[18] In ‘Müller’s Archiv,’ 1834, p. 467. I have already several times referred to M. Martin St. Ange’s excellent Memoir, read before the Academy of Sciences, and subsequently, in 1835, published separately.

Female Organs.—M. Martin St. Ange has described how the peduncle[19] is gorged with an inextricable mass of branching ovarian tubes, filled with granular matter and immature ova. In Conchoderma and Alepas, the ovarian tubes run up in a single plane ([Pl. IX], [fig. 3],) between the two folds of corium round the sack. Here the development of the ova can be well followed: a minute point first branches out from one of the tubes; its head then enlarges, like the bud of a tulip on a footstalk; becomes globular; shows traces of dividing, and at last splits into three, four, or five egg-shaped balls, which finally separate as perfect ova. Within the peduncle, the ovarian tubes branch out in all directions, and within the footstalks of the branches (differently from what takes place round the sack), ova are developed, as well as at their ends. Close together, along the rostral (i. e., ventral) edge of the peduncle, two nearly straight, main ovarian tubes or ducts may be detected, which do not give out any branches till about half way down the peduncle, where they subdivide into branches, which inosculate together, and give rise to the mass filling the peduncle, and sometimes, as we have just seen, sending up branches round the sack. These two main unbranched ovarian ducts, followed up the peduncle, are seen to enter the body of the Cirripede (close along side the great double peduncular nerves), and then separating, they sweep in a large curve along each flank of the prosoma, under the superficial muscles, towards the bases of the first pair of cirri; and then rising up, they run into two glandular masses. These latter rest on the upper edge of the stomach, and touch the cæca where such exist; they were thought by Cuvier to be salivary glands. They are of an orange colour, and form two, parallel, gut-formed masses, having, in Conchoderma, a great flexure, and generally dividing at the end near the mouth into a few blunt branches. I was not able to ascertain whether the two main ducts, coming from the peduncle, expanded to envelope them, or what the precise connection was. The state of these two masses varied much; sometimes they were hollow, with only their walls spotted with a few cellular little masses; at other times they contained or rather were formed of, more or less globular or finger-shaped aggregations of pulpy matter; and lastly, the whole consisted of separate pointed little balls, each with a large inner cell, and this again with two or three included granules. These so closely resembled, in general appearance and size, the ovigerms with their germinal vesicles and spots, which I have often seen at the first commencement of the formation of the ova in the ovarian tubes in the peduncle, that I cannot doubt that such is their nature. Hence I conclude, that these two gut-formed masses are the true ovaria. I may add, that several times I have seen in the two long, unbranched ducts, connecting the true ovaria and the ovarian tubes in the peduncle, pellets of orange-coloured cellular matter (i. e., ovigerms) forming at short intervals little enlargements in the ducts, and apparently travelling into the peduncle.

[19] I may here mention, that in all sessile Cirripedes, the ovarian branching tubes lie between the calcareous or membranous basis and the inner basal lining of the sack, and to a certain height upwards round the sack: the true ovaria and the two ducts occupy the same position as in the Lepadidæ.

The structure here described is quite conformable with that which we have seen in the larva; in the latter, two gut-formed masses of equal thickness extended from the cæca of the stomach to within the future peduncle, where the cement-ducts entered them, and where, after a short period, they were seen to expand into a mass of ovarian tubes. In the mature Cirripede, the cement-ducts can still be found united to the ovarian tubes in the middle of peduncle; and the cause of the wide separation of the true ovaria and ovarian tubes, can be simply accounted for by the internal, almost complete intersection of the animal, which takes place during the last metamorphosis.

The ova, when excluded, remain in the sack of the animal until the larvæ are hatched; they are very numerous, and generally form two concave, nearly circular, leaves, which I have called after Steenstrup and other authors, the ovigerous lamellæ ([Pl. IV], [fig. 2 b]). These lamellæ lie low down on each side of the sack: in [Conchoderma virgata], however, there is often only a single lamella, forming a deeply concave cup: in C. aurita there are generally on each side four lamellæ, one under the other. The ova lie in a layer from two to four deep; and all are held together by a most delicate transparent membrane, which separately enfolds each ovum: this membrane is often thicker and stronger round the margins of the lamellæ, where they are united, in a peculiar manner, presently to be described, to a fold of skin, on each side of the sack: these two folds, I have called the ovigerous fræna ([Pl. IV], [fig. 2 f]).

