such a practice as obtaining among the ants of Sydney; and although it is from the pen of an observer not well known, the observation seems to have been one about which there could scarcely have been a mistake. The observer was Mrs. Hutton, and this is her account. Having killed a number of 'soldier ants,' and returning half an hour afterwards to the place where the dead bodies were lying, she says:

I saw a large number of ants surrounding the dead ones. I determined to watch their proceedings closely. I followed four or five that started off from the rest towards a hillock a short distance off, in which was an ants' nest. This they entered, and in about five minutes they reappeared, followed by others. All fell into rank, walking regularly and slowly two by two, until they arrived at the spot where lay the dead bodies of the soldier ante. In a few minutes two of the ants advanced and took up the dead body of one of their comrades; then two others, and so on, until all were ready to march. First walked two ants bearing a body, then two without a burden; then two others with another dead ant, and so on, until the line was extended to about forty pairs, and the procession now moved slowly onwards, followed by an irregular body of about two hundred ants. Occasionally the two laden ants stopped, and laying down the dead ant, it was taken up by the two walking unburdened behind them, and thus, by occasionally relieving each other, they arrived at a sandy spot near the sea. The body of ants now commenced digging with their jaws a number of holes in the ground, into each of which a dead ant was laid, where they now laboured on until they had filled up the ants' graves. This did not quite finish the remarkable circumstances attending this funeral of the ants. Some six or seven of the ants had attempted to run off without performing their share of the task of digging; these were caught and brought back, when they were at once attacked by the body of ants and killed upon the spot. A single grave was quickly dug, and they were all dropped into it.

The Rev. W. Farren White also, in his papers on ants published in the 'Leisure Hour' (1880), after alluding to the above case, corroborates it by some interesting observations of his own. He says:—

Several of the little sextons I observed with dead in their mandibles, and one in the act of burying a corpse..... I should mention that the dead are not interred without considerable difficulty, in consequence of the sides of the trays being almost perpendicular. The work of the sextons continued until no dead bodies remained upon the surface of the nest, but all were interred in the extramural cemeteries. Afterwards I removed the trays, and turned the contents of the formicarium upside down, and then I placed six trays on the surface of the earth, two of which I filled with sugar for food. All six were used freely as cemeteries, being crowded with the corpses of the little people and their young, the larvÆ which had perished in the disruption of their home.

I have noticed in one of my formicaria a subterranean cemetery, where I have seen some ants burying their dead by placing earth above them. One ant was evidently much affected, and tried to exhume the bodies, but the united exertions of the yellow sextons were more than sufficient to neutralise the effort of the disconsolate mourner. The cemetery was now converted into a large vault, the chamber where the dead were placed, together with the passage which led to it, being completely covered in.

Habits Peculiar to Certain Species.

Leaf-cutting Ants of the Amazon (Œcodoma cephalotes).—The mode of working practised by these ants is thus described by Mr. Bates:—

They mount a tree in multitudes..... Each one places itself on the surface of a leaf, and cuts with its sharp scissor-like jaws a nearly semicircular incision on the upper side; it then takes the edge between its jaws, and by a sharp jerk detaches the piece. Sometimes they let the leaf drop to the ground, where a little heap accumulates, until carried off by another relay of workers; but generally each marches off with the piece it has operated on, and as all take the same road to the colony, the path they follow becomes in a short time smooth and bare, looking like the impression of a cart-wheel through the herbage.

Each ant carries its semicircular piece of leaf upright over its head, so that the home-returning train is rendered very conspicuous. Nearer observation shows that this home-returning or ladened train of workers keeps to one side of the road, while the outgoing or empty-handed train keeps to the other side; so that on every road there is a double train of ants going in opposite directions. When the leaves arrive at the nest they are received by a smaller kind of workers, whose duty it is to cut up the pieces of leaf into still smaller fragments, whereby the leaves seem to be better fitted for the purpose to which, as we shall presently see, they are put. These smaller workers never take any part in the outdoor labours; but they occasionally leave the nest, apparently for the sole purpose of obtaining air and exercise, for when they leave the nest they merely run about doing nothing, and frequently, as if in mere sport, mount some of the semicircular pieces of leaf which the carrier ants are taking to the nest, and so get a ride home.

From his continued observation of these ants, Bates concludes—and his opinion has been corroborated by that both of Belt and MÜller—that the object of all this labour is highly interesting and remarkable. The leaves when gathered do not themselves appear to be of any service to the ants as food; but when cut into small fragments and stored away in the nests, they become suited as a nidus for the growth of a minute kind of fungus on which the ants feed. We may therefore call these insects the 'gardening ants,' inasmuch as all their labour is given to the rearing of nutritious vegetables on artificially prepared soil. They are not particular as to the material which they collect and store up for soil, provided that it is a material on which the fungus will grow. Thus they are very partial to the inside white rind of oranges, and will carry off the flowers of certain shrubs while leaving the leaves untouched. But, to quote again from Bates,—

They are very particular about the ventilation of their underground chambers, and have numerous holes leading up to the surface from them. These they open out or close up, apparently to keep up a regular degree of temperature below. The great care they take that the pieces of leaves they carry into the nest should be neither too dry nor too damp, is also consistent with the idea that the object is the growth of a fungus that requires particular conditions of temperature and moisture to ensure its vigorous growth. If a sudden shower should come on, the ants do not carry the wet pieces into the burrows, but throw them down near the entrances. Should the weather clear up again, these pieces are picked up when nearly dried, and taken inside: should the rain, however, continue, they get sodden down into the ground, and are left there. On the contrary, in dry and hot weather, when the leaves would get dried up before they could be conveyed to the nest, the ants, when in exposed situations, do not go out at all during the hot hours, but bring in their leafy burdens in the cool of the day and during the night. As soon as the pieces of leaves are carried in they must be cut up by the small class of workers into little pieces. Some of the ants make mistakes, and carry in unsuitable leaves. Thus grass is always rejected by them, but I have seen some ants, perhaps young ones, carrying leaves of grass; but after a while these pieces are always brought out again and thrown away. I can imagine a young ant getting a severe ear-wigging from one of the major-domos for its stupidity.

When a nest is disturbed and the masses of ant-food spread about, the ants are in great concern to carry every morsel of it under shelter again; and sometimes, when I had dug into a nest, I found the next day all the earth thrown out filled with little pits, that the ants had dug into it to get out the covered-up food. When they migrate from one part to another, they also carry with them all the ant-food from their old habitations.

In BÜchner's 'Geistesleben der Thiere' there is published an interesting description of the habits of these ants, which was communicated to the author by Dr. Fr. Ellendorf of WiedenbrÜck, who has lived many years in Central America. Dr. Ellendorf says that—

It would be quite impossible for them to creep even through short grass with loads on their heads for miles. They therefore bite off the grass close to the ground for a breadth of about five inches, and throw it on one side. Thus a road is constructed, which is finally made quite smooth and even by the continual passing to and fro of millions upon millions night and day..... If the road is looked down upon from a height with these millions thickly pressed together, and all moving along with their green bannerets over their heads, it looks as though a giant green snake were gliding slowly along the ground; and this picture is all the more striking in that all these bannerets are swaying backwards and forwards.[33]

This observer made the experiment of interrupting the advance of a column of these ants, with the interesting result which he describes:—

I wished to see how they would manage if I put an obstacle in their way. Thick high grass stood on either side of their narrow road, so that they could not pass through it with the load on their heads. I placed a dry branch, nearly a foot in diameter, obliquely across their path, and pressed it down so tightly on the ground that they could not creep underneath. The first comers crawled beneath the branch as far as they could, and then tried to climb over, but failed owing to the weight on their heads. Meanwhile the unloaded ants from the other side came on, and when these succeeded in climbing over the bough there was such a crush that the unladen ants had to clamber over the laden, and the result was a terrible muddle. I now walked along the train, and found that all the ants with their bannerets on their heads were standing still, thickly pressed together, awaiting the word of command from the front. When I turned back to the obstacle, I saw with astonishment that the loads had been laid aside by more than a foot's length of the column, one imitating the other. And now work began on both sides of the branch, and in about half an hour a tunnel was made beneath it. Each ant then took up its burden again, and the march was resumed in the most perfect order.

A migration of these ants is thus described by the same observer:—

The road led towards a cocoa plantation, and here I soon discovered the building which I afterwards visited daily. As I again went thither one day I was met, at a considerable distance from the nest, by a closely pressed column coming thence, and all the ants laden with leaves, beetles, pupÆ, butterflies, &c.; the nearer I came to the nest, the greater was the activity. It was soon plain to me that the ants were in the act of leaving their dwelling, and I walked along the train to discover the new abode. They had gone for some distance along the old road, and had then made a new one through the grass to a cooler place, lying rather higher. The grass on the new road was all bitten off close to the ground, and thousands were busy carrying the path on to the new building. At the new home itself was an unusual stir of life. There were all sorts of labourers—architects, builders, carpenters, sappers, helpers. A number were busy digging a hole in the ground, and they carried out little pellets of earth and laid them together on end to make a wall. Others drew along little twigs, straws, and grass-stalks, and put them near the place of building. I was anxious to know why they had quitted their old home, and when the departure was complete, I dug it up with a spade. At a depth of about a foot and a half I found several tunnels of a large marmot species, the terror of cocoa planters, because in making their passages they gnaw off the thickest roots of the cocoa plants. The interior of the ant-hill had apparently fallen in through these mines. Unfortunately I was unable to follow further the progress of the new building, for I was obliged to leave the next day for San Juan del Sur. When I returned at the end of a week the building was finished, and the whole colony was again busy with the leaves of the coffee plants.

Harvesting Ants (Atta).—The ants which, so far as at present known, practise the peculiar and distinctive habits to be described under this division belong for the most part to one genus, Atta, which, however, comprises a number of species distributed in localised areas over all the four quarters of the globe. Hitherto nineteen species have been detected as having the habits in question. These consist of gathering nutritious seeds of grasses during summer, and storing them in granaries for winter consumption. We owe our present knowledge concerning these insects to Mr. Moggridge,[34] who studied them in the south of Europe, Dr. Lincecum,[35] and Mr. MacCook,[36] who studied them in Texas, and Colonel Sykes[37] and Dr. Jerdon,[38] who made some observations upon them in India. They also occur scattered over a great part of Europe and in Palestine, where they were clearly known to Solomon and other classical writers of antiquity,[39] whose claim to accurate observation, although long disputed (owing to the authority of Huber), has now been amply vindicated.

