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The Formation of Vegetable Mould through the action of worms with
observations of their habits

by Charles Darwin





INTRODUCTION。



The share which worms have taken in the formation of the layer of 
vegetable mould; which covers the whole surface of the land in 
every moderately humid country; is the subject of the present 
volume。  This mould is generally of a blackish colour and a few 
inches in thickness。  In different districts it differs but little 
in appearance; although it may rest on various subsoils。  The 
uniform fineness of the particles of which it is composed is one of 
its chief characteristic features; and this may be well observed in 
any gravelly country; where a recently…ploughed field immediately 
adjoins one which has long remained undisturbed for pasture; and 
where the vegetable mould is exposed on the sides of a ditch or 
hole。  The subject may appear an insignificant one; but we shall 
see that it possesses some interest; and the maxim 〃de minimis non 
curat lex;〃 does not apply to science。  Even Elie de Beaumont; who 
generally undervalues small agencies and their accumulated effects; 
remarks:  {1} 〃La couche tres…mince de la terre vegetale est un 
monument d'une haute antiquite; et; par le fait de sa permanence; 
un objet digne d'occuper le geologue; et capable de lui fournir des 
remarques interessantes。〃  Although the superficial layer of 
vegetable mould as a whole no doubt is of the highest antiquity; 
yet in regard to its permanence; we shall hereafter see reason to 
believe that its component particles are in most cases removed at 
not a very slow rate; and are replaced by others due to the 
disintegration of the underlying materials。

As I was led to keep in my study during many months worms in pots 
filled with earth; I became interested in them; and wished to learn 
how far they acted consciously; and how much mental power they 
displayed。  I was the more desirous to learn something on this 
head; as few observations of this kind have been made; as far as I 
know; on animals so low in the scale of organization and so poorly 
provided with sense…organs; as are earth…worms。

In the year 1837; a short paper was read by me before the 
Geological Society of London; {2} 〃On the Formation of Mould;〃 in 
which it was shown that small fragments of burnt marl; cinders; 
&c。; which had been thickly strewed over the surface of several 
meadows; were found after a few years lying at the depth of some 
inches beneath the turf; but still forming a layer。  This apparent 
sinking of superficial bodies is due; as was first suggested to me 
by Mr。 Wedgwood of Maer Hall in Staffordshire; to the large 
quantity of fine earth continually brought up to the surface by 
worms in the form of castings。  These castings are sooner or later 
spread out and cover up any object left on the surface。  I was thus 
led to conclude that all the vegetable mould over the whole country 
has passed many times through; and will again pass many times 
through; the intestinal canals of worms。  Hence the term 〃animal 
mould〃 would be in some respects more appropriate than that 
commonly used of 〃vegetable mould。〃

Ten years after the publication of my paper; M。 D'Archiac; 
evidently influenced by the doctrines of Elie de Beaumont; wrote 
about my 〃singuliere theorie;〃 and objected that it could apply 
only to 〃les prairies basses et humides;〃 and that 〃les terres 
labourees; les bois; les prairies elevees; n'apportent aucune 
preuve a l'appui de cette maniere de voir。〃 {3}  But M。 D'Archiac 
must have thus argued from inner consciousness and not from 
observation; for worms abound to an extraordinary degree in kitchen 
gardens where the soil is continually worked; though in such loose 
soil they generally deposit their castings in any open cavities or 
within their old burrows instead of on the surface。  Hensen 
estimates that there are about twice as many worms in gardens as in 
corn…fields。 {4}  With respect to 〃prairies elevees;〃 I do not know 
how it may be in France; but nowhere in England have I seen the 
ground so thickly covered with castings as on commons; at a height 
of several hundred feet above the sea。  In woods again; if the 
loose leaves in autumn are removed; the whole surface will be found 
strewed with castings。  Dr。 King; the superintendent of the Botanic 
Garden in Calcutta; to whose kindness I am indebted for many 
observations on earth…worms; informs me that he found; near Nancy 
in France; the bottom of the State forests covered over many acres 
with a spongy layer; composed of dead leaves and innumerable worm…
castings。  He there heard the Professor of 〃Amenagement des Forets〃 
lecturing to his pupils; and pointing out this case as a 〃beautiful 
example of the natural cultivation of the soil; for year after year 
the thrown…up castings cover the dead leaves; the result being a 
rich humus of great thickness。〃

In the year 1869; Mr。 Fish {5} rejected my conclusions with respect 
to the part which worms have played in the formation of vegetable 
mould; merely on account of their assumed incapacity to do so much 
work。  He remarks that 〃considering their weakness and their size; 
the work they are represented to have accomplished is stupendous。〃  
Here we have an instance of that inability to sum up the effects of 
a continually recurrent cause; which has often retarded the 
progress of science; as formerly in the case of geology; and more 
recently in that of the principle of evolution。

Although these several objections seemed to me to have no weight; 
yet I resolved to make more observations of the same kind as those 
published; and to attack the problem on another side; namely; to 
weigh all the castings thrown up within a given time in a measured 
space; instead of ascertaining the rate at which objects left on 
the surface were buried by worms。  But some of my observations have 
been rendered almost superfluous by an admirable paper by Hensen; 
already alluded to; which appeared in 1877。 {6}  Before entering on 
details with respect to the castings; it will be advisable to give 
some account of the habits of worms from my own observations and 
from those of other naturalists。

'FIRST EDITION; October 10th; 1881。'



CHAPTER IHABITS OF WORMS。



Nature of the sites inhabitedCan live long under water
NocturnalWander about at nightOften lie close to the mouths of 
their burrows; and are thus destroyed in large numbers by birds
StructureDo not possess eyes; but can distinguish between light 
and darknessRetreat rapidly when brightly illuminated; not by a 
reflex actionPower of attentionSensitive to heat and cold
Completely deafSensitive to vibrations and to touchFeeble power 
of smellTasteMental qualitiesNature of foodOmnivorous
DigestionLeaves before being swallowed; moistened with a fluid of 
the nature of the pancreatic secretionExtra…stomachal digestion
Calciferous glands; structure ofCalcareous concretions formed in 
the anterior pair of glandsThe 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; {7} 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。 {8}

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 t

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