THE FORMATION OF VEGETABLE MOULD THROUGH THE ACTION OF WORMS
TEXT
Stonehenge. On the whole it appears that the crowns and furrows on land
formerly ploughed, but now covered with grass, tend slowly to disappear
when the surface is inclined; and this is probably in large part due to
the action of worms; but that the crowns and furrows last for a very long
time when the surface is nearly level.
_Formation and amount of mould over the Chalk Formation_.—Worm-castings
are often ejected in extraordinary numbers on steep, grass-covered
slopes, where the Chalk comes close to the surface, as my son William
observed near Winchester and elsewhere. If such castings are largely
washed away during heavy rains, it is difficult to understand at first
how any mould can still remain on our Downs, as there does not appear any
evident means for supplying the loss. There is, moreover, another cause
of loss, namely, in the percolation of the finer particles of earth into
the fissures in the chalk and into the chalk itself. These
considerations led me to doubt for a time whether I had not exaggerated
the amount of fine earth which flows or rolls down grass-covered slopes
under the form of castings; and I sought for additional information. In
some places, the castings on Chalk Downs consist largely of calcareous
matter, and here the supply is of course unlimited. But in other places,
for instance on a part of Teg Down near Winchester, the castings were all
black and did not effervesce with acids. The mould over the chalk was
here only from 3 to 4 inches in thickness. So again on the plain near
Stonehenge, the mould, apparently free from calcareous matter, averaged
rather less than 3½ inches in thickness. Why worms should penetrate and
bring up chalk in some places and not in others I do not know.
In many districts where the land is nearly level, a bed several feet in
thickness of red clay full of unworn flints overlies the Upper Chalk.
This overlying matter, the surface of which has been converted into
mould, consists of the undissolved residue from the chalk. It may be
well here to recall the case of the fragments of chalk buried beneath
worm-castings on one of my fields, the angles of which were so completely
rounded in the course of 29 years that the fragments now resembled
water-worn pebbles. This must have been effected by the carbonic acid in
the rain and in the ground, by the humus-acids, and by the corroding
power of living roots. Why a thick mass of residue has not been left on
the Chalk, wherever the land is nearly level, may perhaps be accounted
for by the percolation of the fine particles into the fissures, which are
often present in the chalk and are either open or are filled up with
impure chalk, or into the solid chalk itself. That such percolation
occurs can hardly be doubted. My son collected some powdered and
fragmentary chalk beneath the turf near Winchester; the former was found
by Colonel Parsons, R. E., to contain 10 per cent., and the fragments 8
per cent. of earthy matter. On the flanks of the escarpment near Abinger
in Surrey, some chalk close beneath a layer of flints, 2 inches in
thickness and covered by 8 inches of mould, yielded a residue of 3.7 per
cent. of earthy matter. On the other hand the Upper Chalk properly
contains, as I was informed by the late David Forbes who had made many
analyses, only from 1 to 2 per cent. of earthy matter; and two samples
from pits near my house contained 1.3 and 0.6 per cent. I mention these
latter cases because, from the thickness of the overlying bed of red clay
with flints, I had imagined that the underlying chalk might here be less
pure than elsewhere. The cause of the residue accumulating more in some
places than in others, may be attributed to a layer of argillaceous
matter having been left at an early period on the chalk, and this would
check the subsequent percolation of earthy matter into it.
From the facts now given we may conclude that castings ejected on our
Chalk Downs suffer some loss by the percolation of their finer matter
into the chalk. But such impure superficial chalk, when dissolved, would
leave a larger supply of earthy matter to be added to the mould than in
the case of pure chalk. Besides the loss caused by percolation, some
fine earth is certainly washed down the sloping grass-covered surfaces of
our Downs. The washing-down process, however, will be checked in the
course of time; for although I do not know how thin a layer of mould
suffices to support worms, yet a limit must at last be reached; and then
their castings would cease to be ejected or would become scanty.
