FEMALE. Pl. 22.

I may premise that after the sketch of the leading peculiarities of Alcippe, and after the discussion on its affinities, just given under the Family, I think it would be superfluous to institute a full generic description, separately from the following detailed account of this most anomalous cirripede.

General Appearance.—The whole animal is from .2 to .3 of an inch in length, of a soft texture, colourless or yellowish, and lives concealed in a cavity of its own formation in the shells of certain Gasteropods. This cavity communicates with the water by a narrow fissure-like orifice (Pl. 22, fig. 4), broadest at the posterior end, where the cirri are exserted; narrow, closed, and generally curved at the other (a) end: the two sides of the fissure (b) are commonly bordered by a calcareous inorganic deposit: the walls of the cavity are worn so thin over the peduncle, at the narrow end of the fissure, that the orange-coloured ovaria can generally be seen through the shell of the mollusc, and hence there is here a distinct fan-shaped stain (fig. 3) on the surface. The animal consists of a compressed capitulum, without valves, and of a sort of peduncle depressed on its rostral face, and covered with a broad, oval, thin, horny disc. We must remember that in the Lepadidæ the peduncle does not essentially differ from the capitulum, being only the flexible lower or anterior end of the animal, and is separated from the capitulum only by shape, and generally by the direction of the lines of growth. The disc, when most regular (fig. 1, H), lies in a plane at right angles to the sides of the capitulum, and almost in a line with the orifice leading into the sack; but the peduncle is often very irregular (fig. 2), and the disc comes even to occupy a position nearly parallel to one or the other side of the capitulum. On the carinal side, the capitulum is generally separated from the peduncle by a rather deep fold (f, in the section fig. 5), but this depends in some degree upon the state of distension of the mass of ovarian cæca. I have given a drawing, fig. 1 (partly taken from Mr. Hancock), of a very regular individual, and of an extremely distorted specimen (fig. 2). The distortion, I believe, is generally caused by the animal, during its excavation, breaking into some old cavity.

External Structure.—The orifice leading into the sack is about one third of the total length of the animal: its edges or lips are thickened, horny, and brownish: at the lower end, exactly where the orifice ends, the lips are formed, from being deeply notched, into two sharp projections (a, figs. 1, 5, 6), unlike anything occurring in any other Cirripede. The external membrane (c, fig. 6) of the lip supports an irregular but nearly straight band of sharp, thick spines of chitine, about 1/1000th of an inch in length, together with a few hairs: at the carinal or upper end of the orifice the spines are largest and most numerous; at the other and lower end, they decrease in size; and on the two projections (a, fig. 6), and on the adjoining parts of the external membrane, they graduate into the small dentated points which cover the whole surface of the animal. The inner tunic of the sack (b), on each side along the upper half of the orifice, is remarkable from having a moderately broad, curved band of short, sharp spines, not quite so thick as those on the external surface, closely adpressed together and pointing upwards, like the javelins of an ancient phalanx, thus probably preventing the ingress of any intruding animal. This band of spines curves at the upper end, conformably with the shape of the orifice. The inner tunic of the sack in this upper part is yellowish, and, what is very unusual, is thicker than the external membrane. A little way down, within the orifice, and more especially in front of an elegant row of hairs on the two sides of the great labrum, there is a band of very fine but stiff hairs (5/1000ths of an inch in length), pointing upwards, and making together with those on the labrum a hedge, barring ingress into the sack.

