The Formation of Vegetable Mould Through the Action of Worms · Charles Darwin
CHAPTER II. HABITS OF WORMS—_continued_.
Manner in which worms seize objects—Their power of suction—The instinct of plugging up the mouths of their burrows—Stones piled over the burrows—The advantages thus gained—Intelligence shown by worms in their manner of plugging up their burrows—Various kinds of leaves and other objects thus used—Triangles of paper—Summary of reasons for believing that worms exhibit some intelligence—Means by which they excavate their burrows, by pushing away the earth and swallowing it—Earth also swallowed for the nutritious matter which it contains—Depth to which worms burrow, and the construction of their burrows—Burrows lined with castings, and in the upper part with leaves—The lowest part paved with little stones or seeds—Manner in which the castings are ejected—The collapse of old burrows—Distribution of worms—Tower-like castings in Bengal—Gigantic castings on the Nilgiri Mountains—Castings ejected in all countries.
IN the pots in which worms were kept, leaves were pinned down to the soil, and at night the manner in which they were seized could be observed. The worms always endeavoured to drag the leaves towards their burrows; and they tore or sucked off small fragments, whenever the leaves were sufficiently tender. They generally seized the thin edge of a leaf with their mouths, between the projecting upper and lower lip; the thick and strong pharynx being at the same time, as Perrier remarks, pushed forward within their bodies, so as to afford a point of resistance for the upper lip. In the case of broad flat objects they acted in a wholly different manner. The pointed anterior extremity of the body, after being brought into contact with an object of this kind, was drawn within the adjoining rings, so that it appeared truncated and became as thick as the rest of the body. This part could then be seen to swell a little; and this, I believe, is due to the pharynx being pushed a little forwards. Then by a slight withdrawal of the pharynx or by its expansion, a vacuum was produced beneath the truncated slimy end of the body whilst in contact with the object; and by this means the two adhered firmly together. {53} That under these circumstances a vacuum was produced was plainly seen on one occasion, when a large worm lying beneath a flaccid cabbage leaf tried to drag it away; for the surface of the leaf directly over the end of the worm’s body became deeply pitted. On another occasion a worm suddenly lost its hold on a flat leaf; and the anterior end of the body was momentarily seen to be cup-formed. Worms can attach themselves to an object beneath water in the same manner; and I saw one thus dragging away a submerged slice of an onion-bulb.
The edges of fresh or nearly fresh leaves affixed to the ground were often nibbled by the worms; and sometimes the epidermis and all the parenchyma on one side was gnawed completely away over a considerable space; the epidermis alone on the opposite side being left quite clean. The veins were never touched, and leaves were thus sometimes partly converted into skeletons. As worms have no teeth and as their mouths consist of very soft tissue, it may be presumed that they consume by means of suction the edges and the parenchyma of fresh leaves, after they have been softened by the digestive fluid. They cannot attack such strong leaves as those of sea-kale or large and thick leaves of ivy; though one of the latter after it had become rotten was reduced in parts to the state of a skeleton.
Worms seize leaves and other objects, not only to serve as food, but for plugging up the mouths of their burrows; and this is one of their strongest instincts. They sometimes work so energetically that Mr. D. F. Simpson, who has a small walled garden where worms abound in Bayswater, informs me that on a calm damp evening he there heard so extraordinary a rustling noise from under a tree from which many leaves had fallen, that he went out with a light and discovered that the noise was caused by many worms dragging the dry leaves and squeezing them into the burrows. Not only leaves, but petioles of many kinds, some flower-peduncles, often decayed twigs of trees, bits of paper, feathers, tufts of wool and horse-hairs are dragged into their burrows for this purpose. I have seen as many as seventeen petioles of a Clematis projecting from the mouth of one burrow, and ten from the mouth of another. Some of these objects, such as the petioles just named, feathers, &c., are never gnawed by worms. In a gravel-walk in my garden I found many hundred leaves of a pine-tree (_P. austriaca_ or _nigricans_) drawn by their bases into burrows. The surfaces by which these leaves are articulated to the branches are shaped in as peculiar a manner as is the joint between the leg-bones of a quadruped; and if these surfaces had been in the least gnawed, the fact would have been immediately visible, but there was no trace of gnawing. Of ordinary dicotyledonous leaves, all those which are dragged into burrows are not gnawed. I have seen as many as nine leaves of the lime-tree drawn into the same burrow, and not nearly all of them had been gnawed; but such leaves may serve as a store for future consumption. Where fallen leaves are abundant, many more are sometimes collected over the mouth of a burrow than can be used, so that a small pile of unused leaves is left like a roof over those which have been partly dragged in.
