Earthworms and Their Allies

CHAPTER I. Structural and Systematic

STRUCTURAL AND SYSTEMATIC

The group of segmented, bristle-bearing, worms, termed Oligochaeta by zoologists, comprises what are popularly known as earthworms together with certain forms, inhabitants of ponds, lakes and rivers, which are not so well known as to have received a more distinctive name than merely 'worms.' Their next allies are apparently the leeches and--a little more remote--marine bristle-bearing worms termed Polychaeta; the three groups, together with perhaps a certain number of other forms belonging to smaller groups, constitute the Annelida which are a distinct and separate assemblage of invertebrate animals.

The most interesting features about these Oligochaetous worms are their very great anatomical variation and the facts of their distribution over the globe. Their importance as geological agents in levelling the ground was made known a long time ago by Darwin, and that aspect of earthworms has remained in much the same position as Darwin left it. We shall concern ourselves here only with the structure, habits, and range of the earthworms and their immediate allies, the aquatic Oligochaeta. These three aspects of the animals dovetail into each other more thoroughly than is the case with some other groups. This is due to the fact that they have of late years been very thoroughly studied from the anatomical and distributional side. So lately as 1889, M. Vaillant in a very comprehensive treatise was only able to enumerate 369 species, of which a large number were but incompletely differentiated, and some are no longer admitted. There are at the moment of writing perhaps 1500 species, the vast majority of which are well known owing to careful investigation. Furthermore there are but few parts of the world, and these are not of large area, from which earthworms at any rate have not been gathered. Though there can be no doubt that a very considerable number of species await discovery, it would seem that we are in possession of information which is not likely to be seriously affected by future researches.

THE ANATOMY OF EARTHWORMS.

Although it is not contemplated to make the present volume a guide to the structure of this group of worms, it is necessary to give some little anatomical sketch of the group in order first of all to illustrate their diversity of structure, secondly to give reasons for the classification of them, and thirdly to enable the reader to realise certain structural details which it is absolutely necessary to give some account of in order to explain other matters.

It is for example impossible to attempt any account of the fitness of some of these animals for their terrestrial life and of others for an aquatic life without treating of anatomy to some extent.

I shall take one particular species as a type and indicate later the principal divergencies shown by other forms. According to the general opinion among those who have studied the Oligochaeta I take as a representative form a Megascolecid (this and the other families are dealt with seriatim on p. 14 et seq.), as this group is presumed to be the oldest, and within that group a representative of the genus Notiodrilus which is with some reason held to be the most primitive genus in the group. Finally I have no particular reason for selecting the species Notiodrilus tamajusi except that there happens to be a longer and fuller description of it than of many.

This earthworm is a native of Guatemala and is some six inches in length with a diameter of perhaps a quarter of an inch. The front part of the body is thicker than posteriorly. The body will be seen to be divided into some 218 rings by circular furrows which run right round the body. These divisions are termed segments or somites. At the head the mouth is surrounded by the first of these, and on the dorsal surface of that segment is a projection like an incomplete segment which is known as the prostomium. From the XIIIth segment to the middle of the XXth the body has a different appearance, and this region is known as the clitellum. Each of the 218 segments of the body except the first, and possibly one or two at the hinder end, is furnished with eight minute projecting bristles, the setae; these are disposed in pairs and all lie upon the ventral aspect of the worm. The movement of these by special muscles aids in locomotion.

An examination with even a hand lens shows a number of external pores which are important. Anteriorly there is the mouth which is overhung by the prostomium referred to above. At the extreme hind end--and surrounded by the last segment of the body--is the vent. Along the middle line of the back are a series of pores, one just at the very anterior edge of each segment, through which, when the worm is dried and then slightly pressed, liquid is seen to be ejected. These are called the dorsal pores and they belong one to each segment with the exception of the first seven, or--in some cases--more, segments. In front of one or other of the pair of setae which is situated most laterally, i.e. furthest from the ventral median line, is an orifice on each side in all but the first one or two segments of the body. These paired pores are the external outlets of the excretory organs frequently termed on account of their regular repetition with the segments 'segmental organs,' but more conveniently to be named nephridia. In the clitellar region and in fact on each of the segments XVII, XVIII, XIX are a pair of pores of which those on the XVIIIth segment are the least conspicuous. The large pair of pores on each of segments XVII and XIX occupy the position of the ventralmost pair of setae, which are here absent, or rather replaced by a very long curved and ornamented seta, which projects out of the orifice. These two pairs of pores are the outlets of the prostatic glands as they have been termed. The minute pair of pores on segment XVIII do not take up the position of the ventral setae; for these are present and to the inside of each pore. A groove, shaped something like a reversed 3 or the Greek letter [Greek: S], connects the orifices of each side of the body, the middle part of the groove, where the two semicircular halves of which it is composed meet, coinciding with the minute pores on segment XVIII which are the orifices of the sperm ducts.