Mr. Moggridge, who was a careful and industrious observer, found the following points of interest in the habits of the European harvesters. From the nest in various directions there proceed outgoing trains, which may be from twenty to thirty or more yards in length, and each consists of a double row of ants, moving, like the leaf-cutting ants, in opposite directions. Those in the outgoing row are empty-handed, while those in the incoming row are laden. But here the burdens are grass seeds. The roads terminate in the foraging ground, or ant-fields, and the insects composing the columns there become dispersed by hundreds among the seed-yielding grasses. The following is their method of collecting seeds; I quote from Moggridge:—

It is not a little surprising to see that the ants bring in not only seeds of large size and fallen grain, but also green capsules, the torn stalks of which show that they have been freshly gathered from the plant. The manner in which they accomplish this feat is as follows. An ant ascends the stem of a fruiting plant of shepherd's-purse (Capsella bursa-pastoris), let us say, and selects a well-filled but green pod about midway up the stem, those below being ready to shed their seeds at a touch. Then, seizing it in its jaws, and fixing its hind legs firmly as a pivot, it contrives to turn round and round, and so strain the fibres of the fruit-stalk that at length they snap. It then descends to the stem, patiently backing and turning upwards again as often as the clumsy and disproportionate burden becomes wedged between the thickly set stalks, and joins the line of its companions on their way to the nest. In this manner capsules of chickweed (Stellaria media) and entire calyces, containing the nutlets of calamint, are gathered; two ants also sometimes combine their efforts, when one stations itself near the base of the peduncle and gnaws it at the point of greatest tension, while the other hauls upon and twists it. I have never seen a capsule severed from its stalk by cutting alone, and the mandibles of this ant are perhaps incompetent to perform such a task. I have occasionally seen ants engaged in cutting the capsules of certain plants, drop them, and allow their companions below to carry them away; and this corresponds with the curious account given by Ælian of the manner in which the spikelets of corn are severed and thrown down 'to the people below,' t? d?? t? ??t?.

The recognition of the principle of the division of labour which the latter observation supplies, is further proved by the following quotation from the same author. A dead grasshopper which was being carried into their nest was—

Too large to pass through the door, so they tried to dismember it. Failing in this, several ants drew the wings and legs as far back as possible, while others gnawed through the muscles where the strain was greatest. They succeeded at last in thus pulling it in.

The same thing is strikingly shown by the following quotation from LespÈs:—

If the road from the place where they are gathering their harvest to the nest is very long, they make regular depÔts for their provisions under large leaves, stones, or other suitable places, and let certain workers have the duty of carrying them from depÔt to depÔt.

BÜchner (loc. cit. p. 101) also makes the following references to the statements of previous observers:—

The subterranean workers of this remarkable genus are very clever. The Rev. H. Clark reports from Rio de Janeiro, that the Sa-ubas have made a regular tunnel under the bed of the river Parahyba, which is there as broad as the Thames at London, in order to reach a storehouse which is on the opposite bank. Bates tells us that close to the Magoary rice-mills, near Para, the ants bored through the dam of a large reservoir, and the water escaped before the mischief could be remedied. In the Para Botanical Gardens an enterprising French gardener did everything he could to drive the Sa-ubas away. He lit fires at the chief entrances of their nests, and blew sulphur vapour into their galleries by means of bellows. But how astonished was Bates when he saw the vapour come out at no less a distance than seventy yards! Such an extension have the subterranean passages of the Sa-ubas.

The recognition of the principle of the division of labour, which is shown by the above observations, is further corroborated by the following quotation from Belt:—

Between the old burrows and the new one was a steep slope. Instead of descending this with their burdens, they cast them down on the top of the slope, whence they rolled down to the bottom, where another relay of labourers picked them up and carried them to the new burrow. It was amusing to watch the ants hurrying out with bundles of food, dropping them over the slope, and rushing back immediately for more.

The same thing has been observed, as already stated, of the leaf-cutting ants—those engaged in cutting frequently throwing down the fragments of leaf which they cut to the carriers below. The prevalence of this habit among various species of ants therefore renders credible the following statements of Vincent Gredler of Botzen which are thus recorded in 'der Zool. Gart.,' xv. p. 434:—

In Herr Gredler's monastery one of the monks had been accustomed for some months to put food regularly on his window-sill for ants coming up from the garden. In consequence of Herr Gredler's communications he took it into his head to put the bait for the ants, pounded sugar, into an old inkstand, and hung this up by a string to the cross-piece of his window, and left it hanging freely. A few ants were in with the bait. These soon found their road out over the string with their grains of sugar, and so their way back to their friends. Before long a procession was arranged on the new road from the window-sill along the string to the spot where the sugar was, and so things went on for two days, nothing fresh occurring. But one day the procession stopped at the old feeding-place on the window-sill, and took the food thence, without going up to the pendent sugar-jar. Closer observation revealed that about a dozen of the rogues were in the jar above, and were busily and unwearyingly carrying the grains of sugar to the edge of the pot, and throwing them over to their comrades down below.

Many other instances of the division of labour might be given besides these, and those to be mentioned hereafter in other connections throughout the course of the present chapter; but enough has been said to show that the principle is unquestionably acted upon by sundry species of ants.

That ants are liable to make mistakes, and, when they do, that they profit by experience, is shown by the following experiment made by Moggridge; and many other instances might be given were it desirable:—

It sometimes happens that an ant has manifestly made a bad selection, and is told on its return that what it has brought home with much pains is no better than rubbish, and is hustled out of the nest, and forced to throw its burden away. In order to try whether these creatures were not fallible like other mortals, I one day took out with me a little packet of grey and white porcelain beads, and scattered these in the path of a harvesting train. They had scarcely lain a minute on the earth before one of the largest workers seized upon a bead, and with some difficulty clipped it with its mandibles and trotted back at a great pace to the nest. I waited for a little while, my attention being divided between the other ants who were vainly endeavouring to remove the beads, and the entrance down which the worker had disappeared, and then left the spot. On my return in an hour's time, I found the ants passing unconcernedly by and over the beads which lay where I had strewed them in apparently undiminished quantities; and I conclude from this that they had found out their mistake, and had wisely returned to their accustomed occupations.

When the grain is thus taken into the nest, it is stored in regular granaries, but not until it has been denuded of its 'husks' or 'chaff.' The denuding process is carried on below ground, and the chaff is brought up to the surface, where it is laid in heaps to be blown away by the wind.

It is a remarkable thing, and one not yet understood, why the seed, when thus stored in subterranean chambers just far enough below the surface to favour germination, does not germinate. Moggridge says that out of twenty-one nests and among many thousands of seeds that he examined, he only found twenty-seven cases of incipient germination. Moreover, all these cases occurred in months from November to February, while in the nests opened in October, March, April, and May, no sprouted seeds were discovered, though these are the months highly favourable to germination. He is at a loss to suggest the treatment to which the ants expose the seeds in order to prevent their sprouting. 'Apparently it is not that moisture or warmth or the influence of atmospheric air is denied to the seeds, for we find them in damp soil in genial weather, and often at but a trifling distance below the surface of the ground;' and he has proved that the vitality of the seeds is not impaired, for he succeeded in raising crops of young plants from seeds removed from the granaries.

He also says,—

By a fortunate chance I have been able to prove that the seeds will germinate in an undisturbed granary when the ants are prevented from obtaining access to it: and this goes to show not only that the structure and nature of the granary chamber is not sufficient of itself to prevent germination, but also that the presence of the ants is essential to secure the dormant condition of the seeds.

I discovered in two places portions of distinct nests of Atta structor which had been isolated owing to the destruction of the hollow wall behind which they lay, and then the granaries well filled up and literally choked with growing seeds, though the earth in which they lay completely enclosed and concealed them until by chance I laid them bare. In one case I knew that the destruction of the wall had only taken place ten days before, so that the seeds had sprouted in the interval.

My experiments also tend to confirm this, and to favour the belief that the non-germination of the seeds is due to some direct influence voluntarily exercised by the ants, and not merely to the conditions found in the nest, or to acid vapours which in certain cases are given off by the ants themselves.

These experiments consisted in confining a large number of harvesting ants with their queen and larvÆ in a glass test-tube partly filled with damp soil and various seeds, the whole being closed with a cork in the mouth of the test-tube. Under these circumstances the seeds all sprouted, showing that mere confinement in an atmosphere of exhalations from the ants did not prevent germination. Another series of experiments, undertaken at the suggestion of Mr. Darwin, on the effects of an atmosphere of formic acid, showed that although this vapour was very injurious to the seeds, it did not prevent their incipient germination. Therefore it yet remains to be ascertained why the seeds do not germinate in the granaries of the ants.

But in whatever way the ants manage to prevent germination, it is certain that they are aware of the importance in this connection of keeping the seeds as dry as possible; for Moggridge repeatedly observed that when the seeds which had been stored proved over-moist, the ants again took them out and spread them in the sun to dry, to be again brought into the nest after a sufficient exposure.

Lastly, he also repeatedly observed the most surprising and interesting fact that when, as we have seen was occasionally the case, the seeds did begin to germinate in the nests, the ants knew the most effective method of preventing the germination from proceeding; for he found that in these cases the ants gnawed off the tips of the radicles. This fact deserves to be considered as one of the most remarkable among the many remarkable facts of ant-psychology.

Passing on now to the harvesting or agricultural ants of Texas, attention was first called to the habits of this insect by Mr. Buckley in 1860,[40] and by Dr. Lincecum, who sent an account of his observations to Mr. Darwin, by whom they were communicated to the LinnÆan Society in 1861. Five years later a paper was published in the Proceedings of the Academy of Natural Sciences of Philadelphia from the MS. of Dr. Lincecum. Lastly, in 1877 Mr. MacCook went to Texas expressly to study the habits of these insects, and he has recently embodied the results of his observations in a book of three hundred pages.[41] These observations are for the most part confirmatory of those of Lincecum, and for this as well as for reasons to be deduced from the work itself, they deserve to be accepted as trustworthy, notwithstanding that in some cases they are provokingly incomplete. The following is an epitome of these observations.

The ants clear away all the herbage above their nest in the form of a perfect circle, or 'disk,' 15 or 20 feet in diameter, by carefully felling every stalk of grass or weed that may be growing thereon. As the nests are placed in thickly grown localities, the effect of these bald or shaven disks is highly conspicuous and peculiar, exactly resembling in miniature the clearings which the settlers make in the American backwoods. The disk, however, is not merely cleared of herbage, but also carefully levelled, all inequalities of the surface being reduced by building pellets of soil into the hollows to an extent sufficient to make a uniformly flat surface. The action of rain and the constant motion of multitudes of ants cause this flat surface to become hard and smooth. In the centre of the disk is the gateway of the nest. This may be either a simple hole or a hollow cone.

From the disk in various directions there radiate ant-roads or avenues, which are cleared and smoothed like the disk itself, and which course through the thick surrounding grass, branching and narrowing as they go till they eventually taper away. These roads are usually three or four in number before they begin to branch, but may be as many as seven. They are usually two to three inches wide at their origin, but in large nests may be as much as five. MacCook found no road longer than sixty feet, but Lincecum describes one of three hundred feet. Along these hard and level roads there is always passing, during the daytime of the harvesting season, a constant stream of ants—those going from the nest being empty-handed, and those returning to it being laden with seeds. Of course the incoming ants, converging from all quarters upon the road, and therefore increasing in numbers as they approach the nest, require greater space for free locomotion; while the outgoing ants, diverging as they get further from home, also require greater proportional space the less their distance from the nest: hence the gradual swelling in the width of the roads as they approach the nests.