A leaf in being dragged a little way into a cylindrical burrow is necessarily much folded or crumpled. When another leaf is drawn in, this is done exteriorly to the first one, and so on with the succeeding leaves; and finally all become closely folded and pressed together. Sometimes the worm enlarges the mouth of its burrow, or makes a fresh one close by, so as to draw in a still larger number of leaves. They often or generally fill up the interstices between the drawn-in leaves with moist viscid earth ejected from their bodies; and thus the mouths of the burrows are securely plugged. Hundreds of such plugged burrows may be seen in many places, especially during the autumnal and early winter months. But, as will hereafter be shown, leaves are dragged into the burrows not only for plugging them up and for food, but for the sake of lining the upper part or mouth.
When worms cannot obtain leaves, petioles, sticks, &c., with which to plug up the mouths of their burrows, they often protect them by little heaps of stones; and such heaps of smooth rounded pebbles may frequently be seen on gravel-walks. Here there can be no question about food. A lady, who was interested in the habits of worms, removed the little heaps of stones from the mouths of several burrows and cleared the surface of the ground for some inches all round. She went out on the following night with a lantern, and saw the worms with their tails fixed in their burrows, dragging the stones inwards by the aid of their mouths, no doubt by suction. “After two nights some of the holes had 8 or 9 small stones over them; after four nights one had about 30, and another 34 stones.” {58} One stone—which had been dragged over the gravel-walk to the mouth of a burrow weighed two ounces; and this proves how strong worms are. But they show greater strength in sometimes displacing stones in a well-trodden gravel-walk; that they do so, may be inferred from the cavities left by the displaced stones being exactly filled by those lying over the mouths of adjoining burrows, as I have myself observed.
Work of this kind is usually performed during the night; but I have occasionally known objects to be drawn into the burrows during the day. What advantage the worms derive from plugging up the mouths of their burrows with leaves, &c., or from piling stones over them, is doubtful. They do not act in this manner at the times when they eject much earth from their burrows; for their castings then serve to cover the mouths. When gardeners wish to kill worms on a lawn, it is necessary first to brush or rake away the castings from the surface, in order that the lime-water may enter the burrows. {59a} It might be inferred from this fact that the mouths are plugged up with leaves, &c., to prevent the entrance of water during heavy rain; but it may be urged against this view that a few, loose, well-rounded stones are ill-adapted to keep out water. I have moreover seen many burrows in the perpendicularly cut turf-edgings to gravel-walks, into which water could hardly flow, as well plugged as burrows on a level surface.
It is not probable that the plugs or piles of stones serve to conceal the burrows from scolopendras, which, according to Hoffmeister, {59b} are the bitterest enemies of worms, or from the larger species of Carabus and Staphylinus which attack them ferociously, for these animals are nocturnal, and the burrows are opened at night. May not worms when the mouth of the burrow is protected be able to remain with safety with their heads close to it, which we know that they like to do, but which costs so many of them their lives? Or may not the plugs check the free ingress of the lowest stratum of air, when chilled by radiation at night, from the surrounding ground and herbage? I am inclined to believe in this latter view: firstly, because when worms were kept in pots in a room with a fire, in which case cold air could not enter the burrows, they plugged them up in a slovenly manner; and secondarily, because they often coat the upper part of their burrows with leaves, apparently to prevent their bodies from coming into close contact with the cold damp earth. Mr. E. Parfitt has suggested to me that the mouths of the burrows are closed in order that the air within them may be kept thoroughly damp, and this seems the most probable explanation of the habit. But the plugging-up process may serve for all the above purposes.
Whatever the motive may be, it appears that worms much dislike leaving the mouths of their burrows open. Nevertheless they will reopen them at night, whether or not they can afterwards close them. Numerous open burrows may be seen on recently-dug ground, for in this case the worms eject their castings in cavities left in the ground, or in the old burrows instead of piling them over the mouths of their burrows, and they cannot collect objects on the surface by which the mouths might be protected. So again on a recently disinterred pavement of a Roman villa at Abinger (hereafter to be described) the worms pertinaciously opened their burrows almost every night, when these had been closed by being trampled on, although they were rarely able to find a few minute stones wherewith to protect them.
_Intelligence shown by worms in their manner of plugging up their burrows_.—If a man had to plug up a small cylindrical hole, with such objects as leaves, petioles or twigs, he would drag or push them in by their pointed ends; but if these objects were very thin relatively to the size of the hole, he would probably insert some by their thicker or broader ends. The guide in his case would be intelligence. It seemed therefore worth while to observe carefully how worms dragged leaves into their burrows; whether by their tips or bases or middle parts. It seemed more especially desirable to do this in the case of plants not natives to our country; for although the habit of dragging leaves into their burrows is undoubtedly instinctive with worms, yet instinct could not tell them how to act in the case of leaves about which their progenitors knew nothing. If, moreover, worms acted solely through instinct or an unvarying inherited impulse, they would draw all kinds of leaves into their burrows in the same manner. If they have no such definite instinct, we might expect that chance would determine whether the tip, base or middle was seized. If both these alternatives are excluded, intelligence alone is left; unless the worm in each case first tries many different methods, and follows that alone which proves possible or the most easy; but to act in this manner and to try different methods makes a near approach to intelligence.
