Part 10

Though the structure of the cirri is very uniform, yet we meet with some peculiarities. In Chelonobia, the segments of the posterior cirri bear only two pairs of main spines; whereas in some varieties of _Balanus balanoides_, they carry as many as ten pairs in a longitudinal row; but in this latter species, the number of these spines varies, in a singular manner, from six to ten pairs. In Tubicinella, the pairs of spines on the segments of the posterior cirri are arranged so closely one under the other, that they appear almost like a single transverse row. Considering the whole family, the third pair of cirri differs most in structure in the different genera. Thus, in _Chthamalus antennatus_, the anterior (or outer) ramus (Pl. 29, fig. 3) is thicker and much longer than the posterior (or inner) ramus; the number of the segments in one instance being, in the two rami, 53 and 18; in the longer ramus, the spines are arranged abnormally, tending to form a little circle round each segment; and the whole ramus may be said to be antenniformed, and I believe acts as an organ of touch: the relative number of the segments, I may add, in the two rami and the arrangement of their spines varies greatly in this species. In two other species of the same genus Chthamalus, we have _occasionally_ the anterior ramus in some degree antenniformed, so that this whole structure is variable. In the allied _Chamæsipho columna_, it is the posterior or inner ramus which is antenniformed, but this peculiar development is more plainly marked in the case of the second pair of cirri than in that of the third pair. In _Tetraclita porosa_ it is, also, the posterior ramus of the third pair which is antenniformed; in this third pair, and indeed in the other cirri, the relative numbers of the segments vary extremely. A similar structure in the third pair, but in a lesser and variable degree, may be observed in some of the other species of Tetraclita. In _Balanus vestitus_, also, we have, in the third pair, an analogous structure. It is scarcely possible to believe that the circumstance of the second pair of legs, which answer to the third pair of cirri, being antenniformed in certain decapod Crustaceans, is an accidental coincidence; it must be owing to some special affinity in the two groups.

In Chelonobia, the third pair of cirri is of unusual length compared with the second pair, but does not otherwise differ from the type of its sub-family: in Coronula and its allies, on the other hand, the third pair is very short and broad, as may be seen (Pl. 29, fig. 5) in Xenobalanus: in this latter genus, the front surfaces of the segments of the pedicels (fig. 6) of the posterior cirri, are extremely protuberant, almost as in _Scalpellum vulgare_.

The last peculiarity in the cirri at all worth mentioning, is in the sub-genus Acasta, in which, differently from in all other known Cirripedes, the anterior ramus of the fourth pair does not absolutely resemble the rami of the fifth and sixth pairs; in most of the species, the spines on this anterior ramus are more crowded together, are larger, and are mingled with some short thick points; and the spines in the dorsal tufts are also longer than in the two posterior pairs of cirri; but in _A. sulcata_ (Pl. 29, fig. 2), and in a lesser degree in _A. cyathus_ and _A. purpurata_, the front margins of the lower segments of this anterior ramus, and of the upper segment of the pedicel, are developed into strong, downwardly curved teeth: it is very remarkable that so beautiful a structure should be extremely variable, as it certainly is in _Acasta sulcata_.

_Caudal Appendages._

With extremely few exceptions, these are present in all the Lepadidæ and Verrucidæ; whereas amongst the Balanidæ they occur only in the two species of Pachylasma, and in one species of Catophragmus; these being the genera most closely allied to the Lepadidæ, and where, consequently, their presence might have been anticipated. These appendages are seated close together over the anus; they are multiarticulate, each segment being sub-cylindrical, with a few small bristles round its upper edge.