The following cases show that a considerable quantity of fine earth is
washed down. The thickness of the mould was measured at points 12 yards
apart across a small valley in the Chalk near Winchester. The sides
sloped gently at first; then became inclined at about 20°; then more
gently to near the bottom, which transversely was almost level and about
50 yards across. In the bottom, the mean thickness of the mould from
five measurements was 8.3 inches; whilst on the sides of the valley,
where the inclination varied between 14° and 20°, its mean thickness was
rather less than 3.5 inches. As the turf-covered bottom of the valley
sloped at an angle of only between 2° and 3°, it is probable that most of
the 8.3-inch layer of mould had been washed down from the flanks of the
valley, and not from the upper part. But as a shepherd said that he had
seen water flowing in this valley after the sudden thawing of snow, it is
possible that some earth may have been brought down from the upper part;
or, on the other hand, that some may have been carried further down the
valley. Closely similar results, with respect to the thickness of the
mould, were obtained in a neighbouring valley.
St. Catherine’s Hill, near Winchester, is 327 feet in height, and
consists of a steep cone of chalk about ¼ of a mile in diameter. The
upper part was converted by the Romans, or, as some think, by the ancient
Britons, into an encampment, by the excavation of a deep and broad ditch
all round it. Most of the chalk removed during the work was thrown
upwards, by which a projecting bank was formed; and this effectually
prevents worm-castings (which are numerous in parts), stones, and other
objects from being washed or rolled into the ditch. The mould on the
upper and fortified part of the hill was found to be in most places only
from 2½ to 3½ inches in thickness; whereas it had accumulated at the foot
of the embankment above the ditch to a thickness in most places of from 8
to 9½ inches. On the embankment itself the mould was only 1 to 1½ inch
in thickness; and within the ditch at the bottom it varied from 2½ to 3½,
but was in one spot 6 inches in thickness. On the north-west side of the
hill, either no embankment had ever been thrown up above the ditch, or it
had subsequently been removed; so that here there was nothing to prevent
worm-castings, earth and stones being washed into the ditch, at the
bottom of which the mould formed a layer from 11 to 22 inches in
thickness. It should however be stated that here and on other parts of
the slope, the bed of mould often contained fragments of chalk and flint
which had obviously rolled down at different times from above. The
interstices in the underlying fragmentary chalk were also filled up with
mould.
My son examined the surface of this hill to its base in a south-west
direction. Beneath the great ditch, where the slope was about 24°, the
mould was very thin, namely, from 1½ to 2½ inches; whilst near the base,
where the slope was only 3° to 4°, it increased to between 8 and 9 inches
in thickness. We may therefore conclude that on this artificially
modified hill, as well as in the natural valleys of the neighbouring
Chalk Downs, some fine earth, probably derived in large part from
worm-castings, is washed down, and accumulates in the lower parts,
notwithstanding the percolation of an unknown quantity into the
underlying chalk; a supply of fresh earthy matter being afforded by the
dissolution of the chalk through atmospheric and other agencies.
CHAPTER VII.
CONCLUSION.
Summary of the part which worms have played in the history of the
world—Their aid in the disintegration of rocks—In the denudation of the
land—In the preservation of ancient remains—In the preparation of the
soil for the growth of plants—Mental powers of worms—Conclusion.
WORMS have played a more important part in the history of the world than
most persons would at first suppose. In almost all humid countries they
are extraordinarily numerous, and for their size possess great muscular
power. In many parts of England a weight of more than ten tons (10,516
kilogrammes) of dry earth annually passes through their bodies and is
brought to the surface on each acre of land; so that the whole
superficial bed of vegetable mould passes through their bodies in the
course of every few years. From the collapsing of the old burrows the
mould is in constant though slow movement, and the particles composing it
are thus rubbed together. By these means fresh surfaces are continually
exposed to the action of the carbonic acid in the soil, and of the
humus-acids which appear to be still more efficient in the decomposition
of rocks. The generation of the humus-acids is probably hastened during
the digestion of the many half-decayed leaves which worms consume. Thus
the particles of earth, forming the superficial mould, are subjected to
conditions eminently favourable for their decomposition and
disintegration. Moreover, the particles of the softer rocks suffer some
amount of mechanical trituration in the muscular gizzards of worms, in
which small stones serve as mill-stones.
The finely levigated castings, when brought to the surface in a moist
condition, flow during rainy weather down any moderate slope; and the
smaller particles are washed far down even a gently inclined surface.
Castings when dry often crumble into small pellets and these are apt to
roll down any sloping surface. Where the land is quite level and is
covered with herbage, and where the climate is humid so that much dust
cannot be blown away, it appears at first sight impossible that there
should be any appreciable amount of sub-aerial denudation; but
worm-castings are blown, especially whilst moist and viscid, in one
uniform direction by the prevalent winds which are accompanied by rain.
