The Works of Francis Balfour 1-25

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Foster M. and Sedgwick A. The Works of Francis Balfour Vol. I. Separate Memoirs (1885) MacMillan and Co., London.

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This historic 1885 book edited by Foster and Sedgwick is the first of Francis Balfour's collected works published in four editions. Francis (Frank) Maitland Balfour, known as F. M. Balfour, (November 10, 1851 - July 19, 1882) was a British biologist who co-authored embryology textbooks.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. I. Separate Memoirs (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. II. A Treatise on Comparative Embryology 1. (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. III. A Treatise on Comparative Embryology 2 (1885) MacMillan and Co., London.

Foster M. and Sedgwick A. The Works of Francis Balfour Vol. IV. Plates (1885) MacMillan and Co., London.
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Pages where the terms "Historic Textbook" and "Historic Embryology" appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms and interpretations may not reflect our current scientific understanding.     (More? Embryology History | Historic Embryology Papers)

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Vol I. Separate Memoirs (1885)

XXV. The Anatomy and Development of Peripatus Capensis

(With Plates 4653.)


THE late Professor Balfour was engaged just before his death in investigating the structure and embryology of Peripatus capensis, with the view of publishing a complete monograph of the genus. He left numerous drawings intended to serve as illustrations to the monograph, together with a series of notes and descriptions of a large part of the anatomy of Peripatus capensis. Of this manuscript some portions were ready for publication, others were more or less imperfect ; while of the figures many were without references, and others were provided with only a few words of explanation.

It was obviously necessary that Professor Balfour's work embodying as it did much important discovery should be published without delay; and the task of preparing his material for the press was confided to us. We have printed all his notes and descriptions without alteration 2 . Explanations which appeared to be necessary, and additions to the text in cases in which he had prepared figures without writing descriptions, together with full descriptions of all the plates, have been added by us, and are distinguished by enclosure in square brackets 3 .

We have to thank Miss Balfour, Professor Balfour's sister, for the important service which she has rendered by preparing

1 From the Quarterly Jotirnal of Microscopical Science, April, 1883.

2 Excepting in an unimportant matter of change of nomenclature used with regard to the buccal cavity.

3 The account of the external characters, generative organs, and development, has been written by the editors.


a large part of the beautiful drawings with which the monograph is illustrated. Many of these had been executed by her under Professor Balfour's personal supervision ; and the knowledge of his work which she then acquired has been of the greatest assistance to us in preparing the MSS. and drawings for publication.

Since his death she has spared no pains in studying the structure of Peripatus, so as to enable us to bring out the first part of the monograph in as complete a state as possible. It is due to her skill' that the first really serviceable and accurate representation of the legs of any species of Peripatus available for scientific purposes are issued with the present memoir 1 .

We have purposely refrained from introducing comments on the general bearing of the new and important results set forth in this memoir, and have confined ourselves to what was strictly necessary for the presentation of Mr Balfour's discoveries in a form in which they could be fully comprehended.

Mr Balfour had at his disposal numerous specimens of Peripatus nova zealandia, collected for him by Professor Jeffrey Parker, of Christchurch, New Zealand ; also specimens from the Cape of Good Hope collected by Mr Lloyd Morgan, and brought to England by Mr Roland Trimen in 1881 ; and others given to him by Mr Wood Mason, together with all the material collected by Mr Moseley during the "Challenger" voyage.

A preliminary account of the discoveries as to the embryology of Peripatus has already been communicated to the Royal Society 2 . It is intended that the present memoir shall be followed by others, comprising a complete account of all the species of the genus Peripatus.


1 The drawings on PI. 47, figs. 9 and 10 on PI. 48, and the drawings of the embryos (except fig. 37), have been made by Miss Balfour since Professor Balfour's death.

5 Proc. Royal Soc. 1883.



Peripatus capensis (fig. i).

[The body is elongated, and slightly flattened dorso-ventrally. The dorsal surface is arched, and darkly pigmented ; while the ventral surface is nearly flat, and of a lighter colour.

The mouth is placed at the anterior end of the body, on the ventral surface.

The anus is posterior and terminal.

The generative opening is single and median, and placed in both sexes on the ventral surface, immediately in front of the anus.

There are a pair of ringed antennae projecting from the anterior end of the head, and a pair of simple eyes, placed on the dorsal surface at the roots of the antennae.

The appendages of the body behind the antennae are disposed in twenty pairs.

1. The single pair of jaws placed within the buccal cavity in front of the true mouth opening, and consisting each of a papilla, armed at its termination with two cutting blades.

2. The oral papillae placed on each side of the mouth. At their apices the ducts of the slime glands open.

3. The seventeen pairs of ambulatory appendages, each provided with a pair of chitinous claws at its extremity.

4. The anal papillae placed on each side of the generative opening.

Colour. The following statements on this head are derived from observations of spirit specimens. The colour varies in different individuals. It always consists of a groundwork of green and bluish grey, with a greater or less admixture of brown. The chief variations in the appearance of the animal, so far as colour is concerned, depend on the shade of the green. In some it is dark, as in the specimen figured (fig. i) ; in others it is of a lighter shade.

There is present in most specimens a fairly broad light band on each side of the body, immediately dorsal to the attachment B. 56


of the legs. This band is more prominent in the lighter coloured vaiieties than in the dark, and is especially conspicuous in large individuals. It is due to a diminution in the green pigment, and an increase in the brown.

There is a dark line running down the middle of the dorsal surface, in the middle of which is a fine whitish line.

The ventral surface is almost entirely free from the green pigment, but possesses a certain amount of light brown. This brown pigment is more conspicuous and of a darker shade on the spinous pads of the foot.

In parts of the body where the pigment is scarce, it is seen to be confined to the papillae. This is especially evident round the mouth, where the sparse green pigment is entirely confined to the papillae.

In some specimens a number of white papillae, or perhaps light brown, are scattered over the dorsal surface ; and sometimes there is a scattering of green papillae all over the ventral surface. These two peculiarities are more especially noticeable in small specimens.

Ridges and Papilla of the Skin. The skin is thrown into a number of transverse ridges, along which the primary wartlike papillae are placed.

The papillae, which are found everywhere, are specially developed on the dorsal surface, less so on the ventral. The papillae round the lips differ from the remaining papillae of the ventral surface in containing a green pigment. Each papilla bears at its extremity a well-marked spine.

The ridges of the skin are not continued across the dorsal middle line, being interrupted by the whitish line already mentioned. Those which lie in the same transverse line as the legs are not continued on to the latter, but stop at the junction of the latter with the body. All the others pass round to the ventral surface and are continued across the middle line ; they do not, however, become continuous with the ridges of the other side, but passing between them gradually thin off and vanish.

The ridges on the legs are directed transversely to their long axes, i.e. are at right angles to the ridges of the rest of the body.


The -antennae are ringed and taper slightly till near their termination, where they present a slight enlargement in spirit specimens, which in its turn tapers to its termination.

The rings consist essentially of a number of coalesced primary papillae, and are, therefore, beset by a number of spines like those of the primary papillae (described below). They are more deeply pigmented than the rest of the antenna.

The free end of the antenna is covered by a cap of tissue like that of the rings. It is followed by four or more rings placed close together on the terminal enlargement. There appears to be about thirty rings on the antennae of all adults of this species. But they are difficult to count, and a number of small rings occur between them, which are not included in the thirty.

The antennae are prolongations of the dorso-lateral parts of the anterior end of the body.

The eyes are paired and are situated at the roots of the antennae on the dorso-lateral parts of the head. Each is placed on the side of a protuberance which is continued as the antenna, and presents the appearance of a small circular crystalline ball inserted on the skin in this region.

The rings of papillae on that part of the head from which the antennae arise lose their transverse arrangement. They are arranged concentrically to the antennal rings, and have a straight course forwards between the antennae.

The oral papillae are placed at the side of the head. They are attached ventro-Iaterally on each side of the lips. The duct of the slime gland opens through their free end. They possess two main rings of projecting tissue, which are especially pigmented on the dorsal side ; and their extremities are covered by papillae irregularly arranged.

The buccal cavity, jaws, and lips are described below.

The Ambulatory Appendages. The claw-bearing legs are usually seventeen in number ; but in two cases of small females we have observed that the anal papillae bear claws, and present all the essential features of the ambulatory appendages. In one small female specimen there were twenty pairs of clawbearing appendages, the last being like the claw-bearing anal papillae last mentioned, and the generative opening being placed between them.

56 2


The ambulatory appendages, with the exception of the fourth and fifth pairs in both sexes, and the last pair (seventeenth) in the male, all resemble each other fairly closely. A typical appendage (figs. 2 and 3) will first be described, and the small variations found in the appendages just mentioned will then be pointed out. Each consists of two main divisions, a larger proximal portion, the leg, and a narrow distal claw-bearing portion, the foot.

The leg has the form of a truncated cone, the broad end of which is attached to the ventro-lateral body-wall, of which it appears to be, and is, a prolongation. It is marked by a number of rings of primary papillae, placed transversely to the long axis of the leg, the dorsal of which contain a green and the ventral a brown pigment. These rings of papillae, at the attachment of the leg, gradually change their direction and merge into the body rings. At the narrow end of the cone there are three ventrally placed pads, in which the brown pigment is dark, and which are covered by a number of spines precisely resembling the spines of the primary papillae. These spinous pads are continued dorsally, each into a ring of papillae.

The papillae of the ventral row next the proximal of these spinous pads are intermediate in character between the primary papillae and the spinous pads. Each of these papillae is larger than a normal papilla, and bears several spines (fig. 2). This character of the papilla of this row is even more marked in some of the anterior legs than in the one figured ; it seems probable that the pads have been formed by the coalescence of several rows of papillae on the ventral surface of the legs. On the outer and inner sides of these pads the spines are absent, and secondary papillae only are present.

In the centre of the basal part of the ventral surface of the foot there are present a group of larger papillae, which are of a slightly paler colour than the others. They are arranged so as to form a groove, directed transversely to the long axis of the body, and separated at its internal extremity by a median papilla from a deep pit which is placed at the point of junction of the body and leg. The whole structure has the appearance, when viewed with the naked eye, of a transverse slit placed at the base of the leg. The segmental organs open by the deep pit placed


at the internal end of this structure. The exact arrangement of the papillae round the outer part of the slit does not appear to be constant.