_Alimentary Canal._

I have not much on this head to add to what I have said under the Lepadidæ. As in that family, the strong internal membrane of the œsophagus terminates in a remarkable, bell-shaped expansion (Pl. 26, fig. 3, _g′_), which, as observed by M. St. Ange, serves to keep the upper broad end of the stomach expanded. The œsophagus is well furnished with constrictor and radiating muscles for closing and opening it; and it is thus capable of a strong swallowing movement. The stomach runs down to the lower end of the prosoma, and then doubling back on itself extends to the anus. As the prosoma is much elongated in Tubicinella and Xenobalanus, so is the stomach of unusual length in these genera. In several species of Balanus, the upper edge of the stomach is surrounded by from six to eight cæca; these cæca I ascertained, in _Balanus perforatus_, are branched, and penetrate a considerable way into the body; and some of them at least expand a little at their extremities. Each cæcum, from the manner in which it retained fluid, must, I think, be furnished, at the point where it enters the stomach, with a sphincter muscle. In Tetraclita, Chthamalus, Tubicinella, Coronula, and Xenobalanus, there are no cæca; but in Xenobalanus and _Coronula balænaris_, there are longitudinal, approximate folds in the upper, broad end of the stomach, which would serve to expose the food to a greater extent of digesting surface.[37]

[37] The presence and absence of these cæca in genera so closely allied as Balanus and Tetraclita, shows, I think, that these cavities are not of high importance; and I must doubt whether Von Siebold's view ('Anatomie Comparée,' tom. i, p. 445), that these cæca form a passage to a true or isolated liver, such as exists in the higher Crustacea, can be admitted. Cæca are said by Von Siebold to occur in some of the Entomostraca, as Daphnia, Argulus, &c.

As in the case of the Lepadidæ, a transparent, structureless, epithelial tube, composed of chitine, containing more or less digested food, is found, in specimens preserved in spirits, occupying the whole length of the stomach, and where there are cæca, sending branched prolongations into them. It does not extend into the œsophagus or into the rectum. This epithelial tube or model of the stomach, filled with excrement, is expelled by the rectum, whole, that is in a single piece, as I observed in some living specimens of _Balanus balanoides_: in some specimens, however, of _Chthamalus stellatus_, the excrement was ejected, perhaps from the animal being confined, in fragments, and the sack thus became befouled. Beneath the epithelial layer, the stomach is lined by a delicate, pulpy and cellular mucus layer, which easily peels off in flakes: this is surrounded by a muscular layer with the fibres closely approximated and transverse; and this by a layer of stronger, longitudinal muscles, but more distant from each other. Lastly, outside this double muscular layer, there is a rather thick, somewhat laminated, pulpy layer, abounding with cells, often nucleated, and frequently containing much oily matter. This structure agrees closely with Dr. C. H. Jones's[38] account of the external covering of the stomach in Daphnia, and which he believes to be hepatic: as in Daphnia, there does not seem to be any ducts. I may here observe, that within the upper part of the prosoma, but not immediately connected with the stomach, I have often observed much white pulpy substance, permeated by lacunal passages, and exhibiting no structure except some excessively minute cells.

[38] 'Philosophical Transactions,' 1849, p. 116. Karsten ('Nov. Actorum Acad. Nat. Cur.,' 1845, tab. xx) has excellently figured the testes, as the hepatic glands; and has indicated the ovaria as salivary glands; it is singular that this anatomist overlooked the ducts which lead from his supposed hepatic glands, into the vesiculæ seminales, within which he observed spermatozoa.

The rectum, lined by membrane continuous with that investing the thorax (and seen through it, in Pl. 26, fig. 8, _c_), extends inwards to about opposite the bases of the third or fourth pairs of cirri. It is longitudinally plaited; the ends of the folds forming a sort of valve where joined on to the stomach. It is coated by circular, transverse muscular fibres: judging from the movements, the anus itself is surrounded by a strong sphincter muscle. The anus opens on the dorsal surface of the thorax (fig. 8, _b_); but as in the genera, in which caudal appendages occur, it opens under them, the orifice, I believe, is homologically terminal, and owes its dorsal aspect to the aborted state of the whole abdomen, and to the great development of the probosciformed penis; for the anus may be said to be situated on the dorsal base of this organ.

Altogether we see that the alimentary canal is of a very simple structure. The food, judging from the contents of the stomach, seems generally to be composed of infusoria and minute animals: but in the case of Tetraclita, I have been surprised at the size and number of the included amphipod, isopod, and entomostracan Crustaceans, in one case, together with an annelid. I have, also, sometimes seen some confervoid matter within the stomach.