The external membrane over the whole animal, excepting the horny disc which covers the rostral face of the peduncle, is very thin and transparent; it is periodically and often moulted, as may be inferred from the many old lines of junction round the edges of the horny disc: it is irregularly and pretty thickly (but not so thickly as in fig. 7) studded with star-headed, minute points, from 2 to 5/10,000ths of an inch in diameter, composed of hard chitine, seated on a short footstall, and this on a circular, yellowish, slightly thickened disc of the general investing membrane, appearing like a halo surrounding each little point. These points are directed obliquely upwards. There are none on the horny disc, though particularly numerous close to its margin. Their state varies much: just after a moult, when newly formed, the spines are regularly star-headed, with quite sharp rays, from two to six in number, with some of them occasionally bifid; but these points or rays soon become blunted, and ultimately half the star is worn away, so that the appearance then presented is that of a crescent with a few blunt points on its convex side. At each exuviation, the thickened membrane of the orifice with its strong external spines (the condition of which also varies according to the period elapsed since the last moult), and of course the whole internal tunic of the sack, with its spines and hairs, are all moulted, together with the external membrane and the little star-shaped points. In most specimens a barely distinguishable band or bar of yellowish, slightly thickened membrane, runs from the points (a), at the lower end of the orifice, for some way obliquely downwards; and at the lower end of this bar the weak adductor scutorum muscle (having transverse striæ) is attached. This bar is often strengthened by a prominent external fold of membrane, but yet it is so flexible, and as it is united only to the lower end of the orifice, I can hardly believe that it can, by means of the adductor muscle attached to its opposite extremity, have much power in closing the orifice. I believe that this muscle acts simply in narrowing the whole animal, so as to favour its movement within the cavity in which it is imbedded. Owing to this position of the adductor muscle, and its consequent little power in closing the orifice, we can understand the necessity for the defence afforded by the bands of spines and hairs on the inner tunic of the sack and on the labrum, which do not occur in other Cirripedes.

Horny disc.—The general shape of the disc, its irregularity and position, have been already described. It never extends, as remarked by Mr. Hancock, to the extreme lower point of the peduncle; upwards it reaches to a little below the lower end of the orifice. It consists of successive layers of membrane, either moderately thick and opaque, or only a little thicker than the general membrane of the body, but never furnished with the little sharp points; it increases in size, in like manner as the calcareous valves of other Cirripedes, the undermost and last formed layer extending beyond the others, with its edge united, till the next exuviation, to the general membrane of the body. The disc is attached, at its upper end, apparently in the usual way, by cement, to the roof of the cavity of the shell in which it is imbedded; but the lower parts of the disc are also slightly and partially attached, chiefly along the lines of growth or exuviation; and this, I suspect, is effected by an inorganic calcareous deposit; anyhow I could not perceive here any cement or cement-ducts. Beyond the circumference of the disc the whole animal lies free in its cavity. The lines of growth in the middle part of the disc are generally obliterated by the decay of the older and outer layers. These lines, though of course ordinarily conformable with the general outline of the disc, are not always so, for the disc sometimes becomes during growth slightly changed in form, and the animal, consequently, slightly changed in position; sometimes either one or the other side or the upper end of the disc is left deserted by the new layers of the growing disc; these being formed on the deserted side of less size or extension, instead of larger size, as they normally should be all round the disc.

The upper end of the disc is always produced into a projection of not regular shape, but generally hollowed out or embayed in front (fig. 1), and almost always hollowed out on the two sides. This projection stands directly over the adductor muscle (b in fig. 5), and on the exterior surface is generally convex, being concave on the under side for the attachment of several muscles presently to be described. The horny layers are in this part usually thicker than elsewhere. The disc is thus upwardly produced, owing apparently to the fissure which leads into the cavity of the shell of the mollusc becoming, during the process of excavation, considerably longer than is necessary,—that is longer than the orifice leading into the sack; and consequently, for the protection of the imbedded animal, the lower and narrow end of the fissure is closed on its under side by this upward production of the horny disc, formed of layers of membrane of unusual thickness. A deposition, also, of lime, hereafter to be mentioned, gives further protection.

In the bays on each side of the upward production of the horny disc, and likewise a little lower down on its edges, and therefore somewhat protected by lying within the narrow, pointed, lower end of the fissure in the shell of the mollusc, the short-lived Males (Pl. 22, fig. 1, m) are attached often in groups of two, three, or more together.