The foot is attached to the distal end of the leg. It is slightly narrower at its attached extremity than at its free end, which bears the two claws. The integument of the foot is covered with secondary papillae, but spines and primary papillae are absent, except at the points now to be described.

On each side of the middle ventral line of the proximal end of the foot is placed an elliptical elevation of the integument covered with spines. Attached to the proximal and lateral end of this is a primary papilla. At the distal end of the ventral side of the foot on each side of the middle line is a group of inconspicuous pale elevations, bearing spines.

On the front side of the distal end of the foot, close to the socket in which the claws are placed, are two primary papillae, one dorsal and the other ventral.

On the posterior side of the foot the dorsal of these only is present. The claws are sickle-shaped, and placed on papillae on the terminal portion of the foot. The part of the foot on which they are placed is especially retractile, and is generally found more or less telescoped into the proximal part (as in the figure).

The fourth and fifth pairs of legs exactly resemble the others, except in the fact that the proximal pad is broken up into three, a small central and two larger lateral. The enlarged segmental organs of these legs open on the small central division.

The last (17) leg of the male (PL 47, fig. 4) is characterized by possessing a well-marked white papilla on the ventral surface. This papilla, which presents a slit-like opening at its apex, is placed on the second row of papillae counting from the innermost pad, and slightly posterior to the axial line of the leg.

The anal papillae, or as they should be called, generative papillae, are placed one on each side of the generative aperture. They are most marked in small and least so in large specimens. That they are rudimentary ambulatory appendages is shewn by the fact that they are sometimes provided with claws, and resemble closely the anterior appendages.]



The alimentary canal of Peripatus capensis forms, in the extended condition of the animal, a nearly straight tube, slightly longer than the body, the general characters of which are shewn in figs. 6 and 7.

For the purposes of description, it may conveniently be divided into five regions, viz. (i) the buccal cavity with the tongue, jaws, and salivary glands, (2) pharynx, (3) the oesophagus, (4) the stomach, (5) the rectum.

The Buccal Cavity. The buccal cavity has the form of a fairly deep pit, of a longitudinal oval form, placed on the ventral surface of the head, and surrounded by a tumid lip.

[The buccal cavity has been shewn by Moseley to be formed in the embryo by the fusion of a series of processes surrounding the true mouth-opening, and enclosing in their fusion the jaws.]

The 'lip is covered by a soft skin, in which are numerous organs of touch, similar to those in other parts of the skin having their projecting portions enclosed in delicate spines formed by the cuticle. The skin of the lips differs, however, from the remainder of the skin, in the absence of tubercles, and in the great reduction of the thickness of the dermis. It is raised into a series of papilliform ridges, whose general form is shewn in fig. 5 ; of these there is one unpaired and median behind, and a pair, differing somewhat in character from the remainder, in front, and there are, in addition, seven on each side.

The structures within the buccal cavity are shewn as they appear in surface views in figs. 5 and 7, but their real nature is best seen in sections, and is illustrated by PL 49, figs. II and 12, representing the oral cavity in transverse section, and by PL 49, figs. 17 and 1 8, representing it in horizontal longitudinal sections. In the median line of the buccal cavity in front is placed a thick muscular protuberance, which may perhaps conveniently be called the tongue, though attached to the dorsal instead of the ventral wall of the mouth. It has the form of an elongated


ridge, which ends rather abruptly behind, becoming continuous with the dorsal wall of the pharynx. Its projecting edge is armed by a series of small teeth, which are thickenings of the chitinous covering, prolonged from the surface of the body over the buccal cavity. Where the ridge becomes flatter behind, the row of teeth divides into two, with a shallow groove between them (PI. 48, fig. 7).

The surface of the tongue is covered by the oral epithelium, in parts of which are organs of special sense, similar to those in the skin; but its interior is wholly formed of powerful muscles. The muscles form two groups, intermingled amongst each other. There are a series of fibres inserted in the free edge of the tongue, which diverge, more or less obliquely, towards the skin at the front of the head anteriorly, and towards the pharynx behind. The latter set of fibres are directly continuous with the radial fibres of the pharynx. The muscular fibres just described are clearly adapted to give a sawing motion to the tongue, whose movements may thus, to a certain extent, be compared to those of the odontophor of a mollusc.

In addition to the above set of muscles, there are also transverse muscles, forming laminae between the fibres just described. They pass from side to side across the tongue, and their action is clearly to narrow it, and so cause it to project outwards from the buccal cavity.

On each side of the tongue are placed the jaws, which are, no doubt, a pair of appendages, modified in the characteristic arthropodan manner, to subserve mastication. Their structure has never been satisfactorily described, and is very complicated. They are essentially short papillae, moved by an elaborate and powerful system of muscles, and armed at their free extremities by a pair of cutting blades or claws. The latter structures are, in all essential points, similar to the claws borne by the feet, and, like these, are formed as thickenings of the cuticle. They have therefore essentially the characters of the claws and jaws of the Arthropoda, and are wholly dissimilar to the setae of Chsetopoda. The claws are sickle-shaped and, as shewn in PL 47, fig. 5, have their convex edge directed nearly straight forwards, and their concave or cutting edge pointed backwards. Their form differs somewhat in the different species, and, as will


be shewn in the systematic part of this memoir 1 , forms a good specific character. In Peripatus capensis (PI. 48, fig. 10) the cutting surface of the outer blade is smooth and without teeth, while that of the inner blade (fig. 9), which is the larger of the two, is provided with five or six small teeth, in addition to the main point. A more important difference between the two blades than that in the character of the cutting edge just spoken of, is to be found in their relation to the muscles which move them. The anterior parts of both blades are placed on two epithelial ridges, which are moved by muscles common to both blades (PI. 49, fig. 1 1). Posteriorly, however, the behaviour of the two blades is very different. The epithelial ridge bearing the outer blade is continued back for a short distance behind the blade, but the cuticle covering it becomes very thin, and it forms a simple epithelial ridge placed parallel to the inner blade. The cuticle covering the epithelial ridge of the inner blade is, on the contrary, prolonged behind the blade itself as a thick rod, which, penetrating backwards along a deep pocket of the buccal epithelium, behind the main part of the buccal cavity for the whole length of the pharynx, forms a very powerful lever, on which a great part of the muscles connected with the jaws find their insertion. The relations of the epithelial pocket bearing this lever are somewhat peculiar.

The part of the epithelial ridge bearing the proximal part of this lever is bounded on both its outer and inner aspect by a deep groove. The wall of the outer groove is formed by the epithelial ridge of the outer blade, and that of the inner by a special epithelial ridge at the side of the tongue. Close to the hinder border of the buccal cavity (as shewn in PL 49, fig. 12, on the right hand side), the outer walls of these two grooves meet over the lever, so as completely to enclose it in an epithelial tube, and almost immediately behind this point the epithelial tube is detached from the oral epithelium, and appears in section as a tube with a chitinous rod in its interior, lying freely in the body-cavity (shewn in PI. 49, figs. 13 16 le). This apparent tube is the section of the deep pit already spoken of. It may

1 Some material for this memoir was left by Prof. Balfour, which will be published separately.


be traced back even beyond the end of the pharynx, and serves along its whole length for the attachment of muscles.

The greater part of the buccal cavity is filled with the tongue and jaws just described. It opens dorsally and behind by the mouth into the pharynx, there being no sharp line of demarcation between the buccal cavity and the pharynx. Behind the opening into the pharynx there is a continuation of the buccal cavity shewn in transverse section in fig. 13, and in longitudinal and horizontal section in fig. 17, into which there opens the common junction of the two salivary glands. This diverticulum is wide at first and opens by a somewhat constricted mouth into the pharynx above (PL 49, fig. 13, also shewn in longitudinal and horizontal section in fig. 17). Behind it narrows, passing insensibly into what may most conveniently be regarded as a common duct for the two salivary glands (PL 49, fig. 17).

The Salivary Glands, These two bodies were originally described by Grube, by whom their nature was not made out, and subsequently by Moseley, who regarded them as fat bodies. They are placed in the lateral compartments of the body-cavity immediately dorsal to the ventral nerve cords, and extend for a very variable distance, sometimes not more than half the length of the body, and in other instances extending for nearly its whole length. Their average length is perhaps about twothirds that of the body. Their middle portion is thickest, and they thin off very much behind and to a slight extent in front. Immediately behind the mouth and in front of the first pair of legs, they bend inwards and downwards, and fall (fig. 7) one on each side into the hind end of the narrow section of the oral diverticulum just spoken of as the common duct for the two salivary glands. The glandular part of these organs is that extending back from the point where they bend inwards. This part (fig. 1 6) is formed of very elongated cells supported by a delicate membrana propria. The section of this part is somewhat triangular, and the cells are so long as to leave a comparatively small lumen. The nuclei of the cells are placed close to the supporting membrane, and the remainder of the cells arc filled with very closely packed secretory globules, which have a high index of refraction. It was the presence of these globules which probably led Moseley to regard the salivary glands as fat


bodies. The part of each gland which bends inwards must be regarded as the duct.

The cells lining the ducts are considerably less columnar than those of the gland proper. Their nuclei (fig. 14) are situated at the free extremities instead of at the base of the cells, and they are without secretory globules. The cells lining the ducts of the salivary glands pass, without any sharp line of demarcation, into those of the oral epithelium, which are flatter and have their nuclei placed in the middle.

The Pharynx. The Pharynx is a highly muscular tube (fig. 7) with a triangular lumen (figs. 14, 15), .which extends from the mouth to about half way between the first and second pair of legs. It is lined by a flattish epithelium bounded by a cuticle continuous with that of the mouth. On the dorsal side is a ridge projecting into the lumen of the pharynx. This ridge may be traced forwards (PI. 49, figs. II 14) into the tongue, and the two grooves at the side of this ridge, forming the two upper angles of the triangular lumen, may be followed into those at the sides of the tongue. The muscles of the pharynx are very highly developed, consisting of an intrinsic and an extrinsic set. The former consists, as is best seen in longitudinal sections, of (PI. 51, fig. 23) radial fibres, arranged in somewhat wedgeshaped laminae, between which are rings of circular fibres. The latter are thicker externally than internally, and so also appear wedge-shaped in longitudinal sections. Very characteristic of the pharynx are the two sympathetic nerves placed close to the two dorsal angles of the triangular lumen (fig. 14, sy).