_Circulatory System._

On this subject I can add nothing, except to express my conviction that there is no heart, or true vessels; the circulation being strictly lacunal. A passage has often been quoted from Poli, in which he states that he saw a pulsating organ, close above the anus; but I have seen this movement, which appeared to me to be a convulsive twitching of the sphincter muscle of this orifice. The largest lacunal channel extends down the middle of the rostral compartment of the shell: and this answers to the rostral channel down the peduncle in the Lepadidæ. Large nerves and the main pair of unbranched ovarian tubes (Pl. 25, fig. 1, leading into _g_) extend along this channel. At the basis (at least in Coronula) this channel joins on to a large circular one, running all round the sack, and sending off branches into the mass of ovarian tubes and cæca.

_Nervous System._

It has been shown in my former volume, that in Lepas and in some other genera of the Lepadidæ, there are six main ganglions; one supra-œsophageal, and five infra-œsophageal, or thoracic. In Pollicipes, however, there are only four thoracic ganglions, the last ganglion supplying the three posterior pairs of cirri with nerves, whereas in the other genera, the last ganglion supplies only the fifth and sixth pairs of cirri. In this genus, moreover, the lateral fusion of the ganglions has been so complete, that there is no evidence of their having been formed by the union of two. Amongst sessile cirripedes, we discover evidence of much higher concentration even than in Pollicipes. My chief examination has been confined to _Coronula diadema_, and to _Balanus tintinnabulum_: and in these genera we find (and the fact appears to me highly remarkable) as high a degree of concentration in the infra-œsophageal ganglion as in any decapod Crustacean, for instance, as in Maia, judging from the figure given by Milne Edwards; for all the nerves, with the exception of those connected with the supra-œsophageal ganglions, radiate from a single great ganglion.[39] The nervous system is, moreover, otherwise complicated.

[39] It must, however, be observed that, according to Mr. Dana, there is in certain suctorial Entomostracans, as in Caligus, only one infra-œsophageal ganglion. Mr. Dana speaks of this as resulting from reduction. In Cirripedes, from the gradation which may be observed from Lepas through Pollicipes into Balanus, the ganglions are certainly not reduced but concentrated. In Van de Hoeven's figure of the nervous system in Limulus, there is seen to be no chain of thoracic ganglions; all the nerves rising from the circa-œsophageal collar; but this, on the other hand, seems hardly developed into a ganglion.

To begin with _Coronula diadema_ the great infra-œsophageal ganglion (Pl. 27, fig. 1, A) is seated nearly opposite to the anterior margin of the second pair of cirri, which are homologous with the first pair of legs in the decapod Crustaceans. This ganglion shows no trace of any longitudinal medial suture; its shape is hardly discoverable, for it is formed by the union of eleven principal pairs of nerves, besides several arising from its under surface; in outline, however, it may be said to be divided into a posterior and anterior half; the latter being somewhat heart-shaped, and the posterior half elongated. The nerves going to the five posterior pairs of cirri rise from the posterior margin of the ganglion, and run for some distance in a sheet, parallel and close together; the pair, however, going to the second pair of cirri soon branches off from the others. Each of these nerves enters at the inner and posterior margin of the cirrus to which it belongs, and, at least in the case of the first pair, divides into two branches as it enters. The nerves (Pl. 27, fig. 1, _r^5_, _r^6_) going to the fifth and sixth pairs of cirri are more closely united together than are the others, and appear, till they branch off, like a single large nerve. That which belongs to the sixth cirrus gives off, opposite to the fifth cirrus, a branch (_s_) nearly as large as itself, which enters the probosciformed penis. I may remark, that homologically this is the only abdominal nerve in any cirripede of the Order. From the under side of the nerves which run to the five posterior pairs of cirri, small branches are given off, extending dorsally into the thorax.

