The Formation of Vegetable Mould Through the Action of Worms. Чарльз Дарвин
of Contents
Nature of the sites inhabited—Can live long under water—Nocturnal—Wander about at night—Often lie close to the mouths of their burrows, and are thus destroyed in large numbers by birds—Structure—Do not possess eyes, but can distinguish between light and darkness—Retreat rapidly when brightly illuminated, not by a reflex action—Power of attention—Sensitive to heat and cold—Completely deaf—Sensitive to vibrations and to touch—Feeble power of smell—Taste—Mental qualities—Nature of food—Omnivorous—Digestion—Leaves before being swallowed, moistened with a fluid of the nature of the pancreatic secretion—Extra-stomachal digestion—Calciferous glands, structure of—Calcareous concretions formed in the anterior pair of glands—The calcareous matter primarily an excretion, but secondarily serves to neutralise the acids generated during the digestive process.
Earth-worms are distributed throughout the world under the form of a few genera, which externally are closely similar to one another. The British species of Lumbricus have never been carefully monographed; but we may judge of their probable number from those inhabiting neighbouring countries. In Scandinavia there are eight species, according to Eisen; [8a] but two of these rarely burrow in the ground, and one inhabits very wet places or even lives under the water. We are here concerned only with the kinds which bring up earth to the surface in the form of castings. Hoffmeister says that the species in Germany are not well known, but gives the same number as Eisen, together with some strongly marked varieties. [8b]
Earth-worms abound in England in many different stations. Their castings may be seen in extraordinary numbers on commons and chalk-downs, so as almost to cover the whole surface, where the soil is poor and the grass short and thin. But they are almost or quite as numerous in some of the London parks, where the grass grows well and the soil appears rich. Even on the same field worms are much more frequent in some places than in others, without any visible difference in the nature of the soil. They abound in paved court-yards close to houses; and an instance will be given in which they had burrowed through the floor of a very damp cellar. I have seen worms in black peat in a boggy field; but they are extremely rare, or quite absent in the drier, brown, fibrous peat, which is so much valued by gardeners. On dry, sandy or gravelly tracks, where heath with some gorse, ferns, coarse grass, moss and lichens alone grow, hardly any worms can be found. But in many parts of England, wherever a path crosses a heath, its surface becomes covered with a fine short sward. Whether this change of vegetation is due to the taller plants being killed by the occasional trampling of man and animals, or to the soil being occasionally manured by the droppings from animals, I do not know. [9b] On such grassy paths worm-castings may often be seen. On a heath in Surrey, which was carefully examined, there were only a few castings on these paths, where they were much inclined; but on the more level parts, where a bed of fine earth had been washed down from the steeper parts and had accumulated to a thickness of a few inches, worm-castings abounded. These spots seemed to be overstocked with worms, so that they had been compelled to spread to a distance of a few feet from the grassy paths, and here their castings had been thrown up among the heath; but beyond this limit, not a single casting could be found. A layer, though a thin one, of fine earth, which probably long retains some moisture, is in all cases, as I believe, necessary for their existence; and the mere compression of the soil appears to be in some degree favourable to them, for they often abound in old gravel walks, and in foot-paths across fields.
Beneath large trees few castings can be found during certain seasons of the year, and this is apparently due to the moisture having been sucked out of the ground by the innumerable roots of the trees; for such places may be seen covered with castings after the heavy autumnal rains. Although most coppices and woods support many worms, yet in a forest of tall and ancient beech-trees in Knole Park, where the ground beneath was bare of all vegetation, not a single casting could be found over wide spaces, even during the autumn. Nevertheless, castings were abundant on some grass-covered glades and indentations which penetrated this forest. On the mountains of North Wales and on the Alps, worms, as I have been informed, are in most places rare; and this may perhaps be due to the close proximity of the subjacent rocks, into which worms cannot burrow during the winter so as to escape being frozen. Dr. McIntosh, however, found worm-castings at a height of 1500 feet on Schiehallion in Scotland. They are numerous on some hills near Turin at from 2000 to 3000 feet above the sea, and at a great altitude on the Nilgiri Mountains in South India and on the Himalaya.
Earth-worms must be considered as terrestrial animals, though they are still in one sense semi-aquatic, like the other members of the great class of annelids to which they belong. M. Perrier found that their exposure to the dry air of a room for only a single night was fatal to them. On the other hand he kept several large worms alive for nearly four months, completely submerged in water. [11] During the summer when the ground is dry, they penetrate to a considerable depth and cease to work, as they do during the winter when the ground is frozen. Worms are nocturnal in their habits, and at night may be seen crawling about in large numbers, but usually with their tails still inserted in their burrows. By the expansion of this part of their bodies, and with the help of the short, slightly reflexed bristles, with which their bodies are armed, they hold so fast that they can seldom be dragged out of the ground without being torn into pieces. [12] During the day they remain in their burrows, except at the pairing season, when those which inhabit adjoining burrows expose the greater part of their bodies for an hour or two in the early morning. Sick individuals, which are generally affected by the parasitic larvæ of a fly, must also be excepted, as they wander about during the day and die on the surface. After heavy rain succeeding dry weather, an astonishing number of dead worms may sometimes be seen lying on the ground. Mr. Galton informs me that on one such occasion (March, 1881), the dead worms averaged one for every two and a half paces in length on a walk in Hyde Park, four paces in width. He counted no less than 45 dead worms in one place in a length of sixteen paces. From the facts above given, it is not probable that these worms could have been drowned, and if they had been drowned they would have perished in their burrows. I believe that they were already sick, and that their deaths were merely hastened by the ground being flooded.
It has often been said that under ordinary circumstances healthy worms never, or very rarely, completely leave their burrows at night; but this is an error, as White of Selborne long ago knew. In the morning, after there has been heavy rain, the film of mud or of very fine sand over gravel-walks is often plainly marked with their tracks. I have noticed this from August to May, both months included, and it probably occurs during the two remaining months of the year when they are wet. On these occasions, very few dead worms could anywhere be seen. On January 31, 1881, after a long-continued and unusually severe frost with much snow, as soon as a thaw set in, the walks were marked with innumerable tracks. On one occasion, five tracks were counted crossing a space of only an inch square. They could sometimes be traced either to or from the mouths of the burrows in the gravel-walks, for distances between 2 or 3 up to 15 yards. I have never seen two tracks leading to the same burrow; nor is it likely, from what we shall presently see of their sense-organs, that a worm could find its way back to its burrow after having once left it. They apparently leave their burrows on a voyage of discovery, and thus they find new sites to inhabit.
Morren states [14] that worms often lie for hours almost motionless close beneath the mouths of their burrows. I have occasionally noticed the same fact with worms kept in pots in the house; so that by looking down into their burrows, their heads could just be seen. If the ejected earth or rubbish over the burrows be suddenly removed, the end of the worm’s body may very often be seen rapidly retreating. This habit of lying near the surface leads to their destruction to an immense extent. Every morning during certain seasons of the year, the thrushes and blackbirds on all the lawns throughout the country draw out of their holes an astonishing number of worms, and this they could not do, unless they lay close to the surface. It is not probable that worms behave in this manner for the sake of breathing fresh air, for we have seen that they can live for a long time under water. I believe that they lie near the surface for the sake of warmth, especially in the morning; and we shall hereafter find that they often coat the mouths of their burrows with leaves, apparently to prevent their bodies from coming into close contact with the cold damp earth. It is said that they completely close their burrows during the winter.
Structure.—A few remarks must be made on