The accessory Malpighian bodies, or at any rate one of them, appear to have curious relations to the segmental tubes. The necks of some of the anterior segmental tubes (Pl. 20, fig. 5) close to their openings into the primary Malpighian bodies are provided with a small knob of cells which points towards the preceding segment and is usually connected with it by a fibrous band. This knob is most conspicuous in the male, and in very young animals or almost ripe embryos. In several instances in a ripe male embryo it appeared to me to have a lumen, and to be continued directly forwards into the accessory Malpighian body of the preceding segment. One such case is figured in the middle segment on Pl. 20, fig. 5. In this embryo segmental tubes were present in the segments immediately succeeding those connected with the vasa efferentia, and at the same time these segments contained ordinary and accessory Malpighian bodies. The segmental tubes of these segments were not, however, connected with the Malpighian body of their proper segment, but instead, turned forwards and entered the segment in front of that to which they properly belonged. I failed to trace them quite definitely to the accessory Malpighian body of the preceding segment, but, in one instance at least, there appeared to me to be present a fibrous connection, which is shewn in the figure already referred to, Pl. 20, fig. 5, r.st. In any case it can hardly be doubted that this peculiarity of the foremost segmental tubes is related to what would seem to be the normal arrangement in the next few succeeding segments, where each segmental tube is connected with a Malpighian body in its own segment, and more or less distinctly with an accessory Malpighian body in the preceding segment.
In the male the anterior segmental tubes, which even in the embryo exhibit signs of atrophy, become in the adult completely aborted (as has been already shewn by Semper), and remain as irregular tubes closed at both ends, which for the most part do not extend beyond the Wolffian duct (Pl. 20, fig. 4, r.st). In the adult, the first two or three segments with these aborted tubes contain only accessory Malpighian bodies; the remaining segments, with aborted segmental tubes, both secondary and primary Malpighian bodies. In neither case are the Malpighian bodies connected with the aborted tubes.
The Malpighian bodies in Scyllium present no special peculiarities. The outer layer of their capsule is for the most part formed of flattened cells; but, between the opening of the segmental tube and the efferent tubulus of the kidney, their cells become columnar. Vide Pl. 20, fig. 5. The convoluted tubuli continuous with them are, I believe, ciliated in their proximal section, but I have not made careful investigations with reference to their finer structure. Each segment is connected with the Wolffian duct by a single tube at the hinder end of the segment. In the kidney proper, these tubes become greatly prolonged, and form the ureters.
It has already been stated that the semen is carried by vasa efferentia from the testes to the anterior segments of the Wolffian body, and thence through the coils of the Wolffian body to the Wolffian duct. The nature of the vasa will be discussed in the embryological section of this chapter: I shall here confine myself to a simple description of their anatomical relations. The consideration of their connections naturally falls under three heads: (1) the vasa efferentia passing from the testes to the Wolffian body, (2) the mode in which these are connected with the Wolffian body, and (3) with the testis.
In Pl. 20, fig. 4, drawn for me from nature by my friend Mr Haddon, are shewn the vasa efferentia and their junctions both with the testes and the kidney. This figure illustrates better than any description the anatomy of the various parts. Behind there are two simple vasa efferentia (v.e.) and in front a complicated network of vasa, which might be regarded as formed of either two or four main vessels. It will be shewn in the sequel that it is really formed of four distinct vessels. Professor Semper states that there is but a single vas efferens in Scyllium canicula, a statement which appears to me unquestionably erroneous. All the vasa efferentia fall into a longitudinal duct (l.c), which is connected in succession with the several segments of the Wolffian body (one for each vas efferens) which appertain to the testis. The hind end of the longitudinal duct is simple, and ends blindly close to its junction with the last vas efferens; but in front, where the vasa efferentia are complicated, the longitudinal duct also has a complicated constitution, and forms a network rather than a simple tube. It typically sends off a duct to join the coils of the Wolffian body between each pair of vasa efferentia, and is usually swollen where this duct parts from it. A duct similar to this has been described by Semper as Nierenrandcanal in several Elasmobranchii, but its existence is expressly denied in the case of Scyllium! It is usually found in Amphibia, as we know from Bidder and Spengel's researches. Spengel calls it Längscanal des Hoden; the vessels from it into the kidney he calls vasa efferentia, and the vessels to it, which I speak of as vasa efferentia, he calls Quercanale.
The exact mode of junction of the separate vasa efferentia with the testis is difficult to make out on account of the opacity of the basal portion of the testis. My figure shews that there is a network of tubes (formed of four main tubes connected by transverse branches) which is a continuation of the anterior vasa efferentia, and joined by the two posterior ones. These tubes receive the tubuli coming from the testicular ampullæ. The whole network may be called, with Semper, the testicular network. While its general relations are represented in my figure, the opacity of the testes was too great to allow of all the details being with certainty filled in.
The kidneys of Scyllium stellare, as might be expected, closely resemble those of Scy. canicula. The ducts of the kidney proper, have, in the former species, a larger number of distinct openings into the urinogenital cloaca. In two male examples I counted seven distinct ureters, though it is not impossible that there may have been one or two more present. In one of my examples the ureters had seven distinct openings into the cloaca, in the other five openings. In a female I counted eleven ureters opening into the Wolffian duct by seven distinct openings. In the remaining parts of the excretory organs the two species of Scyllium resemble each other very closely.
As may be gathered from Prof. Semper's monograph, the excretory organs of Scyllium canicula are fairly typical for Elasmobranchii generally. The division into kidney and Wolffian body is universal. The segmental openings may be more numerous and larger, e.g. Acanthias and Squatina, or absent in the adult, e.g. Mustelus and Raja. Bladder-like swellings of the Wolffian duct in the female appear to be exceptional, and seminal bladders are not always present. The variations in the ureters and their openings are considerable, and in some cases all the ureters are stated to fall into a single duct, which may be spoken of as the ureter par excellence[344], with the same relations to the kidneys as the Wolffian duct bears to the Wolffian body. In some cases Malpighian corpuscles are completely absent in the Wolffian body, e.g. Raja.
The vasa efferentia of the testes in Scyllium are very typical, but there are some forms in which they are more numerous as well as others in which they are less so. Perhaps the vasa efferentia are seen in their most typical form in Centrina as described and figured (Pl. XXI) by Professor Semper, or in Squatina vulgaris, as I find it, and have represented it on Pl. 20, fig. 8. From my figure, representing the anterior part of the Wolffian body of a nearly ripe embryo, it will be seen that there are five vasa efferentia (v.e) connected on the one hand with a longitudinal canal at the base of the testes (n.t) and on the other with a longitudinal canal in the Wolffian body. Connected with the second longitudinal canal are four Malpighian bodies, three of them stalked and one sessile; from which again proceed tubes forming the commencements of the coils of the anterior segments of the Wolffian body. These Malpighian bodies are clearly my primary Malpighian bodies, but there are in Squatina, even in the generative segments, secondary Malpighian bodies. What Semper has described for Centrina and one or two other genera, closely correspond with what is present in Squatina.