The pharynx of Peripatus is interesting in that it is unlike, so far as I know, the pharynx of any true Arthropod, in all of which the region corresponding with the pharynx of Peripatus is provided with relatively very thin walls.

The pharynx of Peripatus has, on the other hand, a very close and obvious resemblance to that of many of the Chaetopoda, a resemblance which is greatly increased by the characteristic course of the sympathetic nerves.

The form of the lumen, as already pointed out by Grube, resembles that of the Nematoda.

Ttte (Esophagus. Behind the pharynx there follows a narrow oesophagus (fig. 7, o e] shewn in section in fig. 16. It has some


what folded and fairly thick walls, and lies freely in the central division of the body-cavity without any mesenteric support. Its walls are formed of five layers, viz. from without inwards.

(1) A peritoneal investment.

(2) A layer of longitudinal fibres.

(3) A layer of circular fibres, amongst which are numerous nuclei.

(4) A connective-tissue layer supporting (5) a layer of fairly columnar hyaline epithelium, bounded on its inner aspect by a cuticle continued from that of the pharynx. In front it passes insensibly into the pharynx, and beyond the region where the dorsal walls of the pharynx have clearly commenced, the ventral walls still retain the characters of the cesophageal walls. The oesophagus is vertically oval in front, but more nearly circular behind. Characteristic of the cesophagus is the junction of the two sympathetic nerves on its dorsal wall (fig. 16). These nerves cannot be traced far beyond their point of junction.

The Stomach. The next section of -the alimentary tract is the stomach or rnesenteron (fig. 6). It is by far the largest part of the alimentary tract, commencing at about the second pair of legs and extending nearly to the hind end of the body. It tapers both in front and behind, and is narrowest in the middle, and is marked off sharply both from the cesophagus in front and the rectum behind, and is distinguished from both of these by its somewhat pinker hue. In the retracted condition of the animal it is, as pointed out by Moseley, folded in a single short dorsal loop, at about the junction of its first with its second third, and also, according to my observations, at its junction with the rectum ; but in the extended condition it is nearly straight, though usually the posterior fold at the junction of the rectum is not completely removed. Its walls are always marked by plications which, as both Moseley and Grube have stated, do not in any way correspond with the segmentation of the body. In its interior I have frequently found the chitinous remains of the skins of insects, so that we are not justified in considering that the diet is purely vegetable. It lies free, and is, like the remainder of the alimentary tract, without a mesentery. The structure of the walls of the stomach has not hitherto been very satisfactorily described.


The connective tissue and muscular coats are extremely thin. There is present everywhere a peritoneal covering, and in front a fairly well-marked though very thin layer of muscles formed of an external circular and an internal longitudinal layer. In the middle and posterior parts, however, I was unable to recognize these two layers in section ; although in surface view Grube found an inner layer of circular fibres and an outer layer formed of bands of longitudinal fibres, which he regards as muscular.

The layer supporting the epithelium is reduced to a basement membrane. The epithelial part of the wall of the stomach is by far the thickest (fig. 20), and is mainly composed of enormously elongated, fibre-like cells, which in the middle part of the stomach, where they are longest, are nearly half a millimetre in length, and only about '006 mm. in breadth. Their nuclei, as seen in fig. 20, are very elongated, and are placed about a quarter of the length from the base.

The cells are mainly filled with an immense number of highly refracting spherules, probably secretory globules, but held by Grube, from the fact of their dissolving in ether, to be fat. The epithelial cells are raised into numerous blunt processes projecting into the lumen of the stomach.

In addition to the cells just described there are present in the anterior part of the stomach a fair sprinkling of mucous cells. There are also everywhere present around the bases of the columnar cells short cells with spherical nuclei, which are somewhat irregularly scattered in the middle and posterior parts of the stomach, but form in the front part a definite layer. I have not been able to isolate these cells, and can give no account of their function.

The rectum extends from the end of the stomach to the anus. The region of junction between the stomach and the rectum is somewhat folded. The usual arrangement of the parts is shewn in fig. 6, where the hind end of the stomach is seen to be bent upon itself in a U-shaped fashion, and the rectum extending forwards under this bent portion and joining the front end of the dorsal limb of the U. The structure of the walls of the rectum is entirely different to that of the stomach, and the transition between the two is perfectly sudden.


Within the peritoneal investment comes a well-developed muscular layer with a somewhat unusual arrangement of its layers, there being an external circular layer and an internal layer formed of isolated longitudinal bands. The epithelium is fairly columnar, formed of granular cells with large nuclei, and is lined by a prolongation of the external cuticle. It is raised into numerous longitudinal folds, which are visible from the surface, and give a very characteristic appearance to this part of the alimentary tract. The muscular layers do not penetrate into the epithelial folds, which are supported by a connective tissue layer.


The central nervous system consists of a pair of supra-cesophageal ganglia united in the middle line, and of a pair of widely divaricated ventral cords, continuous in front with the supra-cesophageal ganglia.

It will be convenient in the first instance to deal with the general anatomy of the nervous system and then with the histology.

Ventral Cords. The ventral cords at first sight appear to be without ganglionic thickenings, but on more careful examination they are found to be enlarged at each pair of legs (PI. 48, fig. 8). These enlargements may be regarded as imperfect ganglia. There are, therefore, seventeen such pairs of ganglia corresponding to the seventeen pairs of legs. There is in addition a ganglionic enlargement at the commencement of the cesophageal commissures, where the nerves to the oral papillae are given off (PL 51, fig. 22 or. g.\ and the region of junction between the cesophageal commissures with the supra-cesophageal ganglia, where another pair of nerves are given off to the jaws (PI. 51, fig. 22/0), may be regarded as the anterior ganglion of the ventral cords. There are, therefore, according to the above reckoning, nineteen pairs of ganglia connected with the ventral cords.

The ventral cords are placed each in the lateral compartments of the body-cavity, immediately within the longitudinal layer of muscles.


They are connected with each other, rather like the pedal nerves of Chiton and the lower Prosobranchiata, by a number of commissures. These commissures exhibit a fairly regular arrangement from the region included between the first and the last pair of true feet. There are nine or ten of them between each pair of feet (PI. 52, fig. 26). They pass along the ventral wall of the body, perforating the ventral mass of longitudinal muscles. On their way they give off nerves which innervate the skin.

In Peripatus nova zealandicz, and probably also in P. capensis, two of these nerves, coming off from each pair of ganglia, are distinguished from the remainder by the fact that they are provided with numerous nerve-cells, instead of being composed of nerve-fibres only, like the remaining commissures (PL 52, fig. 26 g co). In correlation with the nerves given off from them to the skin the commissures are smaller in the middle than at the two ends.

Posteriorly the two nerve-cords nearly meet immediately in front of the generative aperture, and between this aperture and the last pair of feet there are about six commissures passing between them (PL 48, fig. 8). Behind the generative aperture the two cords bend upwards, and, as is shewn in fig. 8, fall into each other dorsally to the rectum. The section of the two cords placed dorsally to the rectum is solely formed of nerve-fibres; the nerve-cells, present elsewhere, being here absent.

In front of the ganglion of the first foot the commissures have a more dorsal situation than in the remainder of the body. The median longitudinal ventral muscle here gradually thins out and comes to an end, while the commissures pass immediately below the wall of the pharynx (PL 49, figs. 14, 15). The ventral cords themselves at first approach very close to each other in this region, separating again, however, to envelope between them the pharynx (PL 51, fig. 22).

There are eleven commissures in front of the first pair of legs (PL 51, fig. 22). The three foremost of these are very close together, the middle one arising in a more ventral position than the other two, and joining in the median ventral line a peculiar mass of cells placed in contact with the oral epithelium (fig. 14). It is probably an organ of special sense.


The ventral cords give off a series of nerves from their outer borders, which present throughout the trunk a fairly regular arrangement. From each ganglion two large nerves (figs. 8, 22, 26) are given off, which, diverging somewhat from each other, pass into the feet, and, giving off branches on their way, may be traced for a considerable distance within the feet along their anterior and posterior borders.

In front of each of the pair of pedal nerves a fairly large nerve may be seen passing outwards towards the side of the body (fig. 22). In addition to this nerve there are a number of smaller nerves passing off from the main trunk, which do not appear to be quite constant in number, but which are usually about seven or eight. Similar nerves to those behind are given off from the region in front of the first pair of legs, while at the point where the two ventral cords pass into the oesophageal commissures two large nerves (fig. 22), similar to the pairs of pedal nerves, take their origin. These nerves may be traced forwards into the oral papillae, and are therefore to be regarded as the nerves of these appendages. On the ventral side of the cords, where they approach most closely, between the oral papillae and the first pair of legs, a number of small nerves are given off to the skin, whose distribution appears to be to the same region of the skin as that of the branches from the commissures behind the first pair of legs.

From the cesophageal commissures, close to their junction with the supra-cesophageal ganglia, a nerve arises on each side which passes to the jaws, and a little in front of this, apparently from the supra-cesophageal ganglion itself, a second nerve to the jaws also takes its origin (PI. 51, fig. 22 j n}.. These two nerves I take to be homologous with a pair of pedal nerves.

Between the nerves to the jaws and those to the oral papillae a number of small nerves take their origin. Three of these on each side pass in a dorsal direction and one or two in a ventral one.

The Supra-cesophageal Ganglia. The supra-cesophageal ganglia (figs. 8 and 22) are large, somewhat oval masses, broader in front than behind, completely fused in the middle, but free at their extremities. Each of them is prolonged anteriorly into an antennary nerve, and is continuous behind with one of the


cesophageal commissures. On the ventral surface of each, rather behind the level of the eye, is placed a very peculiar protuberance (fig. 22 d], of which I shall say more in dealing with the histology of the nervous system.

A number of nerves arise from the supra-cesophageal ganglia, mainly from their dorsal surface.

In front are the immense antennary nerves extending along the whole length of each antenna, and giving off numerous lateral twigs to the sense organs. Near the origin of the antennary nerves, and rather on the dorsal surface, there spring a few small twigs, which pass to the skin, and are presumably sensory. The largest of them is shewn in PI. 50, fig. 19 A. About one-third of the way back the two large optic nerves take their origin, also arising laterally, but rather from the dorsal surface (PL 50, fig. 19 D and E). Each of them joins a large ganglionic mass placed immediately behind the retina. Nearly on a level with the optic nerves and slightly nearer the middle dorsal line a pair of small nerves (fig. 19 D) spring from the brain and pass upwards, while nearly in the same line with the optic nerves and a little behind them a larger pair of nerves take their origin.

