Of the internal organs, whilst the Cirripede is still within the larva, I have already mentioned the stomach with its pair of cæca: from the retracted position of the thorax and rudimentary abdomen, and consequently of the anus, compared with these parts in the larva, the alimentary canal is not above half its former length. There is, as yet, no trace of the filaments supposed by some to act as branchiæ, at the base of the first pair of cirri. Nor could I perceive a trace of the testes or vesiculæ seminales: the penis is represented by a minute, apparently imperforate projection. I have already briefly described the pair of large, gut-formed bodies in the larva, into the anterior ends of which the cement-ducts ran, and evidently derived their slightly opaque, cellular contents. At a very early age, before the young Cirripede can be distinctly made out, the posterior ends of these gut-formed bodies are absorbed, so as not to pass beyond the cæca of the stomach. When the young Cirripede is plainly developed within the larva, these bodies in a relatively reduced condition are still distinct near the cæca, and at the opposite or anterior end (i. e. lower, in the position in which Cirripedes are usually figured), they have branched out into a sheet of delicate inosculating tubes; these could be traced by every stage, until, in the young perfected Cirripede, they filled the peduncle as ordinary ovarian tubes. In the larva, the two gut-formed bodies or incipient ovaria keep of equal thickness from one to the other end, but in the mature Cirripede, the ovarian tubes in the peduncle and the small, glandular, grape-like masses, near the stomach-cæca, are connected only by a delicate tube; this I failed in tracing in specimens in the very immature condition of those now under description.
The larva fixes itself with its sternal surface parallel and close to the surface of attachment, and the antennæ become cemented to it: if the Cirripede, after its metamorphosis had remained in this position, the cirri could not have been exserted, or only against the surface of attachment; but there is a special provision, that the young Cirripede shall immediately assume its proper position at right angles to the position which it held whilst within the larva, namely with its posterior end upwards. This is effected in a singular manner by the exuviation of the great compound eyes, which we have seen are fastened to the outer arms of the double °UU°-like, sternal apodemes: these together with the eyes stretch transversely across, and internally far up into, the body of the larva; and, as the whole has to be rejected or moulted, the membrane of the peduncle of the young Cirripede has necessarily to be formed with a wide and deep inward fold, extending transversely across it; this when stretched open, after the exuviation of the larval carapace and apodemes, necessarily causes the sternal side of the peduncle to be longer than the dorsal, and, as a consequence, gives to the young Cirripede its normal position, at right angles to that of the larva when first attached.
I may here state, that I have examined the larvæ in this the final or perfect stage in four species of Lepas, in Conchodermavirgata, [Ibla quadrivalvis], and, though rather less minutely, in Balanus balanoides, and I find all essential points of organisation similar. With the exception of diversities in the proportional sizes of the different parts, and in the patterns on the carapace, the differences, even in the arrangement of the spines on the limbs and antennæ, are less than I should have anticipated.
I have in this abstract treated the metamorphoses at greater length than I should otherwise have done, on account of the great importance of arriving at a correct homological interpretation of the different parts of the mature animal. In Crustacea, according to the ordinary view, there are twenty-one segments; of these I can recognise in the Cirripede, on evidence as good as can generally be obtained, all with the exception of the four terminal abdominal segments; these do not occur in any species known to me, in any stage of its development. If that part of the larva in front of the mouth, bearing the eyes, the prehensile antennæ, and in an earlier stage two pair of antennæ, be formed, as is admitted in all other Crustacea, of three segments, then beyond a doubt, from the absolute correspondence of every part, and even every coloured mark, the peduncle of the Lepadidæ is likewise thus formed. The peduncle being filled by the branching ovarian tubes is no objection to this view, for I am informed on the high authority of Mr. J. D. Dana,[12] that this is the case with the cephalo-thorax in some true Crustaceans, for instance, in Sapphirina. To proceed, the mouth, formed of mandibles, maxillæ, and outer maxillæ, correspond with the fourth, fifth, and sixth segments of the archetype Crustacean. Posteriorly to the mouth, we come, in the larva, to a rather wide interspace without any apparent articulation or organ, and then to the thorax, formed of six segments, bearing the six pair of limbs, of which the first pair differs slightly from the others. The thorax is succeeded by three small segments, differently shaped, with the posterior one alone bearing appendages; these segments, I cannot doubt, from their appearance alone, and from their apparent function of steering the body, are abdominal segments. If this latter view be correct, the thoracic segments are the six posterior ones of the normal seven segments, and there must be two segments missing between the outer maxillæ and first thoracic pair of legs, which latter on this view springs from the ninth segment. Now, in a very singular Cirripede, already alluded to under the name of Proteolepas, the two missing segments are present, the mouth being actually succeeded by eight segments, and these by the three usual abdominal segments,—every segment in the body being as distinct as in an Annelid: hence in Proteolepas, adding the three segments for the mouth and three for the carapace, we have altogether seventeen segments, which, as I stated, is the full number ever observed in any Cirripede, the four missing ones being abdominal, and, I presume, the four terminal segments. That the cavity in which the thorax is lodged, in the larva and therefore in the mature Cirripede, is simply formed by the backward production of the carapace, does not require any discussion. The valves have no homological signification.