M. Martin St. Ange, describes an orifice under the carina, by which he supposes the ova to enter the sack; this, after repeated and most careful examinations, I venture to affirm does not exist; on the contrary, I have every reason to believe that the ova enter the sack in the following curious manner. Immediately before one of the periods of exuviation, the ova burst forth from the the ovarian tubes in the peduncle and round the sack, and, carried along the open circulatory channels, are collected (by means unknown to me) beneath the chitine-tunic of the sack, in the corium, which is at this period remarkably spongy and full of cavities. The corium then forms or rather (as I believe) resolves itself into the very delicate membrane separately enveloping each ovum, and uniting them together into two lamellæ; the corium having thus far retreated, then forms under the lamellæ the chitine-tunic of the sack, which will of course be of larger size than the last-formed one, now immediately to be moulted with the other integuments of the body. As soon as this exuviation is effected, the tender ova, united into two lamellæ, and adhering, as yet, to the bottom of the sack, are exposed: as the membranes harden, the lamellæ become detached from the bottom of the sack, and are attached to the ovigerous fræna. To demonstrate this view, an individual should have been found, with both the old and new chitine tunic of the sack, and with the lamellæ lying between them; this, I believe, I have seen, but it was before I understood the full importance of the fact: a great number of specimens would have to be examined in order to succeed again, for the changes connected with exuviation supervene very quickly. I have, however, several times found the ova so loose under the sack, as to be detached with a touch from the ovarian tubes; and I have twice carefully examined specimens, which had just moulted, as shown by even the mandibles being flexible, in which the lamellæ had not become united to the fræna, but still adhered to the newly-formed chitine tunic of the sack; in these, the ova were so tender, that they broke into pieces rather than be separated from the membrane of the lamella, itself hardly perfectly developed, for pulpy cellular matter adhered outside some of the ova. These and other facts are quite inexplicable on any other view than that advanced.

As the lamellæ are formed without organic union with the parent, they would be liable to be washed out of the widely open sack of the Lepadidæ, if they had not been specially attached to the fræna. These fræna consist of a pair of more or less semicircular folds of skin, depending inside the sack, on each side of the point of attachment of the body. The fræna are often of considerable size, but in Ibla, they are very minute; they are formed of chitine tunic with underlying corium, like the rest of the sack; on their crests, there is a row, or a set of circular groups, or a broad surface, covered, either with minute, pointed, bead-like bodies mounted on long hair-like footstalks, or with staff-formed bodies on very short footstalks. I measured some of the bead-like bodies, in [Lepas anserifera], and they were 1/2000th of an inch in diameter, and the footstalks three or four times as long as the elongated heads. These heads, of whatever shape they may be, have an opaque, and, I believe, glandular centre; I could not make out with certainty an aperture at their ends, but, I believe, such exists, and they seem to secrete a substance, which hardens into a strong membrane, serving to unite the crest of the frænum to the edges of the lamellæ. In one case, this bit of membrane seemed formed of a woven mass of threads. These little glandular bodies, with the membrane formed by them, are cast off at each exuviation, and new glands formed on the crest of the frænum underneath. In some species of Pollicipes, (viz., P. cornucopia and elegans,) the fræna, though present and large, are functionless and destitute of the glands: I believe, they exist in this same functionless condition, and in rather a different position in the sessile Cirripedes, and that in this family they serve as Branchiæ.

The above-described method by which Cirripedia lay their eggs, namely, united together in a common membrane, placed between their old outer and new inner integuments, and the manner in which the lamellæ, when thus formed, are retained for a time fastened to the fræna, and are then cast off, appears to me very curious. In some of the lower Crustacea, it is known, that the ova escape by rupturing the ovisacs formed by the protruded ovarian tubes, and this is the nearest analogy with which I am acquainted. The ova are impregnated (as I infer from the state of the vesiculæ seminales), when first brought into the sack, and whilst the membrane of the lamellæ is very tender: the long probosciformed penis seems well adapted for this end. In the male of [Ibla Cumingii], which has not a probosciformed penis, the whole flexible body, probably, performs the function of the penis: in [Scalpellum ornatum], however, the spermatozoa must be brought in by the action of the cirri, or of the currents produced by them. That cross impregnation may and sometimes does take place, I infer from the singular case of an individual, in a group of Balani, in which the penis had been cut off, and had healed without any perforation; notwithstanding which fact, larvæ were included in the ova.

Exuviation; Rate of Growth; Size.—I have had occasion repeatedly to allude to the exuviation of the Lepadidæ: with the exception of the genus Lithotrya,[20] in which the calcareous scales on the peduncle, together with the membrane connecting them, is cast off, neither the valves nor the membrane uniting them, nor that forming the peduncle with its scales and styles, are moulted; but the surface gradually disintegrates and is removed, perhaps sometimes in flakes, whilst new and larger layers are formed beneath. In Scalpellum, I ascertained that the new membrane, connecting together the newly-formed calcified rims under the valves of the capitulum, was formed as a fold, with the articulated spines which it bears, all adpressed in certain definite directions. This fold of new membrane, when the old membrane splits and yields, of course expands, and thus the size of the capitulum is increased. In the peduncle, lines of splitting can seldom be perceived, except, indeed, in the sub-globular, embedded, downward-growing peduncle of Anelasma, as described under that genus. I do not understand what determines the complicated lines of splitting of the old membrane between the several valves of the capitulum,—without it be simply, that along these lines alone, the old membrane is not strengthened by the new membrane being closely applied under it, the new being formed, as we have just said, in a fold, in order to allow of increase in size. Although, as I believe, there is strictly no exuviation in the outer membranes of mature Lepadidæ, it seems that narrow strips of membrane are cast off from between the valves, for the few first moults, after the final metamorphosis of the larva. I may here remark that, in most sessile Cirripedes, the outside membrane connecting the operculum and shell, is regularly moulted.

[20] The external integuments being moulted in Crustacea, but not in the Cirripedia, may appear, at first, an important difference: but we here see that non-exuviation is not universal amongst the Lepadidæ, and, on the other hand, according to M. Joly, (‘Annales des Sciences Naturelles,’ 2d series, Zoolog.), there is one true crustacean, the Isaura cycladoides, which has a persistent bivalve shell.