The manner of collecting the seeds in the jungle surrounding the roads is thus described by MacCook:—

At last a satisfactory seed is found. It is simply lifted from the ground, or, as often happens, has to be pulled out of the soil into which it has been tightly pressed by the rain or by passing feet. Now follows a movement which at first I thought to be a testing of the seed, and which, indeed, may be partially that; but finally I concluded that it was the adjusting of the burden for safe and convenient carriage. The ant pulls at the seed-husk with its mandibles, turning and pinching or 'feeling' it on all sides. If this does not satisfy, and commonly it does not, the body is raised by stiffening out the legs, the abdomen is curved underneath, and the apex applied to the seed. I suppose this to be simply a mechanical action for the better adjusting of the load. Now the worker starts homeward. It has not lost itself in the mazes of the grass forest. It turns directly towards the road with an unerring judgment. There are many obstacles to overcome. Pebbles, pellets of earth, bits of wood, obtruding rootlets, or bent-down spears of grass block up or hinder the way. These were scarcely noticed when the ant was empty-handed. But they are troublesome barriers now that she is burdened with a seed quite as thick, twice as wide, and half as long as herself. It is most interesting to see the skill, strength, and rapidity with which the little harvester swings her treasure over or around, or pushes it beneath these obstacles. Now the seed has caught against the herbage as the porter dodges under a too narrow opening. She backs out and tries another passage. Now the sharp points of the husk are entangled in the grass. She jerks or pulls the burden loose, and hurries on. The road is reached, and progress is comparatively easy. Holding the grain in her mandibles well above the surface, she breaks into what I may describe with sufficient accuracy as 'a trot,' and with little further interruption reaches the disk and disappears within the gate. There are variations from this behaviour, more or less marked, according to the nature of the grounds, the seeds, and (I suppose) the individuality of the harvesters; but the mode of ingathering the crop is substantially as above. Each ant operated independently. Once only did I see anything like an effort to extend sympathy and aid. A worker minor seeming to have difficulty in testing or adjusting a large seed of buffalo-grass, was assisted (apparently) by one worker major, and then by another, after which she went on her way.

But these ants do not confine their harvesting operations to gathering fallen seeds; they will, like the ants of Europe, also cut seeds from the stalk.

In order to test the disposition of crudelis to garner the seeds from the stem, bunches of millet were obtained from the North, and stalks eighteen inches high, crowned by the boll of close-set seeds, were stuck in the mound of an active formicary. The ants mounted the stems and set to work vigorously to secure the seeds, clusters of twenty or more being engaged at once upon one head. The seeds were carried off and stored within the nest. This experiment proved pretty conclusively that in the seeding season crudelis does not wait for the seeds to drop, but harvests them from the plant.

The 'granaries' into which the seeds are brought are kept distinct from the 'nurseries' for the pupÆ. Their walls, floor, and roof are so hard and smooth, that MacCook thinks the insects must practise upon them 'some rude mason's craft.'

He traced these granaries to a depth of four feet below the surface of the ground, and believes, from the statements of a native peasant, that they, or at least the formicaries, extend to a depth of fifteen feet.

As regards the care that the ants take of the gathered grain, Lincecum describes the same habit as Moggridge and Sykes describe—viz., the sunning of wet seeds to dry. MacCook, however, neglected to make any experiments on this subject. Neither has he been able to throw any light upon the question as to why the stored seeds do not germinate, and is doubtful whether the habit of gnawing the radicle of sprouting seeds, which prevails in the European species, is likewise practised by the American. On two other points of importance MacCook's observations are also incomplete. One of these has reference to an alleged statement, which he is disposed to believe, that when some ants in a community have been killed by poison, the survivors avoid the poison: he, however, made no experiments to test this statement.

The other main point on which his observations are defective has reference to a remarkable statement made by Lincecum in the most emphatic terms. This statement is that upon the surface of their disk the ants sow the seeds of a certain plant, called ant-rice, for the purpose of subsequently reaping a harvest of the grain. There is no doubt that the ant-disks do very often support this peculiar kind of grass, and that the ants are particularly fond of its seed; but whether the plant is actually sown in these situations by the insects, or grows there on account of these situations being more open than the general surface of the ground—this question MacCook has failed to answer, or even to further. We are, therefore, still left with Dr. Lincecum's emphatic assurance that he has witnessed the fact. His account is that the seed of the ant-rice, which is a biennial plant, is sown in time for the autumnal rains to bring up. At the beginning of November a green row or ring of ant-rice, about four inches wide, is seen springing up round the circumference of the disk. In the vicinity of this circular ring the ants do not permit a single spire of any other grass or weed to remain a day, but leave the aristida, or ant-rice, untouched until it ripens, which occurs in June of the next year. After the maturing and harvesting of the seed, the dry stubble is cut away and removed from the pavement or disk, which is thus left unencumbered until the ensuing autumn, when the same species of grass again appears as before, and so on. Lincecum says he has seen the process go on year after year on the same ant-farms, and adds,—

There can be no doubt of the fact that the particular species of grain-bearing grass mentioned above is intentionally planted. In farmer-like manner the ground upon which it stands is carefully divested of all other grasses and weeds during the time it is growing. When it is ripe the grain is taken care of, the dry stubble cut away and carried off, the paved area being left unencumbered until the ensuing autumn, when the same 'ant-rice' reappears within the same circle, and receives the same agricultural attention as was bestowed upon the previous crop—and so on year after year, as I know to be the case, in all situations when the ant's settlements are protected from graminivorous animals.

In a second letter Dr. Lincecum, in reply to an inquiry from Mr. Darwin whether he supposed that the ants plant seeds for the ensuing crop, says:—

I have not the slightest doubt of it. And my conclusions have not been arrived at from hasty or careless observation, nor from seeing the ants do something that looked a little like it, and then guessing at the results. I have at all seasons watched the same ant-cities during the last twelve years, and I know that what I stated in my former letter is true. I visited the same cities yesterday, and found the crop of ant-rice growing finely, and exhibiting also the signs of high cultivation, and not a blade of any other kind of grass or weed was to be seen within twelve inches of the circular row of ant-rice.—(Journ. Linn. Soc., vol. vi. p. 30-1.)

Now, MacCook found the ant-rice growing as described, but only on some nests. Why it does not grow upon all the nests he does not understand. So far, then, as his observations go, they confirm those of Dr. Lincecum; but he does 'not believe that the ants deliberately sow a crop as Lincecum asserts;' he thinks 'that they have for some reason found it to their advantage to permit the aristida to grow upon their disks, while they clear off all other herbage;' but finally concludes 'that there is nothing unreasonable, nor beyond the probable capacity of the emmet intellect, in the supposition that the crop is actually sown. Simply, it is the Scotch verdict—"Not proven."'

The following facts with regard to 'modes of mining' are worth quoting from MacCook:—

In sinking the galleries the difficulty of carrying is not great in a moist or tough soil, which permits the ant to obtain goodly-sized pellets for portage. But when the soil is light and dry, so that it crumbles into dust as it is bitten off, the difficulty is greatly increased. It would be a very tedious task indeed to take out the diggings grain by grain. This difficulty the worker overcomes by balling the small particles against the surface of the gallery, the under side of the head, or within and against the mandibles. The fore-feet are used for this purpose, being pressed against the side face, turned under, and pushed upward with a motion similar to that of a man putting his hand upon his mouth. The abdomen is then swung underneath the body and the apex pressed against the little heap of grains of dirt massed against the under side of the mandibles, or between that and the smooth under surface of the head. Thus the dust is compressed into a ball which is of sufficient size to justify deportation.

The same operation is observed in the side-galleries, where the ants work very frequently upon their sides or backs, precisely as I have seen colliers do in Pennsylvania coal-mines.

The following is likewise worth quoting from the same author:—

Seeds are evidently not the only food of our agriculturals. When the ants at disk No. 2 had broken through the slight mud-sediment that sealed up their gate, as described above, they exhibited a peculiar behaviour. Instead of heading for the roads and pressing along them, they distributed themselves at once over the entire disk, radiating from the gate to all points in the circumference, from which they penetrated the jungle of grass beyond. In a moment a large number were returning across the roads, out of the grass, over the pavement toward the entrance. They bore in their mandibles objects which I presently found to be the males and females of white ants (Termes flavipes), which were filling the air, during and after the rain, in marriage flight. They had probably swarmed just before the shower. The agriculturals were under great excitement, and hurried forth and back at the top of their speed. The number of ants bearing termites was soon so great that the vestibule became choked, and a mass of struggling anthood was piled up around the gate. A stream of eager insects continually poured out of the door, pushing their way through the crowd that vainly but persistently endeavoured to get in with their burdens. The outcoming ants had the advantage, and succeeded in jostling through the quivering rosette of antennÆ, legs, heads, and abdomens. Occasionally a worker gained an entrance by dint of sheer physical force and perseverance. Again and again would the crowd rush from all sides upon the gate, only to be pushed back by the issuing throng. In the meanwhile quite a heap of termites, a good handful at least, had been accumulated at one side of the gate, the ants having evidently dropped them, in despair of entrance, and hurried off to garner more.

In due time the pressure upon the vestibule diminished, the laden workers entered more freely, and in the end this heap was transferred to the interior. The rapidity with which the ants were distributed to all parts of their roads, after the first opening of the gates, was truly surprising. I was greatly puzzled, at the first, to know what the cause of such a rush might be. The whole behaviour was such as to carry the conviction that they knew accurately what effect the rain would have, had calculated upon it, and were acting in accordance with previous experience. I had no doubt at the time, and have none now, that the capturing of insects beaten down by the rain is one of the well-established customs of these ants. I saw a few other insects taken in, and one milliped, but chiefly the white ants.

That very afternoon I found in a formicary which I then opened several large colonies, or parts of one colony of termites, nested within the limits of the disk and quite at home. The next day numbers of the winged white ants were found stored within the granaries of a large formicary. There is no reason to doubt that these insects were intended for food, in accordance with the quite universal habit of the FormicariÆ.

A curious habit has been noticed by most observers to occur in many species of ant, and it is one on which Mr. MacCook has a good deal to say. The habit in question consists in the ants transporting one another from place to place. The carrying ant seizes her comrade by the middle, and hurries along with it held aloft—the ant which is carried remaining quite motionless with all her legs drawn together. Huber supposed the process to be one enjoyable to both the insects concerned, and to be performed by mutual understanding and consent; but MacCook, in common with most other observers, supposes that it is merely a rough and primitive way of communicating to fellow-workers the locality where their services are required. He says:—

Keeping these facts in mind, we have a key to the solution of the press-gang operations which Lincecum observed among the agriculturals, and which have been fully described in other species. In the absence of any common head or directory, and of all executive officers, a change of location or any other concerted movement must be carried forward by the willing co-operation of individuals. At first sight, the act of seizing and carrying off workers does not appear like an appeal to free-will. It is indeed coercive, so far as the first act goes. But, in point of fact, the coercion ceases the moment the captive is set down within the precincts of the new movement. The carrier-ant has depended upon securing her consent and co-operation by thus bringing her within the circle of activity for which her service is sought. As a rule, no doubt, the deported ant at once yields to the influence around her, and drops into the current of fresh enterprise, in which she moves with as entire freedom and as independently as any other worker. But she is apparently under no restraint, and if she so please, may return to her former haunts.