Behind all these nerves there arises from the line of suture between the two supra-cesophageal ganglia a large median nerve which appears to supply the integument of the dorsal part of the head (PL 48, fig. 8 ; PL 49, figs. 11 14 d it).

Sympathetic System. In addition to the nerves just described there are two very important nerves which arise near the median ventral line, close to the hind end of the supracesophageal ganglia. The origin of these two nerves is shewn in the surface view (fig. 22 sy, and in section in fig. n). They at first tend somewhat forwards and pass into the muscles near the epithelium lining the groove on each side of the tongue. Here they suddenly bend backwards again and follow the grooves into the pharynx.

The two grooves are continuous with the two dorsal angles of the pharynx ; and embedded in the muscles of the pharynx, in juxtaposition with the epithelium, these two nerves may easily be traced in sections. They pass backwards the whole length of the pharynx till the latter joins the oesophagus.


Here they at once approach and shortly meet in the median dorsal line (fig. 16). They can only be traced for a very short distance beyond their meeting point. These nerves are, without doubt, the homologues of the sympathetic system of Chaetopods, occupying as they do the exact position which Semper has shewn to be characteristic of the sympathetic nerves in that group, and arising from an almost identical part of the brain 1 .

Histology of the Nervous System.

Ventral Cords. The histology of the ventral cords and cesophageal commissures is very simple and uniform. They consist of a cord almost wholly formed of nerve-fibres, placed dorsally, and a ventral layer of ganglion cells (figs. 16 and 20).

The fibrous portion of the cord has the usual structure, being formed mainly of longitudinal fibres, each probably being a bundle of fibres of various sizes, enveloped in a sponge-work of connective tissue. The larger bundles of fibres are placed near the inner borders of the cords. In this part of the cord there are placed a very small number of ganglion cells.

The layer of ganglion cells is somewhat crescent-shaped in section, and, as shewn in figs. 16 and 20, envelopes the whole ventral aspect of the fibrous parts of the cord, and even creeps up slightly on to the dorsal side. It is thicker on the inner than on the outer side, and increases considerably in bulk at each ganglionic enlargement. The cells of which it is composed are for the most part of a nearly uniform size, but at the border of the fibrous matter a fair sprinkling of larger cells is found.

The tracheal vessels supplying the nervous system are placed amongst the larger cells, at the boundary between the ganglionic and fibrous regions of the cords.

With reference to the peripheral nerve-stems there is not much to be said. They have for the most part a similar structure to the fibrous parts of the main cord, but are provided with a somewhat larger number of cells.

1 Vide Spengel, " Oligognathus Boncllioc." Naples Mittheilungen, Bel. III. pi. iv. fig- 52 B. 57


Sheath of tlie Ventral Cords. The ventral cords are enveloped by a double sheath, the two layers of which are often in contact, while in other cases they may be somewhat widely separated from each other. The inner layer is extremely thin and always very closely envelopes the nerve-cords. The outer layer is thick and fibrous, and contains a fair sprinkling of nuclei.

Supra-cesophageal Ganglia. In the present state of our knowledge a very detailed description of the histology of the supracesophageal ganglia would be quite superfluous, and I shall confine myself to a description of the more obvious features in the arrangement of the ganglionic and fibrous portions (PI. 50, fig. 19 A G).

The ganglion cells are in the first place confined, for the most part, to the surface. Along the under side of each ganglion there is a very thick layer of cells, continuous behind, with the layer of ganglion cells which is placed on the under surface of the cesophageal commissures. These cells have, moreover, an arrangement very similar to that in the ventral cords, so that a section through the supra-cesophageal ganglia has an obvious resemblance to what would be the appearance of a section through the united ventral cords. On the outer borders of the ganglia the cells extend upwards, but they end on about the level of the optic nerve (fig. 19 D). Immediately dorsal to this point the fibrous matter of the brain is exposed freely on the surface (fig. 19 A, B, &c., a}. I shall call the region of fibrous matter so exposed the dorso-lateral horn of white matter.

Where the two ganglia separate in front the ganglion cells spread up the inner side, and arch over so as to cover part of the dorsal side. Thus, in the anterior part, where the two ganglia are separate, there is a complete covering of ganglionic substance, except for a narrow strip, where the dorso-lateral lobe of white matter is exposed on the surface (fig. 19 A). From the point where the two ganglia meet in front the nerve-cells extend backwards as a median strip on the dorsal surface (fig. 19 D and E). This strip, becoming gradually smaller behind, reaches nearly, though not quite, the posterior limit of the junction of the ganglia. Behind it there is, however, a region where


the whole dorsal surface of the ganglia is without any covering of nerve-cells.

This tongue of ganglion cells sends in, slightly behind the level of the eyes, a transverse vertical prolongation inwards into the white matter of the brain, which is shewn in the series of transverse sections in fig. 19 E, and also in the vertical longitudinal section (PL 51, fig. 21), and in horizontal section in PL 51, fig. 22.

On the ventral aspect of each lobe of the brain there is present a very peculiar, bluntly conical protuberance of ganglion cells (PL 51, fig. 22), which was first detected by Grube (No. 10), and described by him as "a white thick body of a regular tetrahedral form, and exhibiting an oval dark spot in the middle of two of the faces." He further states that it is united by a delicate nerve to the supra-cesophageal ganglion, and regards it as an organ of hearing.

In Peripatus capensis the organ in question can hardly be described as tetrahedral. It is rather, of a flattened oval form, and consists, as shewn in sections (PL 50, fig. 19 C and D, d\ mainly of ganglion cells. In its interior is a cavity with a distinct bounding membrane : the cells of which it is composed vary somewhat in size, being smallest near the point of attachment. At its free end is placed a highly refractive, somewhat oval body, probably forming what Grube describes as a dark spot, half embedded in its substance, and kept in place by the sheath of nervous matter surrounding it. This body appears to have fallen out in my sections. The whole structure is attached to the under surface of the brain by a very short stalk formed of a bundle of cells and nervous fibres.

It is difficult to offer any interpretation of the nature of this body. It is removed considerably from the surface of the animal, and is not, therefore, so far as I can see, adapted to serve as an organ of hearing.

The distribution of the white or fibrous matter of the ganglia is not very easy to describe.

There is a central lobe of white matter (fig. 19 E), which is continuous from ganglion to ganglion, where the two are united. It is smaller behind than in front. On its ventral side it exhibits fairly well-marked transverse commissural fibres, con 572


necting the two halves of the ganglion. Laterally and somewhat ventrally it is prolonged into a horn (fig. 19 D, E, b], which I propose calling the ventro-lateral horn. In front it is placed in a distinct protuberance of the brain, which is placed ventrally to and nearly in the same vertical plane as the optic nerve. This protuberance is best shewn in the view of the brain from below given in PL 51, fig. 22. This part of the horn is characterized by the presence of large vertically-directed bundles of nerve-fibres, shewn in transverse section in fig. 190. Posteriorly the diameter of this horn is larger than in front (fig. 19, E, F, G), but does not give rise to a protuberance on the surface of the brain owing to the smaller development of the median lobe behind.

The median lobe of the brain is also prolonged into a dorsolateral lobe (fig. 19, a], which, as already mentioned, is freely exposed on the surface. On its ventral border there springs the optic nerve, and several pairs of sensory nerves already described (fig. 19 D, E), while from its dorsal border a pair of sensory nerves also spring, nearly in the same vertical plane as the optic nerves.

Posteriorly where the dorsal surface of the brain is not covered in with ganglion cells the dorso-lateral horn and median lobe of the brain become indistinguishable.

In the front part of the brain the median lobe of white matter extends dorsalwards to the dorsal strip of ganglion cells, but behind the region of the transverse prolongation of these cells, into the white matter already described (p. 890), there is a more or less distinctly defined lobe of white matter on the dorsal surface, which I propose calling the postero-dorsal lobe of white matter. It is shewn in the transverse sections (fig. 19 F and G, c). It gradually thins away and disappears behind. It is mainly characterized by the presence on the ventral border of definite transverse commissural fibres.



The skin is formed of three layers.

1. The cuticle.

2. The epidermis or hypodermis.

3. The dermis.

The cuticle is a layer of about O'CO2 mm. in thickness. Its surface is not, however, smooth, but is everywhere, with the exception of the perioral region, raised into minute secondary papillae, the base of which varies somewhat in diameter, but is usually not far from O'O2 mm. On the ventral surface of the body these papillae are for the most part somewhat blunt, but on the dorsal surface they are more or less sharply pointed. In most instances they bear at their free extremity a somewhat prominent spine. The whole surface of each of the secondary papillae just described is in its turn covered by numerous minute spinous tubercles. In the perioral region, where the cuticle is smooth, it is obviously formed of two layers which easily separate from each other, and there is I believe a similar division elsewhere, though it is not so easy to see. It is to be presumed that the cuticle is regularly shed.

The epidermis, placed immediately within the cuticle, is composed of a single row of cells, which vary, however, a good deal in size in different regions of the body. The cells excrete the cuticle, and, as shewn in fig. 32, they stand in a very remarkable relation to the secondary papillae of the cuticle just described. Each epidermis cell is in fact placed within one of these secondary papillae, so that the cuticle of each secondary papilla is the product of a single epidermis cell. This relation is easily seen in section, while it may also be beautifully shewn by taking a part of the skin which is not too much pigmented, and, after staining it, examining from the surface.

In fig. 32 a region of the epidermis is figured, in which the cells are exceptionally columnar. The cuticle has, moreover, in the process of cutting the section, been somewhat raised and carried away from the subjacent cells. The cells of the epidermis are provided with large oval nuclei, which contain a well


developed reticulum, giving with low powers a very granular appearance to the nuclei. The protoplasm of the cells is also somewhat granular, and the granules are frequently so disposed as to produce a very well-marked appearance of striation on the inner end of the cells. The pigment which gives the characteristic colour to the skin is deposited in the protoplasm of the outer ends of the cells in the form of small granules. An attempt is made to shew this in fig. 32.

