I may here mention that I tried a few simple experiments on the senses of [Balanus balanoides], [B. crenatus], and [Chthamalus stellatus]. I found these three species very sensitive to shadows, that is, to an object like my hand passing even quickly, and at the distance of about a foot, between them and the source of the light.[42] They were indifferent to a gradual change from bright to obscure light; but instantly perceived and drew in their cirri, when my hand was passed between the basin in which they were kept and the window, even when this was tried rather late on a dusky evening; and likewise when my hand was passed between them and a single candle. I took, of course, the precaution of passing my hand in other directions, but this never produced any effect. These species are moderately sensible to any vibration in the vessel in which they were kept, but they were indifferent to noises made in the air, or in the water. I found it impossible to touch, under water, an individual shell ever so lightly with a needle, without all the immediately surrounding individuals, when several adhered together, perceiving it, and retracting their cirri: it made no difference whether the one touched had already withdrawn its cirri and was motionless: from this fact, and from seeing that a similar but slighter effect was produced by touching the rock on which the specimens adhered, I infer that the perception by the others of the one being touched, is communicated by vibration. When an individual was touched under water, not by a needle, but by a pointed camel-hair brush, it generally withdrew its cirri, but the neighbouring specimens took no notice: when touched by a single hair of the brush, no notice was taken, unless the skin of the orifice leading into the sack was so touched. In these trials, it is of course necessary carefully to avoid intercepting the light. I could not make out that cirripedes perceived odours diffused in the water.
[42] I find that this fact was long ago observed by Von Siebold, ‘Anatomie Comparée,’ tom. i, p. 434.
Acoustic Organs.
These are situated in the same position as in the Lepadidæ, namely, in a slight swelling on the sides of the thorax (Pl. [25], fig. [1], d′) just beneath the basal articulation of the first pair of cirri. The orifice in [Tubicinella] and [Xenobalanus] is slightly produced, or is tubular; the free part in the former genus projecting 5/100ths of an inch. The structure of all the parts is essentially the same as in the Lepadidæ, but I think all are proportionally larger. The external membrane of the body is turned inwards at the orifice, as a short flattened tube, which widens considerably (being, in a middle-sized specimen of [Coronula], 4/100ths of an inch in width) before it abruptly terminates. The meatus, as I have called the sack-like cavity which encloses the true acoustic sack or vesicle, is formed of pulpy membrane, and is apparently continuous with the corium of the whole body, but by dissection it can be separated entire. The acoustic vesicle is of various shapes, as we shall immediately see; but in all essential respects it is identical with the same part in the Lepadidæ; it is formed of the same peculiar, soft, elastic, brownish, transparent tissue, which seems to be composed of fine, transverse pillars, becoming towards the outside fibrous, and at their inner ends appearing when viewed vertically from above, like hyaline points. In [Coronula diadema], I observed on the outside of the acoustic vesicle, some excessively minute bristles, only 1/3000ths of an inch in length, seated on little eminences. I examined carefully the contents of the vesicle in this species, in specimens well preserved in spirits, and there was nothing within but a very little, thin, pulpy fluid, and a few yellowish nucleated cells, here and there aggregated into small groups. In [Coronula], the flattened acoustic vesicle is elongated, with a somewhat sinuous, but not very irregular margin (Pl. [27], fig. [4]), and is without any ridges on the surface; its neck or orifice projects at right angles to the elongated portion, which stands obliquely to the tubular orifice of the meatus. In a moderately-sized specimen of [Coronula diadema], the elongated portion of the acoustic vesicle was, 6/100ths of an inch in length. In [Tubicinella], the acoustic vesicle is heart-shaped, with the neck attached to its broader end; and the surface is covered by zig-zag ridges. In [Balanus tintinnabulum] (fig. [3]), the acoustic vesicle is almost square at the lower end, with the neck placed at one of the upper corners; on the external surface, there is an oblique prominent ridge or fold, which sends off downwards another ridge; its length, in a large individual, was 5/100ths of an inch.