Certain Ants of Africa.—Livingstone says of certain ants of Africa:—

They have established themselves on the plain where water stands so long annually as to allow the lotus and other aqueous plants to come to maturity. When all the ant-horizon is submerged a foot deep, they manage to exist by ascending to little houses built of black tenaceous loam on stalks of grass, and placed higher than the line of inundation. This must have been the result of experience, for, if they had waited till the water actually invaded their terrestrial habitations, they would not have been able to procure materials for their aËrial quarters, unless they dived down to the bottom for every mouthful of clay.[42]

The Tree Ant of India and New South Wales.—These ants are remarkable from their habit of forming nests only in trees. According to Col. Sykes' account, the shape of the nest is more or less globular, and about ten inches in diameter. It is formed entirely of cow-dung, which the insects collect from the ground beneath, and work into the form of thin scales. These are then built together in an imbricated manner, like tiles or slates upon the roof of a house, the upper or outer scale, however, being one unbroken sheet, which covers the whole nest like a skull-cap. Below this the scales are placed one upon another in a wavy or scalloped manner, so that numerous little arched entrances are left, and yet, owing to the imbricated manner in which the scales are arranged, the interior of the nest is perfectly protected from rain. This interior consists of a number of irregular cells, the walls of which are formed by the same process as the exterior.

In New South Wales there is another species of ant which also frequents trees, but builds within the stem and branches. In the report of Captain Cook's expedition its habits are thus described:—'Their habitations are the insides of the branches of a tree, which they contrive to excavate, by working out the pith almost to the extremity of the slenderest twig; the tree at the same time flourishing as if it had no such inmate.' On breaking one of the branches the ants swarm out in legions. Some of our native species also have the habit of excavating the interior of trees, though not on so extensive a scale.

Honey-making Ant (Myrmecocystus mexicanus).—This ant is found in Texas and New Mexico. Capt. W. B. Fleeson has observed its habits, and his observations have been communicated to the Californian Academy of Sciences, and also, by Mr. Henry Edwards, to Mr. Darwin. The following are the chief points of interest in Capt. Fleeson's results:—

The community appears to consist of three distinct kinds of ants, probably of two separate genera, whose offices in the general order of the nest would seem to be entirely apart from each other, and who perform the labour allotted to them without the least encroachment upon the duties of their fellows. These three kinds are—

I. Yellow workers; nurses and feeders of II.

II. Yellow honey-makers; sole function to secrete a kind of honey in their large globose abdomens, on which the other ants are supposed to feed. They never quit the nest, and are fed and tended by I.

III. Black workers, guards, and purveyors; surround the nest as guards or sentinels, in a manner presently to be described, and also forage for the food required for I. They are much larger and stronger insects than either I. or II., and are provided with very formidable mandibles.

The nest is placed in sandy soil in the neighbourhood of shrubs and flowers, is a perfect square, and occupies about four or five square feet of ground, the surface of which is kept almost unbroken. But the boundaries of the nest are rendered conspicuous by the guard of black workers (III.), which continuously parade round three of its sides in a close double line of defence, moving in opposite directions. In the accompanying diagram this sentry path is represented by the thick black lines. These always face the same points of the compass, and the direction in which the sentries march is one column from south-west to south-east, and the other column from south-east to south-west—each column, however, moving in regular order round three sides of a square. The southern side of the encampment is left unguarded; but if any enemy approaches on this or any other side, a number of the guards leave their stations, and sally forth to face the foe—raising themselves on their hind tarsi on meeting the enemy, and moving their large mandibles in defiance. Spiders, wasps, beetles, and other insects, if they venture too near the nest, are torn to pieces by the guard in a most merciless manner, and the dead body of the vanquished is speedily removed from the neighbourhood of the nest—the guard then marching back to resume their places in the line of defence, their object in destroying other insects being the defence of their encampment, and not the obtaining of food.

The object of leaving the southern side of the square encampment open is as follows. While some of the black workers are engaged on duty as guard, another and larger division are engaged on duty as purveyors. These enter and leave the quadrangle by its open or southern side along the dotted line marked a to the central point c. The incoming line is composed of individuals each bearing a burden of fragments of flowers or aromatic leaves. These are all deposited in the centre of the quadrangle c. Along the other diagonal e there is a no less incessantly moving double line of yellow workers (I.), whose office it is to convey the supplies deposited by the black workers at c to b, which is the gateway of the fortress. It is remarkable that no black ant is ever seen upon the line e, and no yellow one upon the line a; each keeps his own separate station, and follows his own particular duty with a steadfastness and apparent adherence to discipline that are most astonishing. The hole at d seems to be a ventilating shaft; it is never used as a gateway.

Diagram Fig. 7.

Section of the nest reveals, besides galleries, a small chamber about three feet below the surface, across which is spread, like a spider's web, a network of squares spun by the insects, the squares being about ¼ inch across, and the ends of the whole net being fastened to the earthen walls of the chamber. In each one of the squares, supported by the web, sits one of the honey-making ants (II.). Here these honey makers live in perpetual confinement, and receive a constant supply of flowers, pollen, &c., which is continually being brought them by (I.), and which, by a process analogous to that performed by the bee, they convert into honey.

Such is an epitome of the only account that the world has yet received of the habits and economy of this wonderful insect, whose instincts of military organization seem to be not less wonderful than those of the Ecitons, though in this case they are developed with reference to defence, and not to aggression. It is especially noteworthy that the black and yellow workers are believed to belong to 'two separate genera;' for if this is the case, it is the only one I can recall of two distinct species co-operating for a common end; for even the nearest parallel which we find supplied in other species of ants maintaining aphides, is not quite the same thing, seeing that the aphides are merely passive agents, like Class II., of the honey-making ant, and not actively co-operating members of the community, like Class I.

Ecitons.—We have next to consider the habits of the wonderful 'foraging,' or, as it might be more appropriately called, the military ant of the Amazon. These insects, which belong to several species of the same genus, have been carefully watched by Belt, Bates, and other naturalists. The following facts must therefore be accepted as fully established.

Eciton legionis moves in enormous armies, and everything that these insects do is done with the most perfect instinct of military organization. The army marches in the form of a rather broad and regular column, hundreds of yards in length. The object of the march is the capture and plunder of other insects, &c., for food, and as the well-organised host advances, its devastating legions set all other terrestrial life at defiance. From the main column there are sent out smaller lateral columns, the composing individuals of which play the part of scouts, branching off in various directions, and searching about with the utmost activity for insects, grubs, &c., over every log, under every fallen leaf, and in every nook and cranny where there is any chance of finding prey. When their errand is completed, they return into the main column. If the prey found is sufficiently small for the scouts themselves to manage, it is immediately seized, and carried back to the main column; but if the amount is too large for the scouts to deal with alone, messengers are sent back to the main column, whence there is immediately dispatched a detachment large enough to cope with the requirements. Insects which when killed are too large for single ants to carry, are torn in pieces, and the pieces conveyed back to the main army by different individuals. Many insects in trying to escape run up bushes and shrubs, where they are pursued from branch to branch and twig to twig by their remorseless enemies, until on arriving at some terminal ramification they must either submit to immediate capture by their pursuers, or drop down amid the murderous hosts beneath. As already stated, all the spoils that are taken by the scouts or by the detachments sent out in answer to their demands for assistance, are immediately taken back to the main column. When they arrive there, they are taken to the rear of that column by two smaller columns of carriers, which are constantly running, one on either side of the main column, with the supplies that are constantly pouring in from both sides. Each of these outside columns is a double line, the ants composing one of the two lines all running in the same direction as the main army, and the ants composing the other line all running in the opposite direction. The former are empty-handed carriers, which having deposited their burdens in the rear, are again advancing to the van for fresh burdens. Those composing the other line are all laden with the mangled remains of insects, pupÆ of other ants, &c. On either side of the main column there are also constantly running up and down a few individuals of smaller size and lighter colour than the other ants, which seem to play the part of officers; for they never leave their stations, and while running up and down the outsides of the column, they every now and again stop to touch antennÆ with some member of the rank and file, as if to give instructions. When the scouts discover a wasp's nest in a tree, a strong force is sent out from the main army, the nest is pulled to pieces, and all the larvÆ carried to the rear of the army, while the wasps fly around defenceless against the invading multitude. Or, if the nest of any other species of ant is found, a similarly strong force, or perhaps the whole army is deflected towards it, and with the utmost energy the innumerable insects set to work to sink shafts and dig mines till the whole nest is rifled of its contents. In these mining operations the ants work with an extraordinary display of organized co-operation; for those low down in the shafts do not lose time by carrying up the earth which they excavate, but pass on the pellets to those above; and the ants on the surface, when they receive the pellets, carry them, 'with an appearance of forethought that quite staggered' Mr. Bates, only just far enough to ensure that they shall not roll back again into the shaft, and, after depositing them, immediately hurry back for more. But there is not a rigid division of labour, although the work 'seems to be performed by intelligent co-operation amongst a host of eager little creatures;' for some of them act 'sometimes as carriers of pellets, and at another as miners, and all shortly afterwards assume the office of conveyors of the spoil.' Again, as showing the instincts of co-operation, the following may also be quoted from Bates's account:—

On the following morning no trace of ants could be found near the place where I had seen them the preceding day, nor were there signs of insects of any description in the thicket; but at the distance of eighty or one hundred yards, I came upon the same army, engaged evidently on a razzia of a similar kind to that of the previous evening; but requiring other resources of their instinct, owing to the nature of the ground. They were eagerly occupied on the face of an inclined bank of light earth in excavating mines, whence, from a depth of eight or ten inches, they were extracting the bodies of a bulky species of ant of the genus Formica. It was curious to see them crowding round the orifices of the mines, some assisting their comrades to lift out the bodies of the FormicÆ, and others tearing them in pieces, on account of their weight being too great for a single Eciton; a number of carriers seizing each a fragment, and carrying it off down the slope.

These Ecitons have no fixed nest themselves, but live, as it were, on a perpetual campaign. At night, however, they call a halt and pitch a camp. For this purpose they usually select a piece of broken ground, in the interstices of which they temporarily store their plunder. In the morning the army is again on the march, and before an hour or two has passed not a single ant is to be seen where the countless multitudes had previously covered the ground.

