It has been seen that I divide the Cirripedia into three orders,—the [Thoracica], [Abdominalia], and [Apoda]; between which the fundamental difference consists in the limbs or cirri being thoracic in the first, abdominal in the second; and entirely absent in the third. For the sake of showing the range of character in Cirripedes, to which allusion has been made, I will briefly indicate the leading differences in each order. In the [Thoracica], three families are included,—the [Balanidæ], or sessile Cirripedes, the [Verrucidæ], remarkable from their quite asymmetrical shell, and the Lepadidæ, or pedunculated Cirripedes. The great difference in external appearance between these three families is known to all naturalists. Even within the one family of Lepadidæ there are great differences in external appearance, as will be admitted on comparison of Lepas, Pollicipes, Conchoderma, &c.; but we have also important internal differences, as in the case of Anelasma, in which the cirri are barely articulated, and are not capable of seizing prey, whilst the mouth is almost probosciformed, with the outer maxillæ and palpi rudimentary: still more important are the differences in [Alcippe], in which the cirri of the first pair act as brushes; the second, third, and fourth pairs being quite aborted; and the fifth and sixth pairs consist only of four segments, with one of the two normal rami converted into a crenated, button-like projection, for the sake apparently of triturating food; [Alcippe], also, is very remarkable in being destitute of a rectum and anus. In this same genus [Alcippe], in Ibla and Scalpellum, there are either separate males or Complemental males, some of which are so utterly abnormal in their characters, that by no definition which I could frame, could they be included even in their proper Order, much less in their proper Family.
In the second order of [Abdominalia] (Pl. [23] and [24]) the seventh or last cephalic segment is quite distinct, and bears rudimentary organs, answering to the first pair of maxillipeds of ordinary Crustaceans, of which organs, and of the segment supporting them, there is no trace in the [Thoracica]: the seven succeeding thoracic segments are destitute of any appendages; but the three segments of the abdomen bear three pairs of cirri. The mouth is peculiar in the labrum being developed into very large, moveable, lancet-formed organ; and the lower end of the œsophagus is armed with beautiful discs of teeth, and brushes of hairs,—a structure confined to this order. The male resembles the male of [Alcippe]; and the latter genus seems to be the connecting link between the [Thoracica] and [Abdominalia]. But the most important character of this latter order, in which it differs from [Alcippe], and all other known Cirripedes, is in its metamorphoses; all the first changes are merely indicated by changes in form in an egg-like larva, without the development of distinct organs; and the last, or pupal condition, which is attained within the sack of the parent, is very peculiar, by the entire absence of natatory limbs.
The third order of [Apoda] is the most peculiar of all; it contains, like the last, only one known species: the most acute naturalist, I am convinced, if he had not made the class his special study, would never even have suspected that this animal was a Cirripede. We see much magnified in Pl. [25], fig. [7] a naked, plainly-articulated animal, resembling the larva or maggot of a fly, attached by two threads; and these threads, on analysis, can be clearly shown to be the last rudiment of the carapace, specially modified. The last cephalic, the seven thoracic, and the three abdominal segments, are all equally destitute of appendages. The mouth is suctorial, and constructed on a plan unlike, I believe, anything known in the articulate kingdom; for the mandibles and maxillæ have rotated on their axes, and stand back to back; they can act only by tearing open a slit, and this action is performed in a hood, formed by the confluence of the broad palpi and labrum. Although the œsophagus is distinct, there is no stomach or anus. Lastly, owing to there being no carapace, the ova are developed, differently from in all other Cirripedes, within the thorax.
I will close this preliminary discussion on the confines and type of the sub-class, by recalling attention, now that a sketch has been given of the three Orders, to the remark before made, that a wide range of structure is included within it, and by reurging that the Cirripedia should be ranked, not as one of the subordinate groups, but as one of the main divisions of the Crustacea.
On the Sexual Relation of Cirripedes.
Cirripedes are commonly bisexual or hermaphrodite, but in Ibla, Scalpellum, and [Alcippe], members of the Lepadidæ in the order [Thoracica], and in [Cryptophialus] in the order [Abdominalia], the sexes are separate. As two of these genera were described in my former volume, and two others ([Alcippe] and [Cryptophialus]) are described in this volume, I may as well here give a brief summary of the facts as yet known on this very curious subject. The Males, in the above four genera, present a wonderful range of structure; they are attached in the usual way by cement proceeding from the not-moulted antennæ of the pupa, to different parts, in the different species, of the female. These males are minute, often exceedingly minute, and consequently generally more than one is attached to a single female; and I have seen as many as fourteen adhering on one female! In several species the males are short-lived, for they cannot feed, being destitute of a mouth and stomach. As the females are longer lived, successive crops of males, at each period of propagation, become attached to her. It is the females in the above genera which retain the characters of the genus, family, and order to which they belong; the males often departing widely from the normal type. Some of the males are rudimentary to a degree, which I believe can hardly be equalled in the whole animal kingdom; they may, in fact, be said to exist as mere bags of spermatozoa. So widely do some of them depart in every character from their class, that twice it has happened to me to examine specimens with a little care, and not even to suspect, until a long period afterwards, that these males were Cirripedes.[18]
[18] In my volume on the Lepadidæ (p. 200) in searching for analogies for the permanently epizoic and rudimentary condition of the male Cirripedes, I quoted two cases, which I believe are now known not to be analogous; namely, the Syngamus trachealis of Von Siebold, and the worm-like Hectocotyle, which latter was quite lately supposed to be a male Cephalopod, but has now been ascertained to be only one of the arms of the male wonderfully adapted and organised as a sperm-receptacle. The Asplanchna, the mouthless male of a Rotifer, (p. 292) alone remains for me.
In Scalpellum Peronii, and villosum, the males are but little abnormal, for if classified independently of their sexual relations, they would be considered as immature specimens of a new genus, standing next to Scalpellum; in Scalpellum rostratum, the male would form another and rather more distinct genus. The males, in the latter, are attached to the other sex, between the basal edge of the labrum and the adductor scutorum muscle; but in S. Peronii and villosum they are attached lower down, in the furrow between the two scuta, and are thus protected: in these three species, the internal parts of the male present nothing particular. In Ibla, the males are attached low down within the sack of the female; they may be said to consist of a mouth surmounted on a long peduncle, for there is no capitulum or general covering, and the whole thorax is in a rudimentary condition, the cirri being reduced to two distorted pairs. As these males certainly moult several times and grow a little, they must feed; and as they have no cirri fit for action, they must seize their food by the contortions of their peduncle, which we know homologically consists of the three first segments of the head. The movements of the peduncle must, also, supply those of the probosciformed penis, almost invariably present with other Cirripedes, but here absent. If compelled to class these males without regard to the female, great difficulty would be experienced; we could hardly place in the family of the Lepadidæ, a Cirripede without a capitulum, and without cirri, those very organs which give their name to the class, and with a thorax reduced to the dimensions of a lower lip; yet, if the presence of a peduncle did determine the classifier to place these males amongst the Lepadidæ, then undoubtedly the character of the mouth, &c. would fix their position next to Ibla.