MALE. Pl. 23.

On every specimen of the female Alcippe, which I carefully examined, I found some minute parasites (or epizoons) attached to the lateral edges of the upper part of the horny disc, and therefore lying within the narrow end of the fissure leading into the chamber excavated in the shell of the Buccinum. Although having had some experience in the very anomalous forms which male cirripedes assume, yet when I first casually inspected these parasites under a weak lens, from their transparency, their elongated and lobed body, including an internal folded up organ, I actually threw them away, thinking that they were probably Bryozoa. Subsequently, a more careful inspection immediately showed the cemented prehensile antennæ, and their cirripedial nature was demonstrated. I soon found specimens with the perfect still adherent exuviæ of the locomotive pupa, undistinguishable from the pupa already described as probably belonging to the female Alcippe. But as this latter fact, may perhaps be doubted, I must show that there is other evidence sufficient to prove that these cirripedial parasites are the males of the female Alcippe. Of the females, I inspected many specimens, and all certainly were without external male organs; and in the four or five specimens which I rigidly examined, there were no testes or vesiculæ seminales, the latter being in all hermaphrodite and male cirripedes so conspicuous. On the other hand, I examined at least thirty specimens of the parasite, and they were all exclusively males, for all had a probosciformed penis, and the greater number had their vesiculæ seminales filled with spermatozoa, and hence were ready to perform the act of impregnation, but undoubtedly they contained no ova. It would, then, be very strange, if these two cirripedes of opposite sexes, thus attached together, were not sexually related. Wonderfully different as the parasite is from the female Alcippe, yet, in one very important character it is related to Alcippe, and to no other member of the Family, namely, in the sack extending down to the extreme lower point of the peduncle; the male organs, I may add, occupying an analogous position with the peculiar position of the female organs in Alcippe. The lateral lobes of the peduncle in the parasite seem to represent the sides of the broad depressed peduncle in Alcippe; and in both the peduncle grows at its lower end—a very rare circumstance—observed only in two genera in this Family, namely, in Anelasma, and in a slight degree in Lithotrya. Besides these points of resemblance between Alcippe and its parasite, which are striking, considering their external utter dissemblance, the affinities of both point, judging from certain small characters, in the same direction, namely, towards Ibla and Alepas. Finally, then, I think, we may confidently admit that this parasite or epizoon is the male of the female Alcippe: indeed, considering the facts given in my former volume, on Ibla and Scalpellum, I have, perhaps, here discussed the question at unnecessary length.

The males are generally attached, as already stated, to the two hollowed out sides of the upward prolongation of the horny disc; they adhere by means of little patches of cement, proceeding from the terminal segments of their antennæ, to the overlapping edges of the few later-formed zones of the disc; hence, they lie protected, within the narrow end and a little under the edges of the fissure leading into the cavity in which the female is lodged. In some specimens, however, the males are attached rather lower down on the disc, and are not confined exclusively to its upper margin, so that they live fairly under the roof of shell which covers the main part of the disc: but they are never attached very low down, so as to lie far from the lower end of the orifice leading into the sack of the female. I have two or three times seen as many as three males on each side, but sometimes there is only one on each side, or none on one side. A large distorted specimen actually had twelve males, and two pupæ on the point of undergoing their final metamorphosis, all fourteen attached on one side, and all evidently must have been alive together! Another specimen had nearly the same number, a few on one side, and the rest on the other side.

The male immediately, after the exuviation of the pupal carapace, 25/1000th of an inch in length, is only 23/1000th of an inch long, but ultimately it becomes, chiefly from the growth of the lower end of the peduncle, nearly twice this length; for the largest specimen which I have seen, that figured, was 45/1000th of an inch long (i. e. under 1/20th of an inch), and 1/100th of an inch in breadth across the peduncle, beneath the lateral lobes. The whole external membrane of the animal (as well as the internal membrane of the sack), is very thin, quite structureless, and as transparent as glass; hence, even the spermatozoa, within the vesicula seminalis, can be seen from the outside. The whole structure of the animal is very simple. The ventral surface can be at once recognised by the attachment of the antennæ (fig. 19, a), and these organs mark the point which was the anterior end of the male, just at the period of its metamorphosis, and before the lower end of the peduncle had grown. These antennæ have already been fully described; they are conspicuous from being composed of membrane, rather thicker than that investing the body of the male, and which external membrane can be traced entering these organs, and appearing like cement-ducts; but within these tubular prolongations of the outer membrane, I could obscurely see the real cement-ducts.