Another and larger species of Eciton (E. humata) hunts sometimes in dense armies, and sometimes in columns, according to the kind of prey of which they are in search. When in columns they are seeking for the nests of a certain species of ant which have their young in holes of rotten logs. These Ecitons when seeking for these nests hunt about, like those just described, in columns, which branch off in various directions. When a fallen log is reached, the column spreads over it, searching through all the holes and cracks. Mr. Belt says of them:—

The workers are of various sizes, and the smallest are here of use, for they squeeze themselves into the narrowest holes, and search out their prey in the furthest ramifications of the nests. When a nest of the Hypoclinea is attacked, the ants rush out, carrying the larvÆ and pupÆ in their jaws, but are immediately despoiled of them by the Ecitons, which are running about in every direction with great swiftness. Whenever they come across a Hypoclinea carrying a larva or pupa, they take it from it so quickly, that I could never ascertain exactly how it was done.

As soon as an Eciton gets hold of its prey, it rushes off back along the advancing column, which is composed of two sets, one hurrying forward, the other returning laden with their booty, but all and always in the greatest haste and apparent hurry. About the nest which they are harrying, all appears in confusion, Ecitons running here and there and everywhere in the greatest haste and disorder; but the result of all this apparent confusion is that scarcely a single Hypoclinea gets away with a pupa or larva. I never saw the Ecitons injure the Hypoclineas themselves, they were always contented with despoiling them of their young.

The columns of this species 'are composed almost entirely of workers of different sizes;' but, as in the species previously mentioned, 'at intervals of two or three yards there are larger and lighter coloured individuals that often stop, and sometimes run a little backward, stopping and touching some of the ants with their antennÆ,' and looking 'like officers giving orders and directing the march of the column.'

Concerning the other habits of this species, the same author writes:—

The eyes in the Ecitons are very small, in some of the species imperfect, and in others entirely absent; in this they differ greatly from the Pseudomyrma ants, which hunt singly and which have the eyes greatly developed. The imperfection of eyesight in the Ecitons is an advantage to the community, and to their particular mode of hunting. It keeps them together, and prevents individual ants from starting off alone after objects that, if their eyesight was better, they might discover at a distance; the Ecitons and most other ants follow each other by scent, and, I believe, they can communicate the presence of danger, of booty, or other intelligence, to a distance by the different intensity or qualities of the odours given off. I one day saw a column of Eciton hamata running along the foot of a nearly perpendicular tramway cutting, the side of which was about six feet high. At one point I noticed a sort of assembly of about a dozen individuals that appeared in consultation. Suddenly one ant left the conclave, and ran with great speed up the perpendicular face of the cutting without stopping. It was followed by others, which, however, did not keep straight on like the first, but ran a short way, then returned, then again followed a little further than the first time. They were evidently scenting the trail of the pioneer, and making it permanently recognisable. These ants followed the exact line taken by the first one, although it was far out of sight. Wherever it had made a slight dÉtour they did so likewise. I scraped with my knife a small portion of the clay on the trail, and the ants were completely at fault for a time which way to go. Those ascending and those descending stopped at the scraped portion, and made short circuits until they hit the scented trail again, when all their hesitation vanished, and they ran up and down it with the greatest confidence. On gaining the top of the cutting, the ants entered some brushwood suitable for hunting. In a very short space of time the information was communicated to the ants below, and a dense column rushed up to search for their prey. The Ecitons are singular amongst the ants in this respect, that they have no fixed habitations, but move on from one place to another, as they exhaust the hunting grounds around them. I think Eciton hamata does not stay more than four or five days in one place. I have sometimes come across the migratory columns; they may easily be known. Here and there one of the light-coloured officers moves backwards and forwards directing the columns. Such a column is of enormous length, and contains many thousands if not millions of individuals. I have sometimes followed them up for two or three hundred yards without getting to the end.

They make their temporary habitations in hollow trees, and sometimes underneath large fallen trunks that offer suitable hollows. A nest that I came across in the latter situation was open at one side. The ants were clustered together in a dense mass, like a great swarm of bees, hanging from the roof but reaching to the ground below. Their innumerable long legs looked like brown threads binding together the mass, which must have been at least a cubic yard in bulk, and contained hundreds of thousands of individuals, although many columns were outside, some bringing in the pupÆ of ants, others the legs and dissected bodies of various insects. I was surprised to see in this living nest tubular passages leading down to the centre of the mass, kept open just as if it had been formed of inorganic materials. Down these holes the ants who were bringing in booty passed with their prey. I thrust a long stick down to the centre of the cluster, and brought out clinging to it many ants holding larvÆ and pupÆ, which probably were kept warm by the crowding together of the ants. Besides the common dark-coloured workers and light-coloured officers, I saw here many still larger individuals with enormous jaws. These they go about holding wide open in a threatening manner.

It was this ant which, as previously stated, showed sympathy and fellow-feeling with companions in difficulties.

The habits of E. drepanophora are closely similar to those of the species already described; and, indeed, except in matters of detail, all the species of Ecitons have much the same habits. Mr. Bates records an interesting observation which he made on one of the moving columns of this species. He says: 'When I interfered with the column or abstracted an individual from it, news of the disturbance was quickly communicated to a distance of several yards to the rear, and the column at that point commenced retreating.' The main column is in this species narrower, viz., 'from four to six deep,' but extends to a great length, viz., half a mile or more. It was this species of Eciton that the same naturalist describes as enjoying periods of leisure and recreation in the 'sunny nooks of the forest.'

Next we have to consider E. prÆdator, of which the same observer writes:—

This is a small dark reddish species, very similar to the common red stinging ant of England. It differs from all other Ecitons in its habit of hunting, not in columns, but in dense phalanxes consisting of myriads of individuals, and was first met with at Ega, where it is very common. Nothing in insect movements is more striking than the rapid march of these large and compact bodies. Wherever they pass, all the rest of the animal world is thrown into a state of alarm. They stream along the ground and climb to the summits of all the lower trees, searching every leaf to its apex, and whenever they encounter a mass of decaying vegetable matter, where booty is plentiful, they concentrate, like other Ecitons, all their forces upon it, the dense phalanx of shining and quickly-moving bodies, as it spreads over the surface, looking like a flood of dark red liquid. They soon penetrate every part of the confused heap, and then, gathering together again in marching order, onward they move. All soft-bodied and inactive insects fall an easy prey to them, and, like other Ecitons, they tear their victims in pieces for facility of carriage. A phalanx of this species, when passing over a tract of smooth ground, occupies a space of from four to six square yards; on examining the ants closely they are seen to move, not all together in one straightforward direction, but in variously spreading contiguous columns, now separating a little from the general mass, now reuniting with it. The margins of the phalanx spread out at times like a cloud of skirmishers from the flanks of an army. I was never able to find the hive of this species.

Lastly, there are two species of Eciton totally blind, and their habits differ from those of the species which we have hitherto considered. Bates writes of them:—

The armies of E. vastator and E. erratica move, as far as I could learn, wholly under covered roads, the ants constructing them gradually but rapidly as they advance. The column of foragers pushes forward step by step, under the protection of these covered passages, through the thickets, and on reaching a rotting log, or other promising hunting-ground, pour into the crevices in search of booty. I have traced their arcades, occasionally, for a distance of one or two hundred yards; the grains of earth are taken from the soil over which the column is passing, and are fitted together without cement. It is this last-mentioned feature that distinguishes them from the similar covered roads made by termites, who use their glutinous saliva to cement the grains together. The blind Ecitons, working in numbers, build up simultaneously the sides of their convex arcades, and contrive, in a surprising manner, to approximate them and fit in the key-stones without letting the loose uncemented structure fall to pieces. There was a very clear division of labour between the two classes of neuters in these blind species. The large-headed class, although not possessing monstrously lengthened jaws like the worker-majors in E. hamata and E. drepanophora, are rigidly defined in structure from the small-headed class, and act as soldiers, defending the working community (like soldier termites) against all comers. Whenever I made a breach in one of their covered ways, all the ants underneath were set in commotion, but the worker-minors remained behind to repair the damage, whilst the large-heads issued forth in a most menacing manner, rearing their heads and snapping their jaws with an expression of the fiercest rage and defiance.

Annornia arcens.—This is the so-called 'driver' or 'marching' ant of West Africa, which in habits and intelligence closely resembles the military ants of the other hemisphere. I shall therefore not wait again to describe these habits in detail. Like the Ecitons, the marching ants of Africa have no fixed nest, but make temporary halts in the shade of hollow trees, overhanging rocks, &c. They march in large armies, and, like the Ecitons, always in the form of a long close column; but in this case the relative position of the carriers of spoil and larvÆ is reversed, for while these occupy the middle place the soldiers and officers march on either side. These have large heads armed with powerful jaws, and never take part in carrying; their function is to maintain order, act as scouts, and attack prey. The habits of these ants resemble most closely those of the blind Ecitons in that they very frequently, and indeed generally, build covered ways; they do so apparently in order to protect themselves from the heat of the African sun. Their line of march is therefore marked by a continuous arch or tunnel, which is always being constructed by the van of the column. The structure is made of earth moulded together by saliva, and is very quickly built. But it is only built in places where the line of march is exposed to the sunlight; at night, or in the shadow of trees or long grass, it is not made. If their camp is flooded by a tropical rainstorm, the ants congregate in a close mass, with the younger ants in the centre; they thus form a floating island.

It is remarkable that ants of different hemispheres should manifest so close a similarity with respect to all these wonderful habits. The Chasseur ants of Trinidad, and, according to Madame Merian, the ants of visitation of Cayenne, also display habits of the same kind.

General Intelligence of Various Species.

Many of the foregoing facts display an astonishing degree of intelligence as obtaining among ants; for I think that however much latitude we may be inclined to allow to 'blind instinct' in the way of imitating actions elsewhere due to conscious purpose, some at least of these foregoing facts can only be fairly reconciled with the view that the insects know what they are doing and why they are doing it. But as I am myself well aware of the difficulty that arises in all such cases of drawing the line between purposeless instinct and purposive intelligence, I have thought it desirable to reserve for this concluding division of the present chapter several isolated facts which have been observed among sundry species of ants, and which do not seem to admit of being reasonably comprised under the category of instinctive action, if by the latter we mean action pursued without knowledge of the relation between the means adopted and the ends attained.

It will be remembered that our test of instinctive as distinguished from truly intelligent action is simply whether all individuals of a species perform similar adaptive movements under the stimulus supplied by similar and habitual circumstances, or whether they manifest individual and peculiar adaptive movements to meet the exigencies of novel and peculiar circumstances. The importance of this distinction may be rendered manifest by the following illustrations.

diagram Fig. 8.

We have already seen that the ants which Sir John Lubbock observed display many and complex instincts, which together might seem to justify us in anticipating that animals which present such wonderful instincts must also present sufficient general intelligence to meet simple though novel exigencies by such simple adaptations as the unfamiliar circumstances require. Yet experiments which he made in this connection seem to show that such is not the case, but that these ants, with all their wealth of instinctive endowments, are utterly destitute of intelligent resources; they have abundance of common and detailed knowledge (supposing the adaptations to be made consciously) how to act under certain complex though familiar circumstances, but appear quite unable to originate any adaptive action to obviate even the simplest conceivable difficulty, if this is of a kind which they have not been previously accustomed to meet. Thus, on a horizontal rod B supported in a saucer of water S, and therefore inaccessible to the ants from beneath, he placed some larvÆ A. On the nest N he then placed a block of wood C D, constructed so that the portion D should touch the larvÆ at A. When the ants had made a number of journeys over C D A and back again, he raised the block C D so that there was an interval 3/10 of an inch between the end of the block D and the larvÆ at A.