The anterior end of the great infra-œsophageal ganglion is formed by the union of a set of nerves, extending parallel in a bundle in a directly opposite direction to those running to the five posterior pairs of cirri. These nerves consist of an outer larger pair (_r^1_) entering the first pair of cirri; and within these, and rather dorsally to their roots, we have the circa-œsophageal chord (_c_, _c_), or collar nerve; between the roots of the latter, and on the ventral surface (or near side of the figure), there are three closely united, small pairs, running to the gnathites, and, as I believe, to the olfactory sacks. From the under (or dorsal) surface of the anterior end of the ganglion, two nerves, larger even than the circa-œsophageal chord, and which I shall call the splanchnic pair (_d_, _d_) arise; and the singular course of these nerves will presently be described; between this great pair, there is a single (_b_) medial nerve, which runs down and branches into that large diverging muscle, which is attached to the upper ventral surface of the stomach. Posteriorly to these three nerves, we have two pairs of much smaller nerves (not figured), running dorsally into the body, so that we have seven nerves rising from the under surface of the infra-œsophageal ganglion. I need only further add, that on each side of this ganglion, between the nerves going to the first and second pairs of cirri, there is a moderately sized nerve (_k_), which appeared to run into the muscles of the thorax: a nerve in a similar position is figured by Milne Edwards in Maia.

The circa-œsophageal chord (_c_, _c_) nearly equals in length the whole distance from the centre of the main ganglion to the posterior end of the thorax. This collar bows out on each side, where passing the œsophagus (_œ_), which is seated at its anterior end. From the collar a branch is given off on each side, which I traced as far as between the mandibles and maxillæ; from analogy with other Crustaceans, it perhaps runs to the mandibles. The collar has not a transverse commissure, such as described by Milne Edwards in the Podophthalmia, and as figured by Van de Hoeven in Limulus.

The supra-œsophageal ganglions (B) present a singular contrast with the infra-œsophageal ganglion in their little development, size, or degree of confluence. They lie directly under the basal edge of the labrum. They are laterally quite distinct, and consist merely of a slight enlargement of the circa-œsophageal chord. From the anterior edge of each ganglion, a broad nerve (_f_) extends for some distance in a straight line, and, on close examination, can be seen to be formed of two nerves closely united, of which the inner and smaller one, after a space, appears to cross over the larger nerve: both become at this point tortuous, and, giving off branches (_m_, _m_), form a plexus. The two nerves (_f_) then bend inwards, and almost touching each other, run down, together with the two ovarian simple ducts, along the rostral compartment of the shell. No doubt, if the smaller branches from these nerves could be traced, they would be seen to form a network over the whole sack; and would therefore enclose, as in a cage, the rest of the nervous system. These nerves correspond, I believe, to the two pair of antennular nerves of ordinary Crustaceans, and hence I will call them by this name. Just in front, at the outside corners of the two supra-œsophageal ganglions (B), a branch (_e′_) arises, which I traced to the ends of the adductor scutorum muscle, and to those several muscles which serve to retract the interspace of membrane between the mouth and the adductor.

The pair of great splanchnic nerves above alluded to, which arise from the anterior and dorsal surface of the infra-œsophageal ganglion, are in Pl. 27, fig. 1, _d_ _d_, (and in fig. 2), laid flat; but in nature they first bow outwards, and then, penetrating deeper into the body, approach each other, and running nearly parallel, pass round the lower end of the œsophagus: their course consequently is nearly similar to that of the circa-œsophageal chord, with this difference, that the outwardly bowed portion is situated near the infra-œsophageal, instead of near the supra-œsophageal ganglion. The splanchnic nerve, a little beyond the supra-œsophageal ganglion, joins a plexus (_d′_); and into this plexus another large nerve (_e_) which I will call the supra-splanchnic nerve, sends branches; this nerve takes an almost semicircular bend over the ovarian glands. The supra-splanchnic nerves (_e_, _e_), though appearing to spring from the supra-œsophageal ganglions, do really arise, as may be seen by tracing the constituent fibres, from the circa-œsophageal chord. The plexus (_d′_) lies close to the coats of the upper end of the stomach: several branches, proceeding from it, run further on, but I was able to trace only a few of them: one went (at least in the case of _Balanus perforatus_), to the adductor scutorum muscle: another branch spread out on the flanks of the prosoma: I strongly suspect that one branch goes to the acoustic sack: it appeared, also, as if some of the small branches entered the second plexus (_m_), where the inner antennular nerve and ophthalmic nerve cross over the outer antennular nerve.