By these several means the superficial mould is prevented from
accumulating to a great thickness; and a thick bed of mould checks in
many ways the disintegration of the underlying rocks and fragments of
rock.
The removal of worm-castings by the above means leads to results which
are far from insignificant. It has been shown that a layer of earth, 0.2
of an inch in thickness, is in many places annually brought to the
surface; and if a small part of this amount flows, or rolls, or is
washed, even for a short distance, down every inclined surface, or is
repeatedly blown in one direction, a great effect will be produced in the
course of ages. It was found by measurements and calculations that on a
surface with a mean inclination of 9° 26′, 2.4 cubic inches of earth
which had been ejected by worms crossed, in the course of a year, a
horizontal line one yard in length; so that 240 cubic inches would cross
a line 100 yards in length. This latter amount in a damp state would
weigh 11½ pounds. Thus a considerable weight of earth is continually
moving down each side of every valley, and will in time reach its bed.
Finally this earth will be transported by the streams flowing in the
valleys into the ocean, the great receptacle for all matter denuded from
the land. It is known from the amount of sediment annually delivered
into the sea by the Mississippi, that its enormous drainage-area must on
an average be lowered .00263 of an inch each year; and this would suffice
in four and half million years to lower the whole drainage-area to the
level of the sea-shore. So that, if a small fraction of the layer of
fine earth, 0.2 of an inch in thickness, which is annually brought to the
surface by worms, is carried away, a great result cannot fail to be
produced within a period which no geologist considers extremely long.
* * * * *
Archæologists ought to be grateful to worms, as they protect and preserve
for an indefinitely long period every object, not liable to decay, which
is dropped on the surface of the land, by burying it beneath their
castings. Thus, also, many elegant and curious tesselated pavements and
other ancient remains have been preserved; though no doubt the worms have
in these cases been largely aided by earth washed and blown from the
adjoining land, especially when cultivated. The old tesselated pavements
have, however, often suffered by having subsided unequally from being
unequally undermined by the worms. Even old massive walls may be
undermined and subside; and no building is in this respect safe, unless
the foundations lie 6 or 7 feet beneath the surface, at a depth at which
worms cannot work. It is probable that many monoliths and some old walls
have fallen down from having been undermined by worms.
* * * * *
Worms prepare the ground {284} in an excellent manner for the growth of
fibrous-rooted plants and for seedlings of all kinds. They periodically
expose the mould to the air, and sift it so that no stones larger than
the particles which they can swallow are left in it. They mingle the
whole intimately together, like a gardener who prepares fine soil for his
choicest plants. In this state it is well fitted to retain moisture and
to absorb all soluble substances, as well as for the process of
nitrification. The bones of dead animals, the harder parts of insects,
the shells of land-molluscs, leaves, twigs, &c., are before long all
buried beneath the accumulated castings of worms, and are thus brought in
a more or less decayed state within reach of the roots of plants. Worms
likewise drag an infinite number of dead leaves and other parts of plants
into their burrows, partly for the sake of plugging them up and partly as
food.
The leaves which are dragged into the burrows as food, after being torn
into the finest shreds, partially digested, and saturated with the
intestinal and urinary secretions, are commingled with much earth. This
earth forms the dark coloured, rich humus which almost everywhere covers
the surface of the land with a fairly well-defined layer or mantle.
Hensen {285} placed two worms in a vessel 18 inches in diameter, which
was filled with sand, on which fallen leaves were strewed; and these were
soon dragged into their burrows to a depth of 3 inches. After about 6
weeks an almost uniform layer of sand, a centimeter (0.4 inch) in
thickness, was converted into humus by having passed through the
alimentary canals of these two worms. It is believed by some persons
that worm-burrows, which often penetrate the ground almost
perpendicularly to a depth of 5 or 6 feet, materially aid in its
drainage; notwithstanding that the viscid castings piled over the mouths
of the burrows prevent or check the rain-water directly entering them.
They allow the air to penetrate deeply into the ground. They also
greatly facilitate the downward passage of roots of moderate size; and
these will be nourished by the humus with which the burrows are lined.