The ants kept on coming, and tried hard to reach down from D to A, which was only just out of their reach..... After a while they all gave up their efforts and went away, losing their prize in spite of most earnest efforts, because it did not occur to them to drop 3/10 of an inch. At the moment when the separation was made there were fifteen ants on the larvÆ. These could, of course, have returned if one had stood still and allowed the others to get on its back. This, however, did not occur to them; nor did they think of letting themselves drop from the bottom of the paper (P) on to the nest. Two or three, indeed, fell down, I have no doubt by accident; but the remainder wandered about, until at length most of them got into the water.

In another experiment he interposed a light straw bridge on the way between the nest and the larvÆ, and when the ants had well learnt the way, he drew the bridge a short distance towards the nest, so that a small chasm was made in the road. The ants tried hard and ineffectually to reach across it, but it did not occur to them to push the straw into its original position.

The following experiment is still more illustrative of the absence of intelligence, because the adjustive action required would not demand the exercise of such high powers of imagination and abstraction as would have been required for the moving forwards of the paper drawbridge.

To test their intelligence I made the following experiments: I suspended some honey over a nest of Lasius flavus at a height of about ½ an inch, and accessible only by a paper bridge more than 10 feet long. Under the glass I then placed a small heap of earth. The ants soon swarmed over the earth on to the glass, and began feeding on the honey. I then removed a little of the earth, so that there was an interval of about 1/3 of an inch between the glass and the earth; but though the distance was so small, they would not jump down, but preferred to go round by the long bridge. They tried in vain to stretch up from the earth to the glass, which, however, was just out of their reach, though they could touch it with their antennÆ; but it did not occur to them to heap the earth up a little, though if they had moved only half a dozen particles of earth they would have secured for themselves direct access to the food. This, however, never occurred to them. At length they gave up all attempts to reach up to the glass, and went round by the paper bridge. I left the arrangement for several weeks, but they continued to go round by the long paper bridge.

Another and somewhat similar experiment consisted in placing an upright stick A, supporting at an angle another stick B, which nearly but not quite touched the ground at C. At the end of the stick B there were placed some larvÆ in a horizontal glass cell at D. Into this cell were also placed a number of ants along with the larvÆ. The drop from D to C was only ½ an inch; 'still, though the ants reached over and showed a great anxiety to take this short cut home, they none of them faced the leap, but all went round by the sticks, a distance of nearly 7 feet.' Sir John then reduced the interruption to 2/5 of an inch, so that the ants could even touch the glass cell with their antennÆ; yet all day long the ants continued to go the long way round rather than face the drop. Next, therefore, he took still longer sticks and tapes, and arranged them as before, only horizontally instead of vertically. He also placed some fine earth under the glass cell containing the larvÆ. The ants as before continued to go the long way round (16 feet), though the drop could not have hurt either themselves or the larvÆ, and though even this drop might have been obviated by heaping up the fine earth into a little mound 1/8 of an inch high, so as to touch the glass cell.

It is desirable, however, here to state that all species of ants do not show this aversion to allowing themselves to drop through short distances; for Moggridge describes the harvesting ants of Europe as seeming rather to enjoy acrobatic performances of this kind; and the same fact is recorded by Belt of the leaf-cutting ants of the Amazons. Dr. Bastian, in his work on 'Brain as an Organ of Mind,' suggests that the 'seeming lack of intelligence betrayed by our English ants, from their disinclination to take a small leap, may be due simply to their defective sight' (pp. 241-2). But even this consideration does not extenuate the stupidity of the ants which failed to heap up the fine earth to reach the glass cell which they were able to touch with their antennÆ.

That the species of ants on which Sir John Lubbock experimented were not, however, quite destitute of intelligence is proved by the result of the following experiment:—

I put some provisions in a shallow box with a glass top and a single hole in one side; I then put some specimens of Lasius niger to the food, and soon a stream of ants was at work busily carrying supplies off to the nest. When they had got to know their way thoroughly, and from thirty to forty were so occupied, I poured some fine mould in front of the hole, so as to cover it to a depth of about ½ an inch. I then took out the ants which were actually in the box. As soon as the ants had recovered from the shock of this unexpected proceeding on my part, they began to run all round and about the box, looking for some other place of entrance. Finding none, however, they began digging down into the earth just over the hole, carrying off the grains of earth one by one and depositing them without any order all round at a distance of from ½ to 6 inches, until they had excavated down to the doorway, when they again began carrying off the food as before.

This experiment was several times repeated on L. niger and on L. flavus, always with the same result.

Thus, then, we may conclude that the reasoning power of these ants, although shown by the first experiments to be almost nil, is shown by this experiment to be not quite nil; for the attempt to meet the exigencies of the case by first going round the box to seek another entrance, before taking the labour to remove the earth from the known entrance, implies a certain rudimentary degree of adaptive capacity which belongs to the category of the rational.

Another point of considerable interest, as bearing on the general intelligence of ants, is one that was brought out as the result of a laborious series of hourly observations, extending without intermission from 6.30 A.M. to 10 P.M. for a period of three months. The object of these observations was to ascertain whether the principle of the division of labour is practised by the ants. The result of these observations was to show that during the winter-time, when the ants are not active, certain individuals are told off to forage for supplies, and that when any casualty overtakes these individuals, others are told off to supply their places. Thus, in the words of Sir John Lubbock's analysis of his lengthy tables,—

The feeders at the beginning of the experiment were those known to us as Nos. 5, 6, and 7. On the 22nd of November a friend, registered as No. 8, came to the honey, and again on the 11th December; but with these two exceptions the whole of the supplies were carried in by Nos. 5 and 6, with a little help from No. 7. Thinking now it might be alleged that possibly these were merely unusually active or greedy individuals, I imprisoned No. 6 when she came out to feed on the 5th. As will be seen from the table, no other ant had been out to the honey for some days; and it could therefore hardly be accidental that on that very evening another ant (then registered as No. 9) came out for food. This ant, as will be seen from the table, then took the place of No. 6 (No. 5 being imprisoned). On the 11th January No. 9 took in all the supplies, again with a little help from No. 7. So matters continued until the 17th, when I imprisoned No. 9, and then again, i.e. on the 19th, another ant (No. 10) came out for the food, aided, on and after the 22nd, by another (No. 11). This seems to me very curious. From the 1st November to the 5th January, with two or three casual exceptions, the whole of the supplies were carried in by three ants, one of whom, however, did comparatively little. The other two are imprisoned, and then, but not till then, a fresh ant appears on the scene. She carries in the food for a week, and then she being imprisoned, two others undertake the task. On the other hand, in nest 1, when the first foragers were not imprisoned, they continued during the whole time to carry in the necessary supplies.

The facts, therefore, certainly seem to indicate that certain ants are told off as foragers, and that during winter, when but little food is required, two or three such foragers are sufficient to provide it.

Although Sir John Lubbock's ants showed such meagre resources of intelligent adjustment, other species of ants, which we have already had occasion to consider, appear to be as remarkable in this respect as they are in respect of their instinctive adjustments. Unfortunately observations on this subject are very sparse, but such as they are they hold out a strong inducement for any one who has the opportunity to experiment with the view of testing the intelligence of those species in connection with which the following observations have been made.

RÉaumur states that ants will make no attempt to enter an inhabited beehive to get at the contained honey, knowing that the bees will slaughter them if they do so. But if the hive is uninhabited, or the bees all dead, the ants will swarm into the hive as long as any honey is to be found there.

P. Huber records that a wall which had been partly erected by ants was observed by him—

As though it were intended to support the still unfinished arched roof of a large room, which was being built from the opposite side. But the workers which had begun the arch had given it too low an elevation for the wall on which it was to rest, and if it had been continued on the same lines it would have met the partition wall halfway up, and this was to be avoided. I had just made this criticism to myself, when a new arrival, after looking at the work, came to the same conclusion. For it began at once to destroy what had been done, and to heighten the wall on which it was supported, and to make a new arch with the materials of the old one under my very eyes. When the ants begin an undertaking it seems exactly as if an idea slowly ripened into execution in their minds. Thus if one of them finds two stalks lying crosswise on the nest, which make possible the formation of a room, or some little rafters which suggest the walls and the corners, it first observes the various parts accurately, and then quickly and neatly heaps little pellets of earth in the interspaces and alongside the stalks. It brings from every side materials that seem appropriate, and sometimes takes such from the uncompleted works of its companions, so much is it urged on by the idea which it has once conceived, and by the desire to execute it. It goes and comes and turns back again, until its plan is recognisable by the others.

Ebrard, in his 'Etudes de Moeurs' (p. 3), gives the following remarkable instance of the display of intelligence of F. fusca:—

The earth was damp and the workers were in full swing. It was a constant coming and going of ants, coming forth from their underground dwelling, and carrying back little pellets of earth for building. In order to concentrate my attention I fixed my gaze on the largest of the rooms which were being built, wherein several ants were busy. The work had made considerable progress; but although a projection could be plainly seen along the upper edge of the wall, there remained an interspace of about twelve or fifteen millimetres to fill in. Here would have been the place, in order to support the earth still to be brought in, to have had recourse to those pillars, buttresses, or fragments of dried leaves, which many ants are wont to use in building. But the use of this expedient is not customary with the ants I was observing (F. fusca). Our ants, however, were sufficient for the occasion. For a moment they seemed inclined to leave their work, but soon turned instead to a grass-plant growing near, the long narrow leaves of which ran close together. They chose the nearest, and weighted its distal end with damp earth, until its apex just bent down to the space to be covered. Unfortunately the bend was too close to the extremity, and it threatened to break. To prevent this misfortune, the ants gnawed at the base of the leaf until it bent along its whole length and covered the space required. But as this did not seem to be quite enough, they heaped damp earth between the base of the plant and that of the leaf, until the latter was sufficiently bent. After they had thus attained their object, they heaped on the buttressing leaf the materials required for building the arched roof.

The characteristic trait of the building of ants, says Forel, is the almost complete absence of an unchangeable model, peculiar to each species, such as is found in wasps, bees, and others. The ants know how to suit their indeed little perfect work to circumstances, and to take advantage of each situation. Besides, each works for itself and on a given plan, and is only occasionally aided by others when these understand its plan. Naturally many collisions occur, and some destroy that which others have made. This also gives the key to understanding the labyrinth of the dwelling. For the rest, it is always those workers which have discovered the most advantageous method, or which have shown the most patience, which win over to their plan the majority of their comrades and at last the whole colony, although not without many fights for supremacy. But if one succeeds in obtaining a second to follow it, and this second draws the others after it, the first is soon lost again in the crowd.

Espinas also observed ('Thierischen Gesellschaften,' German translation, 1879, p. 371) that each single ant made its own plan and followed it until a comrade, which had caught the idea, joined it, and then they worked together in the execution of the same plan.