The part answering to the capitulum is much flattened and elongated; it widens but little from the upper to the lower end, where it blends with the carinal or dorsal surface (the under surface in fig. 19) of the lobed peduncle. At the upper end there is a small orifice, and close to this, on the ventral or rostral side, there is a thin, apparently double projection (i, fig. 19) or flap of membrane, one flap lying exactly over the other. The whole length of this capitulum probably corresponds with that small portion of the capitulum in the female, between the upward prolongation of the horny disc and the lower end of the orifice; and the two broad flattened projections in the male, probably answer to the two sharp narrow points (a, fig. 1, Pl. 22) in the female. The peduncle has two lateral lobes (h, g, fig. 19), and, whilst young, what may be called a third and medial lobe, but this soon increases largely by growth, and forms the main part of the peduncle. The lateral lobes are intimately connected with the ventral surface; they tend to lie in a plane, at right angles to the compressed capitulum, but owing to the excessive thinness and flexibility of the whole external membrane, it is difficult to ascertain the relative position of the different parts. Moreover, owing to the pupa being so much flattened, these lobes are necessarily formed folded up; and, I believe, it depends on the position, with respect to surrounding objects, which the male ultimately holds, whether the lobes ever assume, their apparently normal position, in a plane at right angles to the sides of the pupa; owing, also, to the form of the pupa, the two lobes seem generally to be actually formed of unequal sizes, that formed in the dorsal region of the pupa being the largest. I believe that these lobes correspond with the lateral margins of the upper end of the peduncle of the female, which margins project laterally beyond the sides of the capitulum. The lower lobe, or end of the peduncle, is depressed in the same plane with the lobes; it is of variable length; when first formed it hardly extends beyond the basal articulation of the prehensile antennæ. Commonly it does not lie quite in a straight line of the capitulum; and I have seen specimens in which it stood at nearly right angles to the capitulum and to what was the ventral surface of the pupa; this irregularity in the relative position and sizes of the different parts of the peduncle, no doubt, to a considerable extent, depends on the form of surface to which the male becomes attached, just in the same way as we have seen that the peduncle of the female becomes altered in shape during the excavation of the chamber in which it is lodged.

I feel some difficulty on one point: in the pupa the single eye of the future male can be clearly distinguished, and it lies some way from the anterior end of the body; but in two males, which certainly had just moulted, and in which none of the internal organs were as yet developed, the eye lay close to the anterior end, directly over the basal articulation of the antennæ. I suspect this is somehow caused by the great change of form which supervenes, during the metamorphosis, at this anterior end of the body; the extremely compressed body of the pupa having to become depressed and lobed in the young male. I have given a figure of a young male, just as it appeared (Pl. 23, fig. 18), somewhat distorted from lying on a flat surface; c, being the eye.

The sack extends, in a very remarkable manner, down to the lower end of the peduncle, the whole inside of the animal being thus freely open to the water. In the upper part, the sack forms a mere narrow tube; it does not appear to have been formed in the same manner as in all other cirripedes, namely, surrounding the thorax and natatory legs of the pupa, but in an abnormal position, along the dorsal surface, above the sack and thorax of the pupa: a transparent line, where the new narrow sack is in process of formation, is the first indication of the coming metamorphosis. The sack in the capitulum of the male is not central, but lies near the dorsal surface; the ventral interspace, between the outside and the sack, is occupied by oblique fibres (l, fig. 19), which may be striæ-less muscles, but I suspect are ligamentous fibres, giving support to the whole projecting capitulum. These fibres enter a little way within the lobed peduncle; they are probably homologous with the strong muscles, which run from beneath the upper end of the horny disc of the female to the lower end of the orifice leading into the sack. Round the lobed peduncle,[148] there are two bands (e, f) of thin muscular fasciæ, slightly oblique to each other, and attached at the ends to the outer membrane; they are evidently homologous with the external transverse muscles, which are best developed round the same part in the female. Some of these muscles present a singular chain-like appearance, from being strangled at intervals: they act probably in aiding the long probosciformed penis to protrude itself out of the sack. I could not detect any longitudinal muscles, and the lower part of the peduncle seems destitute of muscles of any kind.

The internal structure of the animal is very simple. Within the lower end of the peduncle there is a dark purple eye (c), under the 1/1000th of an inch in diameter, a testis (d) and a (b) vesicula seminalis. These organs falsely appear as if suspended in the middle of the peduncle, but they are really attached, I believe within a separate partition, to the ventral surface, occupying the same position as the mass of ovarian cæca in the female. The eye lies on the line of junction between the testis and the vesicula seminalis, and on their ventral side. The testis is rounded, and consists of a mass of cells, on an average 1/5000th of an inch in diameter. The vesicula seminalis varies extremely in condition, being either a mere rather broad vessel, enlarged where it joins the testis, or a bag fully as large as the testis itself, and distended with spermatozoa, all arranged parallel to its longer axis. There was an evident relation between the size of the vesicula seminalis and that of the testis, the number of the cells in the latter decreasing as the mass of the spermatozoa increased: there was also an evident relation between the age of the male and the state of these organs; younger and more opaque individuals, having their testes of large size; and older specimens, with the lower end of the peduncle arrived at its full dimensions, having the vesicula distended. Some few old specimens had evidently discharged their spermatozoa. By dissection I more than once distinctly traced the vesicula seminalis entering the broad lower end of the penis. The membrane, forming the vesicula, is ringed, and I presume is, as in other cirripedes, contractile, so as to expel the spermatozoa. The probosciformed penis (m) is of extraordinary length: it is plainly ringed, or rather articulated, in this respect resembling that organ in Ibla and Alepas; it tapers gradually, and terminates (as usual) with a brush of fine bristles; it is furnished with delicate voluntary muscles, arising from the body round its basis, and extending no doubt up to the apex, but too fine to be traced all the way. Its broad lower end is attached in a slight depression, on the ventral side of the sack, a little above the point of attachment of the pupal antennæ. According to all analogy, the spot whence the penis springs must be considered as representing the thorax and abdomen; and the outer membrane of the penis is here, as on this view it should be, reflexed and is continuous with that lining the sack. Ordinarily the penis lies coiled up in complicated folds, appearing like a large intestinal worm, and fills the lobed part of the peduncle, which apparently serves for no other purpose than its reception. In one case in which I dissected out the penis, I found it in its contracted state; 41/1000th of an inch in length, equal to that of the entire capitulum and peduncle; in a specimen, in which the penis had been naturally exserted, the part which protruded (m) was by itself rather longer than the whole animal; and as this specimen had been placed in spirits of wine, the organ no doubt was contracted; hence I think it probable that the probosciformed penis, when fully stretched out, would equal twice the length of the entire animal.