In all these cases, the acoustic vesicle is mainly attached by its neck, to the upper end of the sack-like meatus; but there is likewise a layer of soft, pulpy, cellular matter, slightly connecting that side of the vesicle which is opposite to the neck, with the walls of the meatus or outer sack. The mouth or orifice of the vesicle is closed by a delicate lid or diaphragm, which can easily be separated; and this diaphragm is formed by the expansion of a large nerve, which here abruptly terminates. In a very large specimen of [Coronula diadema] I clearly made out the existence of this nerve, and traced its course for some distance from the point where the summit of the meatus and the neck of the vesicle are joined together; the nerve first runs posteriorly, and then turns inwards and doubles back or anteriorly; and I clearly followed it to the antero-lateral sides of the uppermost end of the stomach, where it seemed to enter a ganglion, so that I unfortunately cut it off, but found only a slight plexus, with the cut off nerve apparently running onwards with nearly the same diameter. The diameter is great, fully equalling, in its widest part, that of the circa-œsophageal chord; but it is very much flattened, and so has not nearly so much bulk as that nerve. Before it reached the stomach, it gave off one branch, which ran towards the mouth. The only nerves which, from their size, could, I think, be continuous with this from the acoustic sack, are the main branches proceeding from that plexus (d′) formed by the interbranching of the splanchnic and supra-splanchnic nerves.[43]
[43] I have always feared that anatomists would reject my view of these organs being acoustic, owing to the absence of otolithes; but I observe that so high an authority as Von Siebold (‘Anatomie Comparée,’ tom. i, p. 433) does not believe that otolithes occur in the acoustic organs even of the highest Crustacea. He considers an “ampoule volumineuse, a parois mince, remplie d’un liquide transparent,” and a “membrane tympanique,” though having a fissure in the centre, as sufficient. I may here remark, that the nerve proceeding from the acoustic vesicle in Cirripedes, and apparently running to the splanchnic nerve, may easily be placed in connexion with the antennular nerves, by the second plexus (m) in figs. [1] and [2], pl. [27]. I should infer from Von Siebold’s remarks on his ampoule volumineuse in the higher Crustacea, that my acoustic vesicle answered to the labyrinth in higher animals.
Olfactory Sacks.
I can add nothing to the account given of these organs under the Lepadidæ: I saw them in all the genera which I examined for this object. In [Coronula diadema] the orifices are large; they are seated in the usual position (Pl. [26], fig. [4], n), in the confluent segments, beneath the free part of the outer maxillæ, and somewhat exteriorly, or as near as possible to the inner maxillæ. In no sessile cirripede are the orifices produced or tubular, as is the case with several genera amongst the Lepadidæ. I failed, as heretofore, in tracing with certainty the nerve, which appears to enter the base of the sack, to its ganglion.
Male Organs of Generation.
All the Cirripedes of the family we are now describing, are bisexual or hermaphrodite; and no instance has been observed of the presence of males or complemental males. I have very little to add to the observations made by M. Martin St. Ange and R. Wagner,[44] and to those given in my former volume. The testes seem always to be confined to within the thorax, including the prosoma. With their ducts, they resemble club-moss or stag’s horns, with the extremities a little enlarged: a figure[45] of a small portion from [Balanus perforatus] is given in Pl. [25], fig. [2]. It is quite surprising how like in structure and appearance the branching ovarian tubes often are to the testes with their ducts; but the latter are of smaller diameter. Two main ducts generally unite just before entering the broad, often reflexed, end of the vesicula seminalis: in [Coronula balænaris], however, I observed four ducts entering this receptacle. The two vesiculæ seminales, lying within the thorax and prosoma, are usually very long and tortuous: they are formed of a thin inner tunic, which is strengthened by thicker reticulated lines, and of an outer layer of transverse fibres, which are either elastic, or probably muscular, as they serve to expel the contents with force when the end is cut off. The inner tunic is prolonged up the probosciformed penis, at the base of which the two vesiculæ unite.[46] The contents of the vesiculæ are commonly pulpy and cellular; and from the cells the spermatozoa are developed; soon after their development, they are, as it appears, expelled.