At the apex of most, if not all, the primary wart-like papillae there are present oval aggregations, or masses of epidermis cells, each such mass being enclosed in a thickish capsule (fig. 31). The cells of these masses appear to form the wall of a cavity which leads into the hollow interior of a long spine. These spines when carefully examined with high objectives present a rather peculiar structure. The base of the spine is enveloped by the normal cuticle, but the spine itself, which terminates in a very fine point, appears, as shewn in fig. 31, to be continuous with the inner layer of the cuticle. In the perioral region the outer layer of the cuticle, as well as the inner, appear to be continued to the end of the spines. Within the base of the spine there is visible a finely striated substance which may often be traced into the cavity enclosed by the cells, and appears to be continuous with the cells. Attached to the inner ends of most of the capsules of these organs a delicate fibrillated cord may be observed, and although I have not in any instance succeeded in tracing this cord into one of the nervestems, yet in the antennas, where the nerve-stems are of an enormous size, I have satisfied myself that the minute nerves leaving the main nerve-stems and passing out towards the skin are histologically not to be distinguished from these fibrillated cords. I have therefore but little hesitation in regarding these cords as nerves.

In certain regions of the body the oval aggregations of cells are extremely numerous ; more especially is this the case in the antennas, lips, and oral papillae. On the ventral surface of the peripheral rings of the thicker sections of the feet they are also very thick set (fig. 20 P). They here form a kind of pad, and have a more elongated form than in other regions. In the antennae they are thickly set side by side on the rings of skin


which give such an Arthropod appearance to these organs in Peripatus.

The arrangement of the cells in the bodies just described led me at first to look upon them as glands, but a further investigation induced me to regard them as a form of tactile organ. The arguments for this view are both of a positive and a negative kind.

The positive arguments are the following :

(1) The organs are supplied with large nerves, which is distinctly in favour of their being sense organs rather than glands.

(2) The peculiar striae at the base of the spines appear to me like the imperfectly preserved remains of sense hairs.

(3) The distribution of these organs favours the view that they are tactile organs. They are most numerous on the antennas, where such organs would naturally be present, especially in a case like that of Pe'ripatus, where the nerve passing to the antennas is simply gigantic. On the other hand, the antennae would not be a natural place to look for an enormous development of dermal glands.

The lips, oral papillae, and under surface of the legs, where these bodies are also very numerous, are situations where tactile organs would be of great use.

Under the head of negative arguments must be classed those which tell against these organs being glandular. The most important of these is the fact that they have no obvious orifice. Their cavities open no doubt into the spines, but the spines terminate in such extremely fine points that the existence of an orifice at their apex is hardly credible.

Another argument, from the distribution of these organs over the body is practically the converse of that already used. The distribution being as unfavourable to the view that they are glands, as it is favourable to that of their being sense organs.


The apertures of the tracheal system are placed in the depressions between the papillae or ridges of the skin. Each of them leads into a tube, which I shall call the tracheal pit (fig. 30), the walls of which are formed of epithelial cells bounded


towards the lumen of the pit by a very delicate cuticular membrane continuous with the cuticle covering the surface of the body. The pits vary somewhat in depth; the pit figured was about O'CX) mm. It perforates the dermis and terminates in the subjacent muscular layer. The investigation of the inner end of the pit gave me some little trouble.

Transverse sections (fig. 30) through the trunk containing a tracheal opening shew that the walls of the pit expanded internally in a mushroom-like fashion, the narrow part being, however, often excentric in relation to the centre of the expanded part.

Although it was clear that the tracheae started from the expanded region of the walls of the pit, I could not find that the lumen of the pit dilated into a large vesicle in this part, and further investigation proved that the tracheae actually started from the slightly swollen inner extremity of the narrow part of the pit, the expanded walls of the pit forming an umbrella-like covering for the diverging bundles of tracheae.

I have, in fig. 30, attempted to make clear this relation between the expanded walls of the tracheal pits and the tracheae. In longitudinal sections of the trunk the tracheal pits do not exhibit the lateral expansion which I have just described, which proves that the divergence of the bundles of tracheae only takes place laterally and not in an antero-posterior direction. Cells similar in general character to those of the walls of the tracheal pits are placed between the branches of tracheae, and somewhat similar cells, though generally with more elongated nuclei, accompany the bundles of tracheae as far as they can be followed in my sections. The structure of these parts in the adult would, in fact, lead one to suppose that the tracheae had originated at the expense of the cells of pits of the epidermis, and that the cells accompanying the bundles of tracheae were the remains of cords of cells which sprouted out from the blind ends of the epidermis pits and gave rise in the first instance to the tracheae.

The tracheae themselves are extremely minute, unbranched (so far as I could follow them) tubes. Each opening by a separate aperture into the base of the tracheal pit, and measuring about O-QO2 mm. in diameter. They exhibit a faint transverse striation, which I take to be the indication of a spiral fibre.


[Moseley (Phil. Trans., 1874, PI. 73, fig. i) states that the tracheae branch, but only exceptionally.]

Situation of the tracheal apertures. Moseley states (No. 13) that the tracheae arise from the skin all over the surface of the body, but are especially developed in certain regions. He finds "a row of minute oval openings on the ventral surface of the body," the openings being "situate with tolerable regularity in the centres of the interspaces between the pairs of members, but additional ones occurring at irregular intervals. Other similar openings occur in depressions on the inner side of the conical foot protuberance." It is difficult in preserved specimens to make out the exact distributions of the tracheal apertures, but I have been able to make out certain points about them.

There is a double row of apertures on each side of the median dorsal line, forming two sub-dorsal rows of apertures. The apertures are considerably more numerous than the legs. There is also a double row of openings, again more numerous than the legs, on each side of the median ventral line between the insertions of the legs. Moseley speaks of a median row in this position. I think this must be a mistake.

Posteriorly the two inner rows approach very close to each other in the median ventral line, but I have never seen them in my section opening quite in the middle line. Both the dorsal and ventral rows are very irregular.

I have not found openings on the ventral or dorsal side of the feet but there are openings at the anterior and posterior aspects of the feet. There are, moreover, a considerable number of openings around the base of the feet.

The dorsal rows of tracheal apertures are continued into the head and give rise in this situation to enormous bundles of tracheae.

In front of the mouth there is a very large median ventral tracheal pit, which gives off tracheae to the ventral part of the nervous system, and still more in front a large number of such pits close together. The tracheae to the central nervous system in many instances enter the nervous system bound up in the same sheath as the nerves.



The general muscular system consists of (i) the general wall of the body; (2) the muscles connected with the mouth, pharynx, and jaws; (3) the muscles of the feet; (4) the muscles of the alimentary tract.

The muscular wall of the body is formed of (i) an external layer of circular fibres; (2) an internal layer of longitudinal muscles; (3) a layer of transverse fibres.

The layer which I have spoken of as formed of circular fibres is formed of two strata of fibres which girth the body somewhat obliquely (PI. 51, fig. 25). In the outer stratum the rings are arranged so that their ventral parts are behind, while the ventral parts of the rings of the inner stratum are most forward. Both in the median dorsal and ventral lines the layer of circular fibres become somewhat thinner, and where the legs are attached the regularity of both strata is somewhat interfered with, and they become continuous with a set of fibres inserted in the wall of the foot.

The longitudinal muscles are arranged as five bands (vide fig. 1 6), viz. two dorsal, two lateral, and three ventral. The three ventral may be spoken of as the latero-ventral and medioventral bands.

The transverse fibres consist of (i) a continuous sheet on each side inserted dorsally in the cutis, along a line opposite the space between the dorsal bands of longitudinal fibres, and ventrally between the ventro-median and ventro-lateral bands. Each sheet at its insertion slightly breaks up into separate bands. They divide the body-cavity into three regions a median, containing the alimentary tract, slime glands, &c., and two lateral, which are less well developed, and contain the nervous system, salivary glands, segmental organs, &c.

(2) Inserted a little dorsal to the transverse band just described is a second band which immediately crosses the first, and then passes on the outer side of the nervous cord and salivary gland, where such is present, and is inserted ventrally in the space between the ventro-lateral and lateral longitudinal band.


Where the feet are given off the second transverse band becomes continuous with the main retractor muscular fibres in the foot, which are inserted both on to the dorsal side and ventral side.

Muscular system of the feet. This consists of the retractors of the feet connected with the outer transverse muscle and the circular layer of muscles. In addition to these muscles there are intrinsic transverse muscles which cross the cavity of the feet in various directions (PI. 51, fig. 20). There is no special circular layer of fibres.

Histology of the muscle, The main muscles of the body are unstriated and divided into fibres, each invested by a delicate membrane. Between the membrane and muscle are scattered nuclei, which are never found inside the muscle fibres. The muscles attached to the jaws form an exception in that they are distinctly transversely striated.


The body-cavity, as already indicated, is formed of three compartments one central and two lateral. The former is by far the largest, and contains the alimentary tract, the generative organs, and the mucous glands. It is lined by a delicate endothelial layer, and is not divided into compartments nor traversed by muscular fibres.

The lateral divisions are much smaller than the central, and are shut .off from it by the inner transverse band of muscles. They are almost entirely filled with the nerve-cord and salivary gland in front and with the nerve-cord alone behind, and their lumen is broken up by muscular bands. They further contain the segmental organs which open into them. They are prolonged into the feet, as is the embryonic body-cavity of most Arthropoda.

The vascular system is usually stated to consist of a dorsal heart. I find between the dorsal bands of longitudinal fibres a vessel in a space shut off from the body-cavity by a continuation of the endothelial. lining of the latter (fig. 16). The vessel has definite walls and an endothelial lining, but I could not make out whether the walls were muscular. The ventral


part of it is surrounded by a peculiar cellular tissue, probably, as suggested by Moseley, equivalent to the fat bodies of insects. It is continued from close to the hind end of the body to the head, and is at its maximum behind. In addition to this vessel there is present a very delicate ventral vessel, by no means easy to see, situated between the cutis and the outer layer of circular muscles.


A series of glandular organs are found in Peripatus which have their external openings situated on the ventral surface of a certain number of the legs, and which, to the best of my belief, end internally by opening into the lateral compartments of the body-cavity. These organs are probably of an excretory nature, and I consider them homologous with the nephridia or segmental organs of the Chaetopoda.

In Peripatus capensis they are present in all the legs. In all of them (except the first three) the following parts may be recognized :

(1) A vesicular portion opening to the exterior by a narrow passage.

(2) A coiled portion, which is again subdivided into several sections.

(3) A terminal section ending by a somewhat enlarged opening into the lateral compartment of the body-cavity.

The last twelve pairs of these organs are all constructed in a very similar manner, while the two pairs situated in the fourth and fifth pairs of legs are considerably larger than those behind, and are in some respects very differently constituted.