I have called the nerves (_dd_, _ee_) splanchnic and supra-splanchnic, from their course and apparent function in supplying the viscera. In the descriptions of the nervous system of other Crustaceans I can find nothing analogous to my great splanchnic nerve (_dd_); the so-called supra-splanchnic nerves (_ee_), which arise from the circa-œsophageal chord, seem to be the analogues of the ordinary splanchnic nerves, though these latter are always described as uniting into a single medial branch. The plexus (_d′_) is the cervical ganglion of M. Martin St. Ange,[40] who has likewise indicated the course of my splanchnic and supra-splanchnic nerves; but the plexus, when viewed as a transparent object, hardly appears to me to be ganglionic in its nature. In my former volume on the Lepadidæ, I quite misunderstood the course of these splanchnic nerves.

[40] 'Mémoire sur l'Organisation des Cirripèdes,' p. 19.

From the commissure between the two supra-œsophageal ganglions, a straight chord (Pl. 27, fig. 1, _g_) arises, which terminates in a small ganglion (C), barely exhibiting traces of being formed of two laterally confluent ganglions. This is the ophthalmic ganglion. The chord connecting it with the two supra-œsophageal ganglions is accompanied by a small nerve (_h_) which runs on to the muscles round the adductor scutorum muscle; the chord is encased by much fibrous tissue, and its dissection is thereby rendered difficult. From the ophthalmic ganglion, on each side, a nerve (_i_) goes forth and crosses the antennular nerve; these, if I could have traced them, would have been found to run, as may be safely inferred from what is known in _Balanus tintinnabulum_, to a pair (D, D) of eyes.

In _Balanus tintinnabulum_, the structure of the great infra-œsophageal ganglion (Pl. 27, fig. 2, A) is essentially the same as described under Coronula. The great pair of splanchnic nerves springing from its under side, are here actually twice as large as the circa-œsophageal chord. The plexus (_d′_) formed by the splanchnic nerve (_d_), on each side, with the supra-splanchnic nerve (_e_), which arises close posteriorly to the supra-œsophageal ganglion, is here much less complicated, but is perfectly distinct; and there was no appearance of the cervical ganglion of M. Martin St. Ange. The chord (_g_) running from between the two supra-œsophageal ganglions to the ophthalmic ganglion, is nearly as large as the double antennular nerve (_f_) on each side of it. The ophthalmic chord (which is accompanied in its whole course by a small branch running to the adductor scutorum muscle) terminates in a small ophthalmic ganglion (C), which seems to be formed by the almost complete fusion of two ganglions. This ganglion is hardly larger than the chord which it terminates: it appeared to me to give rise to more than one pair of nerves, and a single nerve (to my surprise) joined the branch just mentioned, which goes to the adductor scutorum muscle.

From each supra-œsophageal ganglion, two closely united antennular nerves (_f_) extend, of which the inner one crosses over the main or exterior nerve, nearly opposite to the ophthalmic ganglion, and here forms (_m_) a plexus. The structure of this plexus I was not able, any more than in Coronula, to make out thoroughly, but I traced quite distinctly a long nerve (_i_) running from it into what must be considered as the eye. As in the case of Coronula, I traced a nerve on each side from the ophthalmic ganglion into the plexus, where I lost it; and as here in Balanus, I saw on each side of the ophthalmic ganglion a cut off nerve, of about the size of that which runs from the plexus on each side into the eye, I think we may safely conclude that the latter or optic nerve does really arise from the ganglion here called ophthalmic. I may add that the analogy of the nervous system in the Lepadidæ most strongly confirms the view of this latter being the ophthalmic ganglion.

_Eyes and Vision._