Before proceeding any further, I should observe that figs. [3] and [4], in Pl. [26], represent the membranes of the mouth of [Coronula diadema], perfectly cleaned. In fig. [3], all the front part of the mouth has been removed, the mandible on one side, the labrum with the two palpi, and the œsophagus being alone left, and these are viewed from the inner side; the front part, however, of the supra-œsophageal cavity has been cut away. In fig. [4], the labrum, with the œsophagus, has been removed, whilst the two outer maxillæ, the right-hand inner maxilla and mandible (with the exterior and basal portions, d, d′′, of one palpus) are seen from the outside; but in order that these parts should all be shown, the whole of the right-hand side of the mouth has been spread out, for the teeth of the mandible should have stood in a vertical line between the two outer maxillæ. In the mandibles, the free upper part is separated, by a distinct articulation, from the square piece of thickened membrane (fig. [3], c1) on which it is supported; and this latter is separated by a second articulation from a portion of thickened membrane (c2), the basal edge of which forms the third and lowest articulation, separating the mouth from the body. This basal, thickened portion of membrane curls round and inwards, towards the outer maxillæ or front of the mouth, and its terminal points sometimes even penetrate a little way within the muscles, like apodemes: it is not distinctly separated by any line or suture from the membrane, which forms the whole broad labrum; so that I at first concluded that the labrum dipped under the mandibles, and thus afforded a support on which they were articulated; but this appears so opposed to all analogy, that it is more probable that the above basal thickened portion of membrane is truly the basal segment of the mandibles, completely confluent with the labrum; and it is, I think, not very improbable that even a large portion of what in appearance belongs to the labrum, namely, those concavities to which the muscles of the mandibles are attached, may, also, be part of the basal segment of the mandibles. Whether or no there really are two segments beneath the upper free portion of the mandibles, which have become laterally confluent with other parts, I must think that the square thickened piece of membrane (c1) represents at least one segment. I may here observe, that Prof. Milne Edwards seems to consider the mandible of the higher Crustaceans as answering homologically to the haunch of the leg; but, according to M. Brullé,[35] there ought to be two basal segments (sous-maxillaire and maxillaire) bearing the proper mandible, and giving rise, on the outer side, to the palpus,—a structure which perfectly corresponds with my view of the mandible and palpus in Cirripedes.
[35] ‘Annales des Scienc. Nat.,’ 3d series, Zoolog., tom. ii, p. 271.
Maxillæ: the point whence the long apodeme (b′, Pl. [26], fig. [4] and fig. [7]) arises, according to Audouin’s views, must mark an articulation, and this would separate the upper free segment from the lower segments, which I believe to be laterally confluent with the organs on each side. The thickened membrane, of which the upper free part is formed, extends a little distance beyond the insertion of the apodeme; and this small portion beneath the point of insertion may possibly answer to the square, thickened piece of membrane, or second segment, supporting the mandibles. Beneath it, a rather wide expanse of thin, flexible membrane reaches down to the basal fold surrounding the mouth, and may thus form the third segment.
Outer Maxillæ: the upper free segment has a spinose lobe (a′′, Pl. [26], figs. [2] and [4]), on its inner face, which may indicate a lower and second, almost free segment. Passing over this, we have, on the outside of the mouth, beneath the free, upper segments, an expanse of membrane, which, on the side, close to the inner maxillæ, is perforated (Pl. [26], fig. [4], n) by orifices which I believe are olfactory. In some species, as in [Bal. eburneus] and [improvisus], there is a longitudinal medial suture in this expanse of membrane, which I suppose indicates the lateral confluence of the middle segments of the two outer maxillæ. A short, transverse articulation or fold separates this middle segment (fig. [4], a1) of each maxilla from the third or basal segment; and this latter (a2) is separated from the body by a very distinct fold, which (at least amongst the Lepadidæ) sends inwards a short, medial, tongue-formed apodeme. Here, then, we apparently have, as in the mandibles, two segments under the upper free segment of each outer maxilla, laterally confluent with the adjoining organs. But I must state that, in old specimens, and only in old specimens of [Coronula diadema], I have found under the outer maxilla an additional transverse ridge and fold, which plainly shows how easily a mere thickening of the membrane might be mistaken for an articulation. I can, however, hardly persuade myself that the articulated membrane, under the free part of the mandibles, which has now been figured and described, has no homological signification; and the fusion of the palpus and labrum seems too plain to be mistaken. Hence I must conclude that the mouth, in the Cirripedia, does truly exhibit a compounded structure of a very peculiar nature.