It will be convenient to commence with one of the hinder nephridia. Such a nephridium from the ninth pair of legs is represented in fig. 28. The external opening is placed at the outer end of a transverse groove placed at the base of one of the feet, while the main portion of the organ lies in the body-cavity in the base of the leg, and extends into the trunk to about the level of the outer edge of the nerv.e-cord of its side. The external opening (p s) leads into a narrow tube (s d\ which gradually dilates into a large sack (s).


The narrow part is lined by small epithelial cells, which are directly continuous with and perfectly similar to those of the epidermis (fig. 20). It is provided with a superficial coating of longitudinal muscular fibres, which thins out where it passes over the sack, along which it only extends for a short distance.

The sack itself, which forms a kind of bladder or collecting vesicle for the organ, is provided with an extremely thin wall, lined with very large flattened cells. These cells are formed of granular protoplasm, and each of them is provided with a large nucleus, which causes a considerable projection into the lumen of the sack (figs. 20, 29 s). The epithelial wall of the sack is supported by a membrana propria, over which a delicate layer of the peritoneal epithelium is reflected.

The coiled tube forming the second section of the nephridium varies in length, and by the character of the epithelium lining it may be divided into four regions. It commences with a region lined by a fairly columnar epithelium with smallish nuclei (fig. 28 s c i). The boundaries of the cells of this epithelium are usually very indistinct, and the protoplasm contains numerous minute granules, which are usually arranged in such a manner as to give to optical or real sections of the wall of this part of the tube a transversely striated appearance. These granules are very probably minute balls of excretory matter.

The nuclei of the cells are placed near their free extremities, contrary to what might have been anticipated, and the inner ends of the cells project for very different lengths into the interior, so causing the inner boundary of the epithelium of this part of the tube to have a very ragged appearance. This portion of the coiled tube is continuous at its outer end with the thin-walled vesicle. At its inner end it is continuous with region No. 2 of the coiled tube (fig. 28 s c 2), which is lined by small closely-packed columnar cells. This portion is followed by region No. 3, which has a very characteristic structure (fig. 28 s c 3). The cells lining this part are very large and flat, and contain large disc-shaped nuclei, which are usually provided with large nucleoli, and often exhibit a beautiful reticulum. They may frequently be observed in a state of division. The protoplasm of this region is provided with similar granules to that in the first region, and the boundaries of the cells are usually


very indistinct. The fourth region is very short (fig. 28 s c 4), and is formed of small columnar cells. It gradually narrows till it opens suddenly into the terminal section (s o t], which ends by opening into the body-cavity, and constitutes the most distinct portion of the whole organ. Its walls are formed of columnar cells almost filled by oval nuclei, which absorb colouring matters with very great avidity, and thus renders this part extremely conspicuous. The nuclei are arranged in several rows.

The study of the internal opening of this part gave me some trouble. No specimens ever shew it as rounded off in the characteristic fashion of tubes ending in a cul-de-sac. It is usually somewhat ragged and apparently open. In the best preserved specimens it expands into a short funnel-shaped mouth, the free edge of which is turned back. Sections confirm the results of dissections. Those passing longitudinally through the opening prove its edges are turned back, forming a kind of rudimentary funnel. This is represented in fig. 29, from the last leg of a female. I have observed remains of what I consider to be cilia in this section of the organ. The fourth region of the organ is always placed close to the thin-walled collecting vesicle (figs. 28 and 29). In the whole of the coiled tube just described the epithelium is supported by a membrana propria, which in its turn is invested by a delicate layer of peritoneal epithelium.

The fourth and fifth pairs are very considerably larger than those behind, and are in other respects peculiar. The great mass of each organ is placed behind the leg, on which the external opening is placed, immediately outside one of the lateral nerve-cords. Its position is shewn in fig. 8.

The external opening, instead of being placed near the base of the leg, is placed on the ventral side of the third ring (counting from the outer end) of the thicker portion of the leg. It leads (fig. 27) into a portion which clearly corresponds with the collecting vesicle of the hinder nephridia. This part is not, however, dilated into a vesicle in the same sort of way, and the cells which form the lining epithelium have not the same characteristic structure, but are much smaller. Close to the point where the vesicle joins the coiled section of the nephridium the


former has a peculiar nick or bend in it. At this nick it is firmly attached to the ventral side of the foot by muscles and tracheae, and when cut away from its attachment the muscles and tracheae cannot easily be detached from it. The main part of the coils are formed by region No. i, and the epithelial cells lining this part present very characteristically the striated appearance which has already been spoken of. The large-celled region of the coiled tube (fig. 2 ; ") is also of considerable dimensions, and the terminal portion is wedged in between this and the commencing part of the coiled tube. The terminal portion with its internal opening is in its histological characters exactly similar to the homologous region in the hinder nephridia.

The three pairs of nephridia in the three foremost pairs of legs are very rudimentary, consisting, so far as I have been able to make out, solely of the collecting vesicle and the duct leading from them to the exterior. The external opening is placed on the ventral side of the base of the feet, in the same situation as that of the posterior nephridia, but the histological

part of the body to which it belongs), does not acquire the normal relations of a blastopore, but presents only those rudimentary features (deep groove connected with origin of mesoblast) which the whole blastopore of other tracheates presents.

We think it probable that the larval anus eventually shifts to the hind end of the body, and gives rise to the adult anus. We reserve the account of the internal structure of these embryos (Stages A E) and of the later stages for a subsequent memoir.

We may briefly summarise the more important facts of the early development of Peripatus capensis, detailed in the preceding account.

1. The greater part of the mesoblast is developed from the walls of the archenteron.

2. The embryonic mouth and anus are derived from the respective ends of the original blastopore, the middle part of the blastopore closing up.

3. The embryonic mouth almost certainly becomes the adult mouth, i.e. the aperture leading from the buccal cavity into the pharynx, the two being in the same position. The embryonic anus is in front of the position of the adult anus, but in all probability. shifts back, and persists as the adult anus.

4. The anterior pair of mesoblastic somites gives rise to the swellings of the praeoral lobes, and to the mesoblast of the head 1 .

There is no need for us to enlarge upon the importance of these facts. Their close bearing upon some of the most important problems of morphology will be apparent to all, and we may with advantage quote here some passages from Balfour's Comparative Embryology, which shew that he himself long ago had anticipated and in a sense predicted their discovery.

"Although the mesoblastic groove of insects is not a gastrula, it is quite possible that it is the rudiment of a blastopore, the gastrula corresponding to which has now vanished

1 We have seen nothing in any of our sections which we can identify as of socalled mesenchymatous origin.


from development." (Comparative Embryology, Vol. I. p. 378, the original edition 1 .)

"TRACHEATA. Insecta. It (the mesoblast) grows inwards from the lips of the germinal groove, which probably represents the remains of a blastopore." (Comparative Embryology, Vol. II. p. 291, the original edition 2 .)

"It is, therefore, highly probable that the paired ingrowths of the mesoblast from the lips of the blastopore may have been, in the first instance, derived from a pair of archenteric diverticula." (Comparative Embryology, Vol. II. p. 294, the original edition 3 .)

The facts now recorded were discovered in June last, only a short time before Balfour started for Switzerland ; we know but little of the new ideas which they called up in his mind. We can only point to passages in his published works which seem to indicate the direction which his speculations would have taken.

After speculating as to the probability of a genetic connection between the circumoral nervous system of the Ccelenterata, and the nervous system of Echinodermata, Platyelminthes, Chaetopoda, Mollusca, &c., he goes on to say :

" A circumoral nerve-ring, if longitudinally extended, might give rise to a pair of nerve-cords united in front and behind exactly such a nervous system, in fact, as is present in many Nemertines (the Enopla and Pelagonemertes), in Peripatus and in primitive molluscan types (Chiton, Fissurella, &c.). From the lateral parts of this ring it would be easy to derive the ventral cord of the Chaetopoda and Arthropoda. It is especially deserving of notice, in connection with the nervous system of the above-mentioned Nemertines and Peripatus, that the commissure connecting the two nerve-cords behind is placed on the dorsal side of the intestines. As is at once obvious, by referring to the diagram (fig. 231 B), this is the position this commissure ought, undoubtedly, to occupy if derived from part of a nervering which originally followed more or less closely the ciliated edge of the body of the supposed radiate ancestor." (Comparative Embryology, Vol. II. pp. 311, 312, the original edition 4 .)

1 This edition, Vol. n. p. 457. 2 This edition, Vol. III. p. 352.

3 This edition, Vol. m. p. 356. 4 This edition, Vol. in. pp. 378, 379.


The facts of development here recorded give a strong additional support to this latter view, and seem to render possible a considerable extension of it along the same lines.]


1. M. Lansdown Guilding. "An Account of a New Genus of Mollusca," Zoological Journal, Vol. II. p. 443, 1826.

2. M. Andouin and Milne-Edwards. " Classific. des Anndlides et description de celles qui habitent les cotes de France," p. 411, Ann. Scien. Nat. ser. I. Vol. xxx. 1833.

3. M. Gervais. "Etudes p. servir a 1'histoire naturelle des Myriapodes," Ann. Scien. Nat. ser. n. Vol. vn. 1837, p. 38.

4. Wiegmann. Wiegmann's Archiv, 1837.

5. H. Milne-Edwards. "Note sur le Peripate juluforme" Ann. Scien. Nat. ser. n. Vol. xvm. 1842.

6. Blanchard. "Sur Forganisation des Vers," chap. IV. pp. 137 141, Ann. Scien. Nat. ser. in. Vol. Vlll. 1847.

7. Quatrefages. " Anat. des Hermelles, note on," p. 57, Ann. Scien. Nat. ser. in. Vol. x. 1848.

8. Quatrefages. Hist. Nat. des Anneles, 1865, Appendix, pp. 675 6.

9. De Blainville. SuppL au Diet, des Sc. Nat. Vol. I.

10. Ed. Grube. " Untersuchungen lib. d. Bau von Peripatus Edwardsii? Archiv fur Anat. und Physiol. 1853.

11. Saenger. " Moskauer Naturforscher Sammlung," Abth. Zool. 1869.

12. H. N. Moseley. "On the Structure and Development of Peripatus capensis? Proc. Roy. Soc. N.O. 153, 1874.

13. H. N. Moseley. " On the Structure and Development of Peripatus capensis," Phil. Trans. Vol. CLXIV. 1874.

14. H. N. Moseley. "Remarks on Observations by Captain Hutton, Director of the Otago Museum, on Peripatus novce zealandice," Ann. and Mag. of Nat. History, Jan. 1877.