In the first place 227 withered leaves of various kinds, mostly of English plants, were pulled out of worm-burrows in several places. Of these, 181 had been drawn into the burrows by or near their tips, so that the foot-stalk projected nearly upright from the mouth of the burrow; 20 had been drawn in by their bases, and in this case the tips projected from the burrows; and 26 had been seized near the middle, so that these had been drawn in transversely and were much crumpled. Therefore 80 per cent. (always using the nearest whole number) had been drawn in by the tip, 9 per cent. by the base or foot-stalk, and 11 per cent. transversely or by the middle. This alone is almost sufficient to show that chance does not determine the manner in which leaves are dragged into the burrows.
Of the above 227 leaves, 70 consisted of the fallen leaves of the common lime-tree, which is almost certainly not a native of England. These leaves are much acuminated towards the tip, and are very broad at the base with a well-developed foot-stalk. They are thin and quite flexible when half-withered. Of the 70, 79 per cent. had been drawn in by or near the tip; 4 per cent. by or near the base; and 17 per cent. transversely or by the middle. These proportions agree very closely, as far as the tip is concerned, with those before given. But the percentage drawn in by the base is smaller, which may be attributed to the breadth of the basal part of the blade. We here, also, see that the presence of a foot-stalk, which it might have been expected would have tempted the worms as a convenient handle, has little or no influence in determining the manner in which lime leaves are dragged into the burrows. The considerable proportion, viz., 17 per cent., drawn in more or less transversely depends no doubt on the flexibility of these half-decayed leaves. The fact of so many having been drawn in by the middle, and of some few having been drawn in by the base, renders it improbable that the worms first tried to draw in most of the leaves by one or both of these methods, and that they afterwards drew in 79 per cent. by their tips; for it is clear that they would not have failed in drawing them in by the base or middle.
The leaves of a foreign plant were next searched for, the blades of which were not more pointed towards the apex than towards the base. This proved to be the case with those of a laburnum (a hybrid between _Cytisus alpinus_ and _laburnum_) for on doubling the terminal over the basal half, they generally fitted exactly; and when there was any difference, the basal half was a little the narrower. It might, therefore, have been expected that an almost equal number of these leaves would have been drawn in by the tip and base, or a slight excess in favour of the latter. But of 73 leaves (not included in the first lot of 227) pulled out of worm-burrows, 63 per cent. had been drawn in by the tip; 27 per cent. by the base, and 10 per cent. transversely. We here see that a far larger proportion, viz., 27 per cent. were drawn in by the base than in the case of lime leaves, the blades of which are very broad at the base, and of which only 4 per cent. had thus been drawn in. We may perhaps account for the fact of a still larger proportion of the laburnum leaves not having been drawn in by the base, by worms having acquired the habit of generally drawing in leaves by their tips and thus avoiding the foot-stalk. For the basal margin of the blade in many kinds of leaves forms a large angle with the foot-stalk; and if such a leaf were drawn in by the foot-stalk, the basal margin would come abruptly into contact with the ground on each side of the burrow, and would render the drawing in of the leaf very difficult.
Nevertheless worms break through their habit of avoiding the foot-stalk, if this part offers them the most convenient means for drawing leaves into their burrows. The leaves of the endless hybridised varieties of the Rhododendron vary much in shape; some are narrowest towards the base and others towards the apex. After they have fallen off, the blade on each side of the midrib often becomes curled up while drying, sometimes along the whole length, sometimes chiefly at the base, sometimes towards the apex. Out of 28 fallen leaves on one bed of peat in my garden, no less than 23 were narrower in the basal quarter than in the terminal quarter of their length; and this narrowness was chiefly due to the curling in of the margins. Out of 36 fallen leaves on another bed, in which different varieties of the Rhododendron grew, only 17 were narrower towards the base than towards the apex.
My son William, who first called my attention to this case, picked up 237 fallen leaves in his garden (where the Rhododendron grows in the natural soil) and of these 65 per cent. could have been drawn by worms into their burrows more easily by the base or foot-stalk than by the tip; and this was partly due to the shape of the leaf and in a less degree to the curling in of the margins: 27 per cent. could have been drawn in more easily by the tip than by the base: and 8 per cent. with about equal ease by either end. The shape of a fallen leaf ought to be judged of before one end has been drawn into a burrow, for after this has happened, the free end, whether it be the base or apex, will dry more quickly than the end imbedded in the damp ground; and the exposed margins of the free end will consequently tend to become more curled inwards than they were when the leaf was first seized by the worm. My son found 91 leaves which had been dragged by worms into their burrows, though not to a great depth; of these 66 per cent. had been drawn in by the base or foot-stalk; and 34 per cent. by the tip.