Moggridge says of the harvesters of Europe,—

I have observed on more than one occasion that when in digging into an ants' nest I have thrown out an elater larva, the ants would cluster round it and direct it towards some small opening in the soil, which it would quickly enlarge and disappear down. At other times, however, the ants would take no notice of the elater, and it is my belief that the attentions paid to it on former occasions were purely selfish, and that they intended to avail themselves of the tunnel thus made down into the soil, with the view of reopening communications with the galleries and granaries concealed below, the approaches to which had been covered up. I have frequently watched the ants make use of these passages mined by the elater on these occasions.

And again, as showing apparently intelligent adaptation of their usual habits to altered circumstances, he gives an account of the behaviour of these ants when a great crowd of them were confined by him in a glass jar containing earth. He says:—

On the following morning the openings were ten in number, and the greatly increased heaps of excavated earth showed that they must probably have been at work all night. The amount of work done in this short time was truly surprising, for it must be remembered that, eighteen hours before, the earth presented a perfectly level surface, and the larvÆ and ants, now housed below, found themselves prisoners in a strange place, bounded by glass walls, and with no exit possible.

It seems to me that the ants displayed extraordinary intelligence in having thus at a moment's notice devised a plan by which the superabundant number of workers could be employed at one time without coming in one another's way. The soil contained in the jar was of course less than a tenth part of that comprised within the limits of an ordinary nest, while the number of workers was probably more than a third of the total number belonging to the colony. If therefore but one or two entrances had been pierced in the soil, the workers would have been for ever running against one another, and a great number could never have got below to help in the all-important task of preparing passages and chambers for the accommodation of the larvÆ. These numerous and funnel-shaped entrances admitted of the simultaneous descent and ascent of large numbers of ants, and the work progressed with proportionate rapidity. After a few days only three entrances, and eventually only one, remained open.

Concerning the harvesting ant of Texas, the following quotation may be made, under the present head, from MacCook. After remarking that these ants always select sunny places wherein to build their nests, or disks, he goes on to say that within a few paces of his tent—

A nest was made which was partly shaded by a small mesquite tree that stood just beyond the margin of the clearing. The sapling had probably grown up after the location of the community, and for some reason had been permitted to remain until too old to kill off. The shadow thrown upon the pavement was very slight; nevertheless, fifteen feet distant a new formicary was being established. The path from the ranch to the spring ran between this new hill and the old one, and ants were in communication between the two. An opening had been made in the ground, and the beginnings of a new formicary were quite apparent. This is the only instance observed of what seemed an attempt at colonising or removing, and I associated it with the presence of the small but growing shadow of the young tree.

He also gives us a still more remarkable observation, which indeed, I must candidly say, does not appear to me credible. I am, therefore, glad to add that it does not appear very distinctly from the account whether the author himself made the observation, or had it narrated to him by his guide. But here is the observation in his own words:—

While studying the habits of the cutting ant I was tempted to make a night visit to a farm some distance from camp, by the farmer's story of depredations made by these insects upon certain plants and vegetables. A long, dark tramp, a blind and vain search among the fields, compelled us at last to call out the countryman from his bed. He led us directly to one of the cutting ants' nests, which was overshadowed by a young peach tree. 'There they be, sir,' cried he triumphantly. They were agriculturals! So also were the other nests shown. The reason for this confounding of the two ants on the part of the people hereabouts, and the reason for the 'cutting' operations of our harvesters, will be explained farther on. It is only in point here to say that the farmer affirmed that the ants under the peach tree had stripped off the first tender leaves last spring, so that scarcely one had been left upon the limbs. I am convinced that the reason for this onslaught was the desire to be rid of the obnoxious shade, and open the formicary to the full light of the sun.

From this account it is not very clear whether the writer himself saw evidence of the former denudation of the tree, and if so whether there was any indication, other than the word of the farmer, that the denudation had been effected by the ants. To make this conclusion credible the best conceivable evidence would be required, and this, unfortunately, is just what we find wanting. Somewhat the same remarks may be made on the following quotation from the same writer, though in this case his view is to some extent supported by an observation of Moggridge, as well as by that of Ebrard already quoted:—

Here I observed what appeared to be a new mode of operation. The workers, in several cases, left the point at which they had begun a cutting, ascended the blade, and passed as far out toward the point as possible. The blade was thus borne downward, and as the ant swayed up and down it really seemed that she was taking advantage of the leverage thus gained, and was bringing the augmented force to bear upon the fracture. In two or three cases there appeared to be a division of labour; that is to say, while the cutter at the roots kept on with her work, another ant climbed the grass blade and applied the power at the opposite end of the lever. This position may have been quite accidental, but it certainly had the appearance of a voluntary co-operation. I was sorry not to be able to establish this last inference by a series of observations, as the facts were only observed in this one nest.

The observation of Moggridge, to which I have alluded as in some measure rendering support to the foregoing, is as follows. Speaking of European harvesters which he kept in an artificial nest for the purposes of close observation, he says:

I was also in this way able to see for myself much that I otherwise could not have seen. Thus I was able to watch the operation of removing roots which had pierced through their galleries, belonging to seedling plants growing on the surface, and which was performed by two ants, one pulling at the free end of the root, and the other gnawing at its fibres where the strain was greatest, until at length it gave way.

And again,—

Two ants sometimes combine their efforts, when one stations itself near the base of the peduncle, and gnaws it at the point of greatest tension, while the other hauls upon and twists it..... I have occasionally seen ants engaged in cutting the capsules of certain plants, drop them, and allow their companions below to carry them away.

Lastly, the statements of these three observers taken together serve to render credible the following quotation from Bingley,[43] who says that in Captain Cook's expedition in New South Wales ants were seen by Sir Joseph Banks and others—

As green as a leaf, which live upon trees and build their nests of various sizes, between that of a man's head and his fist. These nests are of a very curious structure: they are formed by bending down several of the leaves, each of which is as broad as a man's hand, and gluing the points of them together so as to form a purse. The viscous matter used for this purpose is an animal juice..... Their method of bending down leaves we had no opportunity to observe; but we saw thousands uniting all their strength to hold them in this position, while other busy multitudes were employed within, in applying this gluten, that was to prevent their returning back. To satisfy ourselves that the leaves were bent and held down by the efforts of these diminutive artificers, we disturbed them in their work; and as soon as they were driven from their station, the leaves on which they were employed sprang up with a force much greater than we could have thought them able to conquer by any combination of their strength.

This remarkable fact also seems to be corroborated by the following independent observation of Sir E. Tennent:—

The most formidable of all is the great red ant, or Dimiya. It is particularly abundant in gardens and on fruit-trees; it constructs its dwellings by gluing the leaves of such species as are suitable from their shape and pliancy into hollow balls, and these it lines with a kind of transparent paper, like that manufactured by the wasp. I have watched them at the interesting operation of forming these dwellings;—a line of ants standing on the edge of one leaf bring another into contact with it, and hold both together with their mandibles till their companions within attach them firmly by means of their adhesive paper, the assistants outside moving along as the work proceeds. If it be necessary to draw closer a leaf too distant to be laid hold of by the immediate workers, they form a chain by depending one from the other till the object is reached, when it is at length brought into contact, and made fast by cement.

I shall now pass on to the remarkable observation communicated to Kirby by Colonel Sykes, F.R.S., and which is thus narrated by Kirby in his 'History, Habits, and Instincts of Animals:'—

When resident at Poona, the dessert, consisting of fruits, cakes, and various preserves, always remained upon a small side table, in a verandah of the dining-room. To guard against inroads, the legs of the table were immersed in four basins filled with water; it was removed an inch from the wall, and, to keep off dust from open windows, was covered with a tablecloth. At first the ants did not attempt to cross the water, but as the strait was very narrow, from an inch to an inch and a half, and the sweets very tempting, they appear, at length, to have braved all risks, to have committed themselves to the deep, to have scrambled across the channel, and to have reached the objects of their desires, for hundreds were found every morning revelling in enjoyment: daily vengeance was executed upon them without lessening their numbers; at last the legs of the table were painted, just above the water, with a circle of turpentine. This at first seemed to prove an effectual barrier, and for some days the sweets were unmolested, after which they were again attacked by these resolute plunderers; but how they got at them seemed totally unaccountable, till Colonel Sykes, who often passed the table, was surprised to see an ant drop from the wall, about a foot above the table, upon the cloth that covered it; another and another succeeded. So that though the turpentine and the distance from the wall appeared effectual barriers, still the resources of the animal, when determined to carry its point, were not exhausted, and by ascending the wall to a certain height, with a slight effort against it, in falling it managed to land in safety upon the table.

Colonel Sykes was a good observer, so that this statement, standing upon his authority, ought not, perhaps, to be questioned. But in all cases of remarkable intelligence displayed by animals, we naturally and properly desire corroboration, however good the authority may be on which the statement of such cases may rest. I will, therefore, add the following instances of the ingenious and determined manner in which ants overcome obstacles, and which so far lend confirmation to the above account.

Professor Leuckart placed round the trunk of a tree, which was visited by ants as a pasture for aphides, a broad cloth soaked in tobacco-water. When the ants returning home down the trunk of the tree arrived at the soaked cloth, they turned round, went up the tree again to some of the overhanging branches, and allowed themselves to drop clear of the obnoxious barrier. On the other hand, the ants which desired to mount the tree first examined the nature of the barrier, then turned back and procured from a distance little pellets of earth, which they carried in their jaws and deposited one after another upon the tobacco-cloth till a road of earth was made across it, over which the ants passed to and fro with impunity.

This interesting, and indeed surprising observation of Leuckart's is, in turn, a corroboration of an almost identical one made more than a century ago by Cardinal Fleury, and communicated by him to RÉaumur, who published it in his 'l'Histoire des Insectes' (1734). The Cardinal smeared the trunk of a tree with birdlime in order to prevent the ants from ascending it; but the insects overcame the obstacle by making a road of earth, small stones, &c., as in the case just mentioned. In another instance the Cardinal saw a number of ants make a bridge across a vessel of water surrounding the bottom of an orange-tree tub. They did so by conveying a number of little pieces of wood, the choice of which material instead of earth or stones, as in the previous case, seems to betoken no small knowledge of practical engineering.

BÜchner, after quoting these cases, proceeds to say (loc. cit., p. 120),—

The ants behaved in yet more ingenious fashion under the following very similar circumstances. Herr G. Theuerkauf, the painter (Wasserthorstr. 49, Berlin), writes to the author, November 18, 1875: 'A maple tree standing on the ground of the manufacturer, Vollbaum, of Elbing (now of Dantzic), swarmed with aphides and ants. In order to check the mischief, the proprietor smeared about a foot width of the ground round the tree with tar. The first ants who wanted to cross naturally stuck fast. But what did the next? They turned back to the tree and carried down aphides, which they stuck down on the tar one after another until they had made a bridge over which they could cross the tarring without danger. The above-named merchant, Vollbaum, is the guarantor of this story, which I received from his own mouth on the very spot whereat it occurred.'