15. Captain Hutton. " Observations on Peripatus novce sealandice," Ann. and Mag. of Nat. History, Nov. 1876.

16. F. M. Balfour. "On Certain Points in the Anatomy of Peripatus capensis" Quart. Journ. of Micr. Science, Vol. xix. 1879.

17. A. Ernst. Nature, March loth, 1881.




A. Anus. a. Dorso-lateral horn of white matter in brain, a.g. Accessory gland of male (modified accessory leg gland), at. Antenna, at. n. Antennary nerve, b. Ventro-lateral horn of white matter of brain. b. c. Body-cavity. bl. Blastopore. C. Cutis. c. Postero-dorsal lobe of white matter of brain. e.g. Supracesophageal ganglia, cl. Claw. c. m. Circular layer of muscles, co. Commissures between the ventral nerve-cords, co. i. Second commissure between the ventral nerve-cords. co 1 . 2. Mass of cells developed on second commissure, cor. Cornea, c. s. d. Common duct for the two salivary glands. . cu. Cuticle, d. Ventral protuberance of brain. d. 1. m. Dorsal longitudinal muscle of pharynx. d. n. Median dorsal nerve to integument from supraoesophageal ganglia, d. o. Muscular bands passing from the ventro-lateral wall of the pharynx at the region of its opening into the buccal cavity. E. Eye. E. Central lobe of white matter of brain, e. n. Nerves passing outwards from the ventral cords, ep. Epidermis, ep.c. Epidermis cells. F. i, F. a, &c. First and second pair of feet, c. f. Small accessory glandular tubes of the male generative apparatus. F.^. Ganglionic enlargement on ventral nerve-cord, from which a pair of nerves to foot pass off. f. gl. Accessory foot-gland. F. n. Nerves to feet. g. co. Commissures between the ventral nerve-cords containing ganglion cells, g. o. Generative orifice. H. Heart, h. Cells in lateral division of body-cavity. hy. Hypoblast, i.j. Inner jaw. j. Jaw. j. n. Nerves to jaws. L. Lips. /. Lens. /. b. c. Lateral compartment of body-cavity, le. Jaw lever (cuticular prolongation of inner jaw lying in a backwardly projecting diverticulum of the buccal cavity). /. m. Bands of longitudinal muscles. M. Buccal cavity. M 1 . Median backward diverticulum of mouth or common salivary duct which receives the salivary ducts, me. Mesenteron. mes. Mesoblastic somite, m. 1. Muscles of jaw lever, m. s. Sheets of muscle passing round the side walls of pharynx to dorsal body wall. od. Oviduct, ce. OZsophagus. a's. co. OZsophageal commissures, o.f. g. Orifice of duct of foot-gland, o.j. Outer jaw. op. Optic ganglion, op. n. Optic nerve, or.g. Ganglionic enlargements for oral papillae, o r. n. Nerves to oral papillae, or. p. Oral papillas. o. s. Orifice of duct of segmental organ, ov. Ovary, p. Pads on ventral side of foot. p. Common duct into which the vasa deferentia open. p. c. Posterior lobe of brain. /. d. c. Posterior commissure passing dorsal to rectum. /./. Internal opening of nephridium into body cavity, ph. Pharynx, pi. Pigment in outer ends of epidermic cells, pi. r. Retinal pigment, p. n. Nerves to feet. p.p. Primary papilla, pr. Prostate. R. Rectum. Re. Retinal rods. R. m. Muscle of claw. s. Vesicle of nephridium. j 1 . Part of 4th or 5th nephridium which corresponds to vesicle of other nephridia.

1 The explanations of the figures printed within inverted commas are by Professor Balfour, the rest are by the Editors.


s. c. i. Region No. i of coiled tube of nephridium. s. c. 2. Region No. i of ditto. s. c. 3. Region No. 3 of ditto. s. c. 4. Region No. 4 of ditto, s. d. Salivary duct. s. g. Salivary gland, si. d. Reservoir of slime gland, sl.g. Tubules of slime gland. s. o. i, 2, 3, &c. Nephridia of ist, 2nd, &c., feet. s. o.f. Terminal portion of nephridium. s.p. Secondary papilla, st. Stomach, sf. e. Epithelium of stomach, sy. Sympathetic nerve running in muscles of tongue and pharynx, sy 1 . Origin of pharyngeal sympathetic nerves. T. Tongue, t. Teeth on tongue, te. Testis. tr. Trach.e0e. tr. c. Cells found along the course of the tracheae. tr. o. Tracheal stigma, tr. p. Tracheal pit. tit. Uterus, v. c. Ventral nerve cord. v. d. Vas deferens. v. g. Imperfect ganglia of ventral cord.


Fig. i. Peripatus capensis, x 4 ; viewed from the dorsal surface. (From a drawing by Miss Balfour. )


Fig. 2. A left leg of Peripatus capensis, viewed from the ventral surface ; x 30. (From a drawing by Miss Balfour.)

P'ig. 3. A right leg of Peripatus capensis, viewed from the front side. (From a drawing by Miss Balfour.)

Fig. 4. .The last left (i7th) leg of a male Peripatus capensis, viewed from the ventral side to shew the papilla at the apex of which the accessory gland of the male, or enlarged crural gland, opens to the exterior. (From a drawing by Miss Balfour.) Prof. Balfour left a rough drawing (not reproduced) shewing the papilla, to which is appended the following note. " Figure shewing the accessory genital gland of male, which opens on the last pair of legs by a papilla on the ventral side. The papilla has got a slit-like aperture at its extremity."

Fig. 5. Ventral view of head and oral region of Peripatus capensis. (From a drawing by Miss Balfour.)


Figs. 6 and 7 are from one drawing.

Fig. 6. Peripatus capensis dissected so as to shew the alimentary canal, slime glands, and salivary glands ; x 3. (From a drawing by Miss Balfour.)

Fig. 7. The anterior end of Fig. 6 enlarged ; x 6. (From a drawing by Miss Balfour.) The dissection is viewed from the ventral side, and the lips, L., have been cut through in the middle line behind and pulled outwards, so as to expose the jaws, /., which have been turned outwards, and the tongue, T. , bearing a median row of chitinous teeth, which branches behind into two. The junction of the salivary ducts, j. d., and the opening of the median duct so formed into the buccal cavity is also shewn. The muscular pharynx, extending back into the space between the ist and 2nd pairs of legs, is followed by a short tubular oesophagus. The latter opens into the large stomach with plicated walls, extending almost to the hind end of the animal. The stomach at its point of junction with the rectum presents an S-shaped ventrodorsal curve.

A. Anus. at. Antenna. F. i, K. 2. First and second feet. /. Jaws. L. Lips. ae. OZsophagus. or. p. Oral papilla, ph. Pharynx. R. Rectum, s. d. Salivary duct. s. g. Salivary gland, si. d. Slime reservoir, si. g. Portion of tubules of slime gland, st. Stomach. T. Tongue in roof of mouth.

Fig. 8. Peripatus capensis, X4; male. (From a drawing by Miss Balfour.) Dissected so as to shew the nervous system, slime glands, ducts of the latter passing into the oral papilla, accessory glands opening on the last pair of legs (enlarged crural glands), and segmental organs, viewed from dorsal surface. The first three pairs of segmental organs consist only of the vesicle and duct leading to the exterior. The fourth and fifth pairs are larger than the succeeding, and open externally to the crural glands. The ventral nerve-cords unite behind dorsal to the rectum.

A. Anus. a. g. Accessory generative gland, or enlarged crural gland of the iyth leg. at. Antenna, c. g. Supra-oesophageal ganglia with eyes. co. Commissures between the ventral nerve-cords, d. n. Large median nerve to dorsal integument from hinder part of brain. F. i, i, &c. Feet. g. o. Generative orifice, <x. (Esophagus. KS. co. QEsophageal commissures, or. p. Oral papilla, p.d.c. Posterior dorsal commissure between the ventral nerve-cords, ph. Pharynx, p. n. Nerves to feet, one pair from each ganglionic enlargement. si. d. Reservoir of slime gland. si. g. Tubules of slime gland. s. o. i, 2, 3, &c. Segmental organs. v. c. Ventral nervecords, "v. g. Imperfect ganglia of ventral cords.

Figs. 9 and 10. Left jaw of Peripatus capensis (male), shewing reserve jaws. (From a drawing by Miss Balfour.)

Fig. 9. Inner jaw. Fig. 10. Outer jaw.


Figs, ii 16. A series of six transverse sections through the head of Peripatus capensis.

Fig. n. The section is taken immediately behind the junction of the supracesophageal ganglia, c. g., and passes through the buccal cavity, M., and jaws, o.j. and i.j.

Fig. 12. The section is taken through the hinder part of the buccal cavity at the level of the opening of the mouth into the pharynx and behind the jaws. The cuticular rod-like continuation (le.) of the inner jaw lying in a backwardly directed pit of the buccal cavity is shewn; on the right hand side the section passes through the opening of this pit.

Fig. 1 3. The section passes through the front part of the pharynx, and shews the opening into the latter of the median backward diverticulum of the mouth (M 1 ), which receives the salivary ducts. It also shews the commencement of the ventral nerve-cords, and the backwardly projecting lobes of the brain.

Fig. 14. The section passes through the anterior part of the pharynx at the level of the second commissure (co. 2), between the ventral nerve-trunks, and shews the mass of cells developed on this commissure, which is in contact with the epithelium of the backward continuation of the buccal cavity (M 1 ).


Fig. 15. Section through the point of junction of the salivary ducts with the median oral diverticulum.