Many seeds owe their germination to having been covered by castings; and
others buried to a considerable depth beneath accumulated castings lie
dormant, until at some future time they are accidentally uncovered and
germinate.
Worms are poorly provided with sense-organs, for they cannot be said to
see, although they can just distinguish between light and darkness; they
are completely deaf, and have only a feeble power of smell; the sense of
touch alone is well developed. They can therefore learn but little about
the outside world, and it is surprising that they should exhibit some
skill in lining their burrows with their castings and with leaves, and in
the case of some species in piling up their castings into tower-like
constructions. But it is far more surprising that they should apparently
exhibit some degrees of intelligence instead of a mere blind instinctive
impulse, in their manner of plugging up the mouths of their burrows.
They act in nearly the same manner as would a man, who had to close a
cylindrical tube with different kinds of leaves, petioles, triangles of
paper, &c., for they commonly seize such objects by their pointed ends.
But with thin objects a certain number are drawn in by their broader
ends. They do not act in the same unvarying manner in all cases, as do
most of the lower animals; for instance, they do not drag in leaves by
their foot-stalks, unless the basal part of the blade is as narrow as the
apex, or narrower than it.
* * * * *
When we behold a wide, turf-covered expanse, we should remember that its
smoothness, on which so much of its beauty depends, is mainly due to all
the inequalities having been slowly levelled by worms. It is a
marvellous reflection that the whole of the superficial mould over any
such expanse has passed, and will again pass, every few years through the
bodies of worms. The plough is one of the most ancient and most valuable
of man’s inventions; but long before he existed the land was in fact
regularly ploughed, and still continues to be thus ploughed by
earth-worms. It may be doubted whether there are many other animals
which have played so important a part in the history of the world, as
have these lowly organized creatures. Some other animals, however, still
more lowly organized, namely corals, have done far more conspicuous work
in having constructed innumerable reefs and islands in the great oceans;
but these are almost confined to the tropical zones.
FOOTNOTES.
{2} ‘Leçons de Géologie Pratique,’ tom. i. 1845, p. 140.
{3} ‘Transactions Geolog. Soc.’ vol. v. p. 505. Read November 1, 1837.
{4a} ‘Histoire des progrès de la Géologie,’ tom. i. 1847, p. 224.
{4b} ‘Zeitschrift für wissenschaft. Zoologie,’ B. xxviii. 1877, p. 361.
{5} ‘Gardeners’ Chronicle,’ April 17, 1869, p. 418.
{6} Mr. Darwin’s attention was called by Professor Hensen to P. E.
Müller’s work on Humus in ‘Tidsskrift for Skovbrug,’ Band iii. Heft 1 and
2, Copenhagen, 1878. He had, however, no opportunity of consulting
Müller’s work. Dr. Müller published a second paper in 1884 in the same
periodical—a Danish journal of forestry. His results have also been
published in German, in a volume entitled ‘Studien über die natürlichen
Humusformen, unter deren Einwirkung auf Vegetation und Boden,’ 8vo.,
Berlin, 1887.
{8a} ‘Bidrag till Skandinaviens Oligochætfauna,’ 1871.
{8b} ‘Die bis jetzt bekannten Arten aus der Familie der Regenwürmer,’
1845.
{9b} There is even some reason to believe that pressure is actually
favourable to the growth of grasses, for Professor Buckman, who made many
observations on their growth in the experimental gardens of the Royal
Agricultural College, remarks (‘Gardeners’ Chronicle,’ 1854, p. 619):
“Another circumstance in the cultivation of grasses in the separate form
or small patches, is the impossibility of rolling or treading them
firmly, without which no pasture can continue good.”
{11} I shall have occasion often to refer to M. Perrier’s admirable
memoir, ‘Organisation des Lombriciens terrestres’ in ‘Archives de Zoolog.
expér.’ tom. iii. 1874, p. 372. C. F. Morren (‘De Lumbrici terrestris
Hist. Nat.’ 1829, p. 14) found that worms endured immersion for fifteen
to twenty days in summer, but that in winter they died when thus treated.
{12} Morren, ‘De Lumbrici terrestris Hist. Nat.’ &c., 1829, p. 67.
{14} ‘De Lumbrici terrestris Hist. Nat.’ &c., p. 14.
{17} Histolog. Untersuchungen über die Regenwürmer. ‘Zeitschrift für
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