[12] This distinguished naturalist has given his opinion in the ‘American Journal of Science,’ March, 1846, that “the pedicel of Anatifa corresponds to a pair of antennæ in the young;” although the peduncle or pedicel is undoubtedly thus terminated, even in mature individuals, I think it has been shown that it is the whole of the anterior part of the larva in front of the mouth, which is directly converted into the peduncle. Professor E. Forbes, in his Lectures, and Professor Steenstrup, in his ‘Untersuchungen über das vorkommen des Hermaphroditismus in der Natur,’ ch. v, have considered the peduncle as a pair of fused legs. Lovén has taken, judging from a single sentence, the same view of the homologies of the external parts as I have done; in his description of Alepas squalicola, (Ofversigt of Kongl. Vetens., &c., Stockholm, 1844, pp. 192-4), he uses the following words: “Capitis reliquæ partes, ut in Lepadibus semper, in pedunculum mutatæ et involucrum,” &c.; his involucrum is the same as the capitulum of this work.
As we have just seen that the first pair of natatory legs is borne on the ninth segment of the body, so it must be with the first pair of cirri, which consequently correspond to the outer maxillipods (the two inner pair of maxillipods or pied-machoires being here aborted) of the higher Crustacea, and hence their difference from the five posterior pair, which correspond with the five, ordinary pair of ambulatory legs in these same Crustacea. The part of the body, which I have called the prosoma, that is the protuberant, non-articulated, lower part of the thorax ([Pl. IX], [fig. 4 n]), is a special development, either of the ninth segment, bearing the first pair of cirri, or of the segments corresponding with the organs of the mouth. The three abdominal segments of the larva are represented in the mature Cirripede, in the Order containing the Lepadidæ, only by a minute, triangular gusset, let in between the V-shaped tergal arches of the last thoracic segment: in this gusset, small as it is, is seated the anus, and on each side the caudal appendages, often rudimentary and sometimes absent. In another order, I may remark, (including, probably, the Alcippe of Mr. Hancock,) the cirri, of which there are only three pair, are abdominal.
I feel much confidence, that the homologies here given are correct. The cause of their having been generally overlooked arises, I believe, from the peculiar manner, already described, in which the animal, during its last metamorphosis, is internally almost intersected: even for some little time after discovering that the larval antennæ were always embedded in the centre of the surface of attachment, I did not perceive, that this was the anterior end of the whole animal. The accompanying woodcut gives at a glance, a view of the homologies of the external parts: the upper figure (from Milne Edwards) is a Stomapod Crustacean, Leucifer of Vaughan Thompson, and the abdomen, which we know becomes in Cirripedes, after the metamorphosis, rudimentary, and therefore does not fairly enter into the comparison, is given only in faint lines: the lower figure is a mature Lepas, with the antennæ and eyes, which are actually present in the larva, retained and supposed to have gone on growing. All that we externally see of a Cirripede, whether pedunculated or sessile, is the three anterior segments of the head of a Crustacean, with its anterior end permanently cemented to a surface of attachment, and with its posterior end projecting vertically from it.