In this case, therefore, the worms judged with a considerable degree of correctness how best to draw the withered leaves of this foreign plant into their burrows; notwithstanding that they had to depart from their usual habit of avoiding the foot-stalk. On the gravel-walks in my garden a very large number of leaves of three species of Pinus (_P. austriaca_, _nigricans_ and _sylvestris_) are regularly drawn into the mouths of worm burrows. These leaves consist of two so-called needles, which are of considerable length in the two first and short in the last named species, and are united to a common base; and it is by this part that they are almost invariably drawn into the burrows. I have seen only two or at most three exceptions to this rule with worms in a state of nature. As the sharply pointed needles diverge a little, and as several leaves are drawn into the same burrow, each tuft forms a perfect _chevaux de frise_. On two occasions many of these tufts were pulled up in the evening, but by the following morning fresh leaves had been pulled in, and the burrows were again well protected. These leaves could not be dragged into the burrows to any depth, except by their bases, as a worm cannot seize hold of the two needles at the same time, and if one alone were seized by the apex, the other would be pressed against the ground and would resist the entry of the seized one. This was manifest in the above mentioned two or three exceptional cases. In order, therefore, that worms should do their work well, they must drag pine-leaves into their burrows by their bases, where the two needles are conjoined. But how they are guided in this work is a perplexing question.
This difficulty led my son Francis and myself to observe worms in confinement during several nights by the aid of a dim light, while they dragged the leaves of the above named pines into their burrows. They moved the anterior extremities of their bodies about the leaves, and on several occasions when they touched the sharp end of a needle they withdrew suddenly as if pricked. But I doubt whether they were hurt, for they are indifferent to very sharp objects, and will swallow even rose-thorns and small splinters of glass. It may also be doubted, whether the sharp ends of the needles serve to tell them that this is the wrong end to seize; for the points were cut off many leaves for a length of about one inch, and fifty-seven of them thus treated were drawn into the burrows by their bases, and not one by the cut-off ends. The worms in confinement often seized the needles near the middle and drew them towards the mouths of their burrows; and one worm tried in a senseless manner to drag them into the burrow by bending them. They sometimes collected many more leaves over the mouths of their burrows (as in the case formerly mentioned of lime-leaves) than could enter them.
On other occasions, however, they behaved very differently; for as soon as they touched the base of a pine-leaf, this was seized, being sometimes completely engulfed in their mouths, or a point very near the base was seized, and the leaf was then quickly dragged or rather jerked into their burrows. It appeared both to my son and myself as if the worms instantly perceived as soon as they had seized a leaf in the proper manner. Nine such cases were observed, but in one of them the worm failed to drag the leaf into its burrow, as it was entangled by other leaves lying near. In another case a leaf stood nearly upright with the points of the needles partly inserted into a burrow, but how placed there was not seen; and then the worm reared itself up and seized the base, which was dragged into the mouth of the burrow by bowing the whole leaf. On the other hand, after a worm had seized the base of a leaf, this was on two occasions relinquished from some unknown motive.
As already remarked, the habit of plugging up the mouths of the burrows with various objects, is no doubt instinctive in worms; and a very young one, born in one of my pots, dragged for some little distance a Scotch-fir leaf, one needle of which was as long and almost as thick as its own body. No species of pine is endemic in this part of England, it is therefore incredible that the proper manner of dragging pine-leaves into the burrows can be instinctive with our worms. But as the worms on which the above observations were made, were dug up beneath or near some pines, which had been planted there about forty years, it was desirable to prove that their actions were not instinctive. Accordingly, pine-leaves were scattered on the ground in places far removed from any pine-tree, and 90 of them were drawn into the burrows by their bases. Only two were drawn in by the tips of the needles, and these were not real exceptions, as one was drawn in for a very short distance, and the two needles of the other cohered. Other pine-leaves were given to worms kept in pots in a warm room, and here the result was different; for out of 42 leaves drawn into the burrows, no less than 16 were drawn in by the tips of the needles. These worms, however, worked in a careless or slovenly manner; for the leaves were often drawn in to only a small depth; sometimes they were merely heaped over the mouths of the burrows, and sometimes none were drawn in. I believe that this carelessness may be accounted for either by the warmth of the air, or by its dampness, as the pots were covered by glass plates; the worms consequently did not care about plugging up their holes effectually. Pots tenanted by worms and covered with a net which allowed the free entrance of air, were left out of doors for several nights, and now 72 leaves were all properly drawn in by their bases.
Text from Project Gutenberg, public domain in the United States.