There must be a nervous system; and there must likewise be a gland (homologous with the ovaria) for secreting the cement; but I could not distinguish parts so small. Certainly there is no mouth, or stomach, or thorax, or limbs of any kind, or abdomen.

It is obvious that these males must be very short-lived: they perform their masculine functions and then perish. We have seen, however, that after the act of metamorphosis they do grow a little, and I have reason to suspect that this is effected, as with other Cirripedes, by moulting. The growth must be absolutely dependent on the store of nutriment laid up within the pupa. The young male, immediately after the exuviation of the integuments, thorax, natatory legs, abdomen, and eyes of the pupa, consists of a pulpy cellular mass, without any internal organs as yet formed.

Judging from the different sizes of the females which included perfectly developed ova, I infer that they must breed more than once during their lives; and therefore, that successive sets of males, as in the genus Scalpellum, must become attached to them. I was not, however, able to discover the prehensile antennæ or other remains of the old males adherent to the females; a circumstance which I presume is accounted for by their attachment being weak. Considering the very small size of the male, it is not surprising that so many,—in one case fourteen,—are required to impregnate the numerous ova of a single female. How the males know the proper period when the ova, lying in a sheet at the very base of the sack of the female, are ready for impregnation, I cannot say, without it be that they perceive the moulting of the external membrane, close to the edge of which they are attached; for this moulting would indicate the period when the ovigerous lamella came to the surface of the sack, and the ova would then be soon ready for impregnation. From the position in which the males are attached, and from the extraordinary length of the probosciformed penis, capable of voluntary movements, I have no doubt the males can insert the tip of this organ within the lower edge of the orifice of the sack, and there discharge the spermatozoa, which, by their own movements, must pass down the sides of the sack of the female till they reach their proper destination. The position of the males, with respect to the female’s body, is almost exactly the same as that occupied by the complemental males of Scalpellum Peronii and villosum; the lower and narrow end of the fissure, worn in the gasteropod shell, here affording that protection to the males, which the edges of the opposed scuta afford to the complemental males of the above two species of Scalpellum. We cannot doubt that these latter males aid in the impregnation of the ova of the hermaphrodites, but they are not furnished with a very long penis, probably for the very reason that they are complemental males, and therefore not so absolutely necessary for the impregnation of the ova as are the males of Alcippe.

I have, in my former volume, expressed my astonishment at the extent to which abortion had been carried in the male Ibla; but it has been carried much further in the male Alcippe. In Ibla, the thorax is reduced to a mere flap, and only two pairs of cirri exist in a most useless and rudimentary state, but there is a well organised mouth, stomach, and anus. In the males of Scalpellum vulgare, ornatum, and rutilum there is no mouth or stomach, but there is a thorax with four pairs of minute, modified cirri, and a large abdominal lobe. Here, in the male Alcippe, all these negatives are united, we have no mouth, no stomach, no thorax, no cirri, no abdomen! The archetype crustacean consists of twenty-one segments; of these the seventeen anterior segments can be clearly made out in the archetype Cirripede: now, in the male Alcippe, the first three segments are largely developed, forming all that is externally visible, but the remaining fourteen segments are absolutely aborted, but in idea may be considered as forming the membranous depression whence the probosciformed penis springs; for this organ normally arises at the extremity of the seventeenth segment. To show the wonderful diversity of nature, even in the same sub-class, I may be permitted to remark, that whilst in Alcippe only the three anterior segments are developed, the fourteen succeeding segments being rudimentary, in Proteolepas (hereafter to be described) these fourteen segments are all largely developed, whilst the three anterior segments are quite aborted, being represented only by a thin envelope to the two threads by which this Cirripede is attached to the supporting object.[149]