Cirri.
There are always six pairs; each biramous and multiarticulated, supported on a pedicel formed of two segments. A shield-like swelling at the exterior bases of these pedicels often appears like another segment; but such, I believe, is not its nature. The five posterior pairs answer to the five pairs of ambulatory legs in the higher Crustaceans; and as in the case of the latter, the three, or the four hindermost pairs almost invariably resemble each other. The first pair, which is homologous with the outer maxillipeds of ordinary Crustaceans, is separated by an interval from the second pair;—though this is not the case with the legs of the pupa, from which the cirri are metamorphosed. These anterior cirri are attached to the lateral edges of the mouth, namely to the thickened rim of membrane, forming the supposed basal segment of the mandibles. They are capable of more diversified movements than the other cirri: the anterior ramus is always elongated, with the terminal segments more or less tapering, and is directed beyond (or anteriorly to) the mouth: the shorter ramus closely resembles in structure the rami of the second pair. In the [Chthamalinæ] the second pair, and in the [Balaninæ] the second and third pairs (as will be more particularly described under these sub-families) differ in structure from the posterior pairs, from which they are separated by a slight interval. The number of segments on the posterior cirri is often great, amounting in [Chelonobia] even to fifty. Each segment normally is furnished on its inner face, which is usually somewhat protuberant, with from two to rarely eight or ten pairs of long spines or bristles, placed in a double row; the two spines in the lower pairs stand nearer to each other, and are shorter than the spines in the upper pairs. Between each pair of spines there is either a single, very thin bristle, or often a tuft of such. The pairs are directed somewhat upwards, and they diverge when the cirri are uncurled; their function is obviously to entangle the prey. On the dorsal or exterior surface of each segment, close to its upper margin, there is a tuft of spines, often composed of thicker and thinner spines; these, I believe, serve to prevent any creature intruding within the sack. On both sides of the upper margin of each segment, there is generally a row of short, blunt, excessively minute spines, which only deserve notice, inasmuch as it is by their increase in number and size, and by the spreading out of the dorsal tufts, and, lastly, by the increase of the little tuft intermediate between the pairs of spines situated in front, that the segments on the two or three anterior pairs of cirri become covered, like brushes, with bristles. The bristles or spines on the second and third cirri are often, especially in [Tetraclita], doubly and coarsely pectinated. The bristles on the pedicels follow the same arrangement as on the rami; namely, being in regular pairs on the posterior cirri, and crowded thickly, like a brush, on the anterior cirri. The segments in the shorter ramus of the first cirrus, and in both rami of the second, and often of the third cirrus, are broader than the segments of the posterior cirri; they are, also, especially in the genus [Balanus], frequently produced in their upper, ante-lateral corners, into remarkable prolongations (see Pl. [29], fig. [4], of the third cirrus of [Bal. perforatus]), clothed on their inner surfaces, and at their extremities, by numerous bristles. The number of the segments in each cirrus is in some degree variable, and increases with age; this is likewise the case, to a certain extent, with the number of the spines borne on each segment.
As compared with ordinary Crustaceans, I presume the two rami answer to the “tige” and “palpe” of Milne Edwards; and the pedicel (as I have called it) to the two basal segments of the leg.[36] The “fouet” or flabellum does not appear to be developed in any Cirripede; for though the filamentary appendages in certain genera of Lepadidæ, might at first be thought to be of this nature, yet their usual position beneath the basal articulation of the first pair of cirri, and the occasional presence of more than one, proves, I think, that such is not the case.
[36] According to this author’s new nomenclature, the pedicel would consist of the coxopodite and basipodite; the tige would be the ischiopodite and following segments; and the palpe would be the exopodite; the epipodite or flabellum being absent. (‘Annales des Sciences Naturelles,’ tom. xviii, 1852.)
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.