BÜchner also gives the following case on the authority of Karl Vogt (loc. cit., p. 128). An apiary of a friend was invaded by ants:—

To make this impossible for the future, the four legs of the beehive-stand were put into small, shallow bowls filled with water, as is often done with food in ant-infested places. The ants soon found a way out of this, or rather a way into their beloved honey, and that over an iron staple with which the stand was attached to a neighbouring wall. The staple was removed, but the ants did not allow themselves to be defeated. They climbed into some linden trees standing near, the branches of which hung over the stand, and then dropped upon it from the branches, doing just the same as their comrades do with respect to food surrounded by water, when they drop upon it from the ceiling of the room. In order to make this impossible, the boughs were cut away. But once more the ants were found in the stand, and closer investigation showed that one of the bowls was dried up, and that a crowd of ants had gathered in it. But they found themselves puzzled how to go on with their robbery, for the leg did not, by chance, rest on the bottom of the bowl, but was about half an inch from it. The ants were seen rapidly touching each other with their antennÆ, or carrying on a consultation, until at last a rather larger ant came forward and put an end to the difficulty. It rose to its full height on its hind legs, and struggled until at last it seized a rather projecting splinter of the wooden leg, and managed to take hold of it. As soon as this was done other ants ran on to it, strengthened the hold by clinging, and so made a little living bridge, over which the others could easily pass.

The same author publishes the following very remarkable observation, quoted from a letter to him by Dr. Ellendorf:—

It is a hard matter to protect any eatables from these creatures, let the custody be ever so close. The legs of cupboards and tables in or on which eatables are kept are placed in vessels of water. I myself did this, but I none the less found thousands of ants in the cupboard next morning. It was a puzzle to me how they crossed the water, but the puzzle was soon solved; for I found a straw in one of the saucers, which lay obliquely across the edge of the pan and touched the leg of the press: this they had used for a bridge. Hundreds were drowned in the water, apparently because disorder had reigned at first, those coming down with booty meeting those going up. But now there was perfect order; the descending stream used one side of the straw, the ascending the other. I now pushed the straw about an inch away from the cupboard leg; a terrible confusion arose. In a moment the leg immediately over the water was covered with hundreds of ants, feeling for the bridge in every direction with their antennÆ, running back again and coming in ever larger swarms, as though they had communicated to their comrades within the cupboard the fearful misfortune that had taken place. Meanwhile the new-comers continued to run along the straw, and not finding the leg of the cupboard the greatest perplexity arose. They hurried round the edge of the pan, and soon found out where the fault lay. With united forces they quickly pulled and pushed at the straw, until it again came into contact with the wood, and the communication was again restored.

This observation is strikingly, though unconsciously, confirmed by a recent writer in the Leisure Hour (1880, pp. 718-19), who having been much troubled by small red ants in the tropics swarming over his provisions, placed the latter in a meat-safe detached from the wall and standing on four legs, each of which was placed in a little tin vessel containing water. Eight or ten days afterwards he found his provisions in the safe swarming with ants as before, and on investigating their mode of access to them found—

Proceeding along the whitewashed wall a string of ants going and coming from the outer door to a height of four feet on my wall, and corresponding with that of the safe; and looking between it and the wall, I discovered the secret—the bridge which these persevering little insects had made. It consisted of a broken bit of straw, which rested with one end on a mud buttress fixed to the wall, and the other on the overhanging or projecting top of the safe, which came within an inch and a half of the wall. So they must have carried the straw up from the floor, and resting their end of it on the support they had prepared, let it fall until its other end reached the safe, and then crossed and completed the structure, for it was fastened at both ends with the mortar composed of their saliva and fine earth. Ruthlessly I destroyed the bridge, and moving the safe farther from the wall, managed to prevent their inroads for that season at least. Since then I have frequently seen short bridges, composed entirely of the concrete or mortar which the white ants use to cover up their workings, extending from a damp earthen wall to anything not more than three-quarters of an inch from it.

Of the Ecitons Mr. Belt says:—

I shall relate two more instances of the use of a reasoning faculty in these ants. I once saw a wide column trying to pass along a crumbling, nearly perpendicular slope. They would have got very slowly over it, and many of them would have fallen, but a number having secured their hold, and reaching to each other, remained stationary, and over them the main column passed. Another time they were crossing a watercourse along a small branch, not thicker than a goose-quill. They widened this natural bridge to three times its width by a number of ants clinging to it and to each other on each side, over which the column passed three or four deep; whereas excepting for this expedient they would have had to pass over in single file, and treble the time would have been consumed. Can it be contended that such insects are not able to determine by reasoning powers which is the best way of doing a thing?

Another observer, writing from the same part of the world to BÜchner, gives a still more wonderful account of the ingenuity of Ecitons in crossing water. This observer is Herr H. Kreplin, of HeidemÜhl (Station Ducherom), 'who lived for nearly twenty years in South America as an engineer, and had often the opportunity of seeing the driver ants in the forests there.' He writes to BÜchner, under date May 10, 1876, as follows:—

On both sides of the train, at about 10 mm. distance from each other, stronger ants are to be seen, distinguishable from the others by their foxy colour and very thick heads with gigantic mandibles. These 'thickheads' play the same rÔle in the ant-state for which they are cast in cultured communities. They look after the order of the march, and allow none to turn either to the right or left. The least confusion in the regularity of the march makes them turn round and put things straight again. While the procession of the brown workers streams on unceasingly with a swarming motion, the 'officers,' as the natives call these thickheads, run constantly backwards and forwards, ready to take the command on meeting any difficulty. The crossing of streams by these creatures is the most interesting point. If the watercourse be narrow, the thickheads soon find trees, the branches of which meet on the bank on either side, and after a short halt the column set themselves in motion over these bridges, rearranging themselves in the narrow train with marvellous quickness on reaching the further side. But if no natural bridge be available for the passage, they travel along the bank of the river until they arrive at a flat sandy shore. Each ant now seizes a bit of dry wood, pulls it into the water, and mounts thereupon. The hinder rows push the front ones even further out, holding on to the wood with their feet and to their comrades with their jaws. In a short time the water is covered with ants, and when the raft has grown too large to be held together by the small creatures' strength, a part breaks itself off and begins the journey across, while the ants left on the bank busily pull their bits of wood into the water, and work at enlarging the ferry-boat until it again breaks. This is repeated as long as an ant remains on shore. I had often heard described this method of crossing rivers, but in the year 1859 I had the opportunity of seeing it for myself.

It is remarkable that the military or driving ants of Africa exhibit precisely similar devices for the bridging of streams, namely, by forming a chain of individuals over which the others pass. By means of similar chains they also let themselves down from trees. It must be observed, however, that these and all the above observations, being independently made and separately recorded, serve to corroborate one another so strongly that we can entertain no reasonable doubt concerning the wonderful facts which they convey.

I shall now bring these numerous instances to a close with a quotation from Mr. Belt, which reveals in the most unequivocal manner surprising powers of observation and rational action on the part of the leaf-cutting ants of South America, whose general habits we have already considered:—

A nest was made near one of our tramways, and to get to the trees the ants had to cross the rails, over which the waggons were continually passing and repassing. Every time they came along a number of ants were crushed to death. They persevered in crossing for some time, but at last set to work and tunnelled underneath each rail. One day, when the waggons were not running, I stopped up the tunnels with stones; but although great numbers carrying leaves were thus cut off from the nest, they would not cross the rails, but set to work making fresh tunnels underneath them.

Anatomy and Physiology of Nerve-centres and Sense-organs.

The foregoing facts concerning the intelligence of ants fully justifies Mr. Darwin's observation that 'the brain of an ant is one of the most marvellous atoms of matter in the world, perhaps more so than the brain of a man.' It may therefore be interesting in this particular case to depart from the lines otherwise laid down throughout the present work, and to devote a short section to the anatomy and physiology of this nerve-centre with its appended organs of sense.

The brain of an ant, then, is proportionally larger than that of any other insect. (See Titus Graber, 'Insects,' vol. i. p. 255.) In structure, also, the brain of an ant is in advance of that of other insects, its nearest analogue being the brain of a bee. The superiority of development is particularly remarkable with reference to the 'stalked bodies' of Dujardin; and these are largest in neuter workers, which are the most intelligent members of the community.

Injury of the brain causes, as in higher animals, tetanic spasms and involuntary reflex movements, followed by stupefaction.

An ant, whose brain has been perforated by the pointed mandibles of an amazon, remains as though nailed to its place; a shudder runs from time to time through its body, and one of its legs is lifted at regular intervals. It occasionally makes a short and quick step, as though driven by an unseen spring, but, like that of an automaton, aimless and objectless. If it is pulled, it makes a movement of avoidance, but falls back into its stupefied condition as soon as it is released. It is no longer capable of action consciously directed to a given object; it neither tries to escape, nor to attack, nor to go back to its home, nor to rejoin its companions, nor to walk away; it feels neither heat nor cold, it knows neither fear nor desire for food. It is merely an automatic and reflex machine, and is exactly similar to one of those pigeons from which Flourens removed the hemispheres of the cerebrum. Just in the same way behaves the body of an ant from which the head has been taken away. In the numerous fights between amazons and other ants, countless cases have been observed of slight injury to the brain, which have caused the most remarkable phenomena. Many of the wounded were seized with a mad rage, and flung themselves at every one that came in their way, whether friend or foe. Others assumed an appearance of indifference, and walked serenely about in the midst of the fighting. Others exhibited a sudden failure of strength; but they still recognised their enemies, approached them, and tried to bite them in cold blood, in a way quite foreign to the behaviour of healthy ants. They were also often observed to run round and round in a circle, the motion resembling the manÈge, or riding-school action of mammals, when one of the crura cerebri has been removed.

If an ant is cut in half through the thorax, so that the great nerve ganglia of the pro-thorax remain untouched, the behaviour of the head shows that intelligence also remains untouched. Ants mutilated in this way try to go forwards with their two remaining legs, and beg with their antennÆ for their companions' aid. If one of these latter lets itself be stopped, then we observe a lively interchange of thanks and sympathy expressed by the actively moving antennÆ. Forel placed near to each other two such mutilated bodies of the F. rufibarbis. They conversed with each other in the above-described way, and appeared each to beg for help. But when he put in some similarly mutilated ants of a hostile species, F. sanguinea, the picture was changed; war broke out between these cripples just in the same way and with the same fury as between perfect ants.[44]

The antennÆ appear to be the most important of the sense-organs, as their removal produces an extraordinary disturbance in the intelligence of the animal. An ant so mutilated can no longer find its way or recognise companions, and therefore is unable to distinguish between friends and foes. It is also unable to find food, ceases to engage in any labour, and loses all its regard for larvÆ, remaining permanently quiet and almost motionless. A somewhat similar disturbance, or rather destruction, of the mental faculties is observable as a result of the same mutilation in the case of bees.[45]


                                                                                                                                                                                                                                                                                                           

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