On segment XIV are a pair of very minute pores a little in front of the ventralmost setae and thus very near together. These are the openings of the oviducts. Finally, near to the anterior border-line of segments VIII and IX and on a line with the ventral pair of setae is a pore on each side through which the cavity of the spermathecae reaches the exterior.

So much then for the external characters of our worm. We next turn to the internal anatomy. When the worm is opened by a longitudinal section from end to end, and the two flaps of skin are turned outwards and pinned down, the internal structure is almost completely revealed. Running from end to end is seen the alimentary canal; the general cavity of the body (coelom) in which it lies, as do of course the other organs to be enumerated, is seen to be divided by cross divisions, the intersegmental septa, into a series of chambers which correspond with the external division into segments. The septa are in fact inserted on to the body-wall along the furrows which mark the divisions between adjacent segments. Anteriorly the large pharynx is responsible for confusing the arrangement of the septa, which become subdivided and fused or are prolonged a greater way backwards and thus present a less obviously segmental disposition. Certain of the more anteriorly placed of these septa are much thicker than the rest. This is the case with the septa which separate segments V to XII. The alimentary canal is perfectly straight and runs in the middle line, being supported by the septa which it perforates. The mouth leads into a buccal cavity which later becomes the pharynx, a portion of the tube which is much thickened by muscular walls dorsally. Then follows a very short section of the oesophagus and in the fifth segment this becomes the gizzard, a very characteristic organ with thick muscular walls quite smooth and with a very thick lining of structureless membrane. After this is a narrower tube, the rest of the oesophagus. Into this open in each of segments VII, VIII, IX a pair of calciferous glands; these are diverticula of the gut with much folded walls, the cells of which secrete carbonate of lime. In the XIIth segment or so, the oesophagus suddenly widens out to form the intestine which runs as such to the end of the body. This wider tube has a ridge running along its dorsal side, the typhlosole. Along the dorsal surface of the intestine and the oesophagus is seen a red tube, contractile during the life of the worm, which is the dorsal blood vessel and whose contained blood is coloured red, as is the blood of vertebrated animals, by haemoglobin. But in the earthworm the colouring matter is not situated in corpuscles as in the vertebrate. The dorsal vessel is connected by a few pairs of equally contractile transverse trunks with a ventral vessel which is not contractile. There are other branches of these main longitudinal trunks and some minor longitudinal trunks which we shall not stop to describe further. The nervous system of the worm consists of a pair of ganglia which lie above the gut in the third segment; they are connected by a commissure running round the gut with a chain of closely fused pairs of ganglia, one for each segment to the very end of the body. In each of the segments, except the first two or three, there are a pair of excretory organs known as nephridia; these are essentially coiled glandular tubes opening on to the exterior by the regularly placed pores already referred to in considering the external characters. The tube ends in a funnel-shaped, and therefore dilated, mouth, which opens into the segment in front of that which contains the rest of the organ; a nephridium therefore lies in two segments. The only other important organs which are left for consideration are those devoted to the reproduction of the species. The essential organs are the spermaries and the ovaries. Of the former there are two pairs of minute whitish bodies which lie in segments X and XI on either side of the nerve cord attached to the anterior septal wall of their segments. The ovaries are not in the following, but in the XIIIth, segment, and occupy an identical position in that segment. A short tube with a funnel or trumpet-shaped and wide orifice opens into the cavity of the XIIIth segment opposite to each, and, perforating the septum, opens on to the exterior on the XIVth segment. A similar but larger and more folded pair of trumpet-shaped funnels opens in the same way opposite to each spermary. But in this case the two tubes of the sperm ducts run backwards for some way and those of each side after joining open on to the XVIIIth segment by the pores already mentioned. On the XVIIth and XIXth segments open two glands which are long and tubular in form and much coiled. These are the spermiducal glands and each opens in common with a muscular sac containing the long and ornamented seta referred to in describing the various external orifices. It will be noticed that the sperm duct has no direct connection with these glands but only indirectly through the external gutter which connects the three male orifices of each side of the body. Segments IX-XII inclusive contain certain sacs which depend from, and are formed as outgrowths of, the septa of those segments. These are the sperm sacs in which the male germ cells undergo their development. A corresponding body (but very much smaller) is sometimes found in relation to the ovary but has not been actually described in the particular species dealt with here. Finally, in segments VIII and IX are a pair (that is four altogether) of roundish sacs, with two or three minute diverticula, known as the spermathecae. In the diverticula of these sacs are stored the sperm derived from another individual.

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