Fig. 1 6. Section behind the pharynx through the oesophagus.

b. c. Body-cavity. C. Cutis. c. b. c. Central compartment of body-cavity, c. g. Supra-oesophageal ganglia, c. m. Layer of circular muscles, co. Commissure between ventral nerve-cords. co. i. Second commissure between the ventral nerve-cords. co 1 . i. Mass of cells developed on second commissure (probably sensory), c. s. d. Common duct for the two salivary glands, d. /. m. Dorsal longitudinal muscles of pharynx, d. o. Muscles serving to dilate the opening of the pharynx. Ep. Epidermis, e. n. Nerve passing outwards from ventral nerve-cord. H. Heart, i.j. Inner jaw. j. p. Jaw papillae. L. Lips of buccal cavity. /. b. c. Lateral compartment of body-cavity, le. Rod-like cuticular continuation of inner jaw, lying in a pit of the buccal cavity. /. m. Bands of longitudinal muscles. M. Buccal cavity. M 1 . Median backward continuation of buccal cavity, m. 1. Muscles of jaw lever, m. s. Muscular sheets passing from side walls of pharynx to dorsal body wall. ce. CEsophagus. ces. co. CEsophageal commissures. o.j. Outer jaw. ph. Pharynx, s. d. Salivary duct. s. g. Salivary gland, si. d. Reservoir of slime gland, sy. Sympathetic nerves running in muscles of tongue or pharynx, sy 1 . Origin of sympathetic nerves to pharynx. T. Tongue, v. c. Ventral nerve-cords.

Figs. 17, 1 8. Two longitudinal horizontal sections through the head of Peripatus capensis. Fig. 17 is the most ventral. They are both taken ventral to the cerebral ganglia. In Fig. 17 dorsal tracheal pits are shewn with tracheae passing off from them. (Zeiss a a, Hartnack's camera.) C. Cutis. c. s. d. Common salivary duct. ep. Epidermis, i.j. Inner jaw. M. Buccal cavity. M 1 . Median backward diverticulum of mouth, o.j. Outer jaw. s. d. Salivary ducts. T. Tongue, t. Teeth on tongue, tr. Tracheae, tr. p. Tracheal pits.


Fig. 19. "A, B, c, D, E, F, G. Seven transverse sections illustrating the structure of the- supra- cesophageal ganglia. (Zeiss A, Hartnack's camera.) a. Dorso-lateral horn of white matter. b. Ventro-lateral horn of white matter, c. Postero-dorsal lobe of white matter, d. Ventral protuberance of brain, e. Central lobe of white matter, o.p. Optic ganglion.

" A. Section through anterior portions of ganglia close to the origin of the antennary nerve. B. Section a little in front of the point where the two ganglia unite, c. Section close to anterior junction of two ganglia. D. Section through origin of optic nerve on the right side. E. Section shewing origin of the optic nerve on the left side. F. Section through the dorso-median lobe of white matter. G. Section near the termination of the dorsal tongue of ganglion cells."


Fig. 10. Portion of a transverse section through the hinder part of Peripatus capensis (male). The section passes through a leg, and shews the opening of the segmental organ (p. s.), and of a crural gland, o.f.g., and the forward continuation of the enlarged crural gland of the i7th leg (/ g!.). (Zeiss a a, Hartnack's camera.) a-g. accessory gland of male (modified crural gland of last leg), c. Cutis. cL Claw. cu. Cuticle, ep. Epidermis, Crural gland, h. Cells in lateral compartment of body cavity, o.f. g. Orifice of accessory foot gland, o. s. Opening of segmental organ, p. Three spinous pads on ventral surface of foot. pr. Prostate. R. M. Retractor muscle of claw. s. Vesicle of nephridium. s. c. i. Region No. i of coiled part of nephridium. si. g. Tubule of slime gland, s. o. t. Terminal portion of nephridium. st. Stomach, st. e. Epithelium of stomach, v. c. Ventral nerve-cord, v. d. Vas deferens.

Fig. 21. "Longitudinal vertical section through the supra-oesophageal ganglion and oesophageal commissures of Peripatus capensis. (Zeiss a a, Hartnack.)" at. Antenna, e. Central lobe of white matter. /. Part of jaw. s. g. Salivary gland.

Fig. 22: drawn by Miss Balfour. Brain and anterior part of the ventral nervecords of Peripatus capensis enlarged and viewed from the ventral surface. The paired appendages (d) of the ventral surface of the brain are seen, and the pair of sympathetic nerves (sy 1 ) arising from the ventral surface of the hinder part.

From the commencement of the cesophageal commissures (as. co. ) pass off on each side a pair of nerves to the jaws (/. .).

The three anterior commissures between the ventral nerve-cords are placed close together; immediately behind them the nerve-cords are swollen, to form the ganglionic enlargements from which pass off to the oral papillce a pair of large nerves on each side (or. n. )

Behind this the cords present a series of enlargements, one pair for each pair of feet, from which a pair of large nerves pass off on each side to the feet (p. n). at. n. Antennary nerves, co. Commissures between ventral cords, d. Ventral appendages of brain. E. Eye. e. n. Nerves passing outwards from ventral cord. F-g- Ganglionic enlargements from which nerves to feet pass off. j. n. Nerves to jaws. or. g. Ganglionic enlargement from which nerves to oral papillce pass off. or. n. Nerves to oral papillae, p.c. Posterior lobe of brain, p. n. Nerves to feet. s.y. Sympathetic nerves.

Fig. 23. "Longitudinal horizontal section through the head of Peripatus capensis, shewing the structure of the brain, the antennary and optic nerves, &c. (Zeiss a a, Hartnack's camera.)" at. Antenna, at. n. Antennary nerve, cor. Cornea, e. Central mass of white matter. /. Lens. op. n. Optic nerve, ph. Pharynx, p.p. Primary papilla covered with secondary papillte and terminating in a long spine, sy. Pharyngeal sympathetic nerves.

Fig. 24. "Eye of Peripatus capensis, as shewn in a longitudinal horizontal section through the head. The figure is so far diagrammatic that the lens is represented as filling up the whole space between the rods and the cornea. In the actual section there is a considerable space between the parts, but this space is probably artificial, being in part caused by the shrinkage of the lens and in part by the action of the razor. (Zeiss c, Hartnack's camera.)" (It appears that the ganglionic region of the eye is covered by a thin capsule, which is omitted in the figure.)

cor. Cornea. /. Lens. op. Optic ganglion, op-, n. Optic nerve. //'. r. Pigment. Re. rods. s. p. Secondary papillae.

Fig. 25. Longitudinal horizontal section through the dorsal skin, shewing the peculiar arrangement of the circular muscular fibres. (Zeiss A, Hartnack's camera.)


Fig. 26. Portion of ventral cord of Peripatus capensis enlarged, shewing two ganglionic enlargements and the origin of the nerves and commissures. (From a drawing by Miss Balfour.)

co. Commissures. E. n. Nerves passing out from ventral cords. F. n. Nerves to feet. g. co. Commissures between the ventral cords containing ganglion cells, v. g. Ganglionic enlargements.

Fig. 27. Segmental organ from the 5th pair of legs of Peripatus capensis. This nephridium resembles those of the 4th legs, and differs from all the others in its large size and in the absence of any dilatation giving rise to a collecting vesicle on its external portion (enlarged). The terminal portion has the same histological characters as in the case of the hinder segmental organs. (From a drawing by Miss Balfour. )

Fig. 28. Segmental organ or nephridium from the 9th pair of legs of Peripatus capensis^ shewing the external opening, the vesicle, the coiled portion and the terminal portion with internal opening (enlarged). (From a drawing by Miss Balfour.)

o. s. External opening of segmental organ, p.f. Internal opening of nephridium into the body-cavity (lateral compartment). s. Vesicle of segmental organ, j 1 . Portion of segmental organ of 4th and 5th legs, corresponding to vesicle of the other nephridia. s. c. i. First or external portion of coiled tube of nephridium, lined by columnar epithelium with small nuclei ; the cells project for very different distances, giving the inner boundary of this region a ragged appearance, s. c. 2. Region No. 2 of coiled tube of nephridium, lined by small closely-packed columnar cells, s. c. 3. Region No. 3 of coiled tube of segmental organ, lined by large flat cells with large disc-shaped nuclei, s. c. 4. Region No. 4 of coiled tube of nephridium ; this region is very short and lined by small columnar cells, s. o. t. Terminal portion of nephridium.

Fig. 29. " Portion of nephridium of the hindermost leg of Peripatus capensis, seen in longitudinal and vertical section. The figure is given to shew the peritoneal funnel of the nephridium. Portions of the collecting sack (s.) and other parts are also represented. (Zeiss B, Hartnack's camera.)"

p.f. Peritoneal funnel, s. Vesicle, s.c.i, s.c.i, s.c.$. Portions of coiled tube.

Fig. 30. " Section through a tracheal pit and diverging bundles of tracheal tubes" taken transversely to the long axis of the body. (Zeiss E, oc. 2.) (From a rough drawing by Prof. Balfour.)

tr. Tracheae, shewing rudimentary spiral fibre, tr. c. Cells resembling those lining the tracheal pits, which occur at intervals along the course of the trachere. tr. s. Tracheal stigma, tr. p. Tracheal pit.

Fig- 31. "Sense organs and nerves attached from antenna of Peripatus capensis (Zeiss, immersion 2, oc. 2.)" (From a rough drawing by Prof. Balfour.) The figure shews the arrangement of the epidermis cells round the base of the spine. The spine is seen to be continuous with the inner layer of the cuticle.


Fig. 32. Section through the skin of Peripatus capensis ; it shews the secondary papillae covered with minute spinous tubercles and the relation of the epidermis to them. (The cuticle in the process of cutting has been torn away from the subjacent cells.) The cells of the epidermis are provided with large oval nuclei, and there is a deposit of pigment in the outer ends of the cells. The granules in the protoplasm of the inner ends of the cells are arranged in lines, so as to give a streaked appearance. (Zeiss E, oc. 2.) (From a rough drawing by Prof. Balfour.)

c. Dermis. cu. Cuticle, ep. c. Epidermis cells, pi. Deposit of pigment in outer ends of epidermis cells, s.p. Secondary papillae.

Fig. 33. Female generative organs of Peripatus capensis, x 5. (From a rough drawing by Prof. Balfour.) The following note was appended to this drawing: "Ovary rather to dorsal side, lying in a central compartment of body-cavity and attached to one of the longitudinal septa, dividing this from the lateral compartment between the penultimate pair of legs and that next in front. The oviducts cross before opening to the exterior, the right oviduct passing under the rectum and the left over it. They meet by opening into a common vestibule, which in its turn opens below the anus. On each side of it are a pair of short papillae (aborted feet ?)."

F. 16, 17. Last two pairs of legs. od. Oviduct, ov. Ovary, ut. Uterus, v. c. Nerve-cord.


Cite this page: Hill, M.A. (2019, September 15) Embryology The Works of Francis Balfour 1-25. Retrieved from

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