It may be asked to what part, in other Cirripedes, does the horny disc answer? Not considering the upward prolongation, which has been developed for a special purpose, the disc is irregularly circular,—is added to all round,—serves for the attachment of the whole animal to the supporting surface,—is covered on the under surface by a conformable and parallel mass of ovarian cæca, and the latter by the inner tunic of the sack; therefore in every character, and in its relation to the other parts of the animal, the disc answers to the end of the peduncle, or to the basal cup in Lithotrya, or still more closely to the basis in sessile cirripedes, with the important exception that it lies in a line with the longitudinal axis of the whole animal instead of at right angles to this axis. We know that all ordinary cirripedes become first permanently attached in their pupal state by their antennæ, which are seated on the ventral or rostral surface, near to the anterior end of the body; and that from the young cirripede, after the act of metamorphosis, being turned vertically upwards, and from the extreme anterior, now lower, end of the body not being rapidly developed, the surface cemented down, or the basis, encroaches almost equally on the dorsal, lateral, and ventral surfaces. But if we were to suppose the extreme anterior point of the body to be rapidly developed, the surface of attachment or basis, without it grew still more rapidly, could not possibly reach the dorsal surface, and would, consequently, be confined to the ventral or rostral surface. I have not seen the young of the ordinary or female Alcippe soon after its metamorphosis, but in the male the development of the extreme anterior end of the body is extraordinarily rapid, and from analogy we may fairly conclude that this is likewise the case with the female. Hence I believe that the horny disc answers to the cemented down, lower end of the peduncle, in other members of the Lepadidæ, and to the basis in the Balanidæ, and that it is confined to the ventral or rostral surface, owing to the anterior or lower end of the body having been rapidly developed. To make all the parts, internal and external, of Alcippe, correspond with those of other cirripedes, the main circular part of the horny disc must be turned up at nearly right angles to its present position (the dorsal or carinal integuments, to the right-hand in fig. 5, being shortened), and then we should have a peduncle, certainly very short and broad, but holding its proper relative position.[143]

Sack and its Muscles.—I have already described the curious phalanx of spines, the long fine hairs, and thickened condition of the inner tunic of the sack along the sides of the orifice. This inner tunic is a reflexion from that enveloping the body of the animal, in the usual manner, as may be seen in the section (Pl. 22, fig. 5). Between the external membrane and the inner tunic of the sack (e), there is of course the usual double fold of corium, these two folds being united by minute, transverse, ligamentous fibres, branched at the two ends, as in other Lepadidæ. Imbedded in the corium there are numerous, longitudinal, striæ-less muscles, which do not run quite up to the orifice, but to an oblique line beneath it. Externally to these muscles there are, as in the other Lepadidæ, fine transverse muscles, confined to the middle part of the animal, and running from the carinal margin more than half way round both sides. Attached to the upper notched or folded end of the orifice (g, fig. 5, above the upper ends of the longitudinal muscles), there is a fan of rather strong, striæ-less muscles, expanding downwards, with their lower extremities attached to the outer membrane of the capitulum; these muscles apparently serve to open the orifice: there is a somewhat analogous muscle in Lithotrya, but in no other member of the Family: in Cryptophialus, however, there is a closely similar muscle. Owing to the action of these several muscles, the tissues forming the capitulum and peduncle are, according to Mr. Hancock, highly contractile.

I have stated that the under surface of the upper produced end of the horny disc is concave, and serves for the attachment of several muscles. Of these some run to the basal margin of the great labrum, and no doubt, as usual, move the whole mouth; others, as usual, run to the skin between the labrum and the lower end of the orifice,—i. e. in fig. 5, between the lower end of the row of fine hairs (see fig. 11), which shows where the basal margin of the labrum is situated, and the lower side of the point (a), where the orifice terminates: others run obliquely on both sides towards the point of attachment of the small adductor scutorum muscle (b, fig. 5): others, of considerable strength, and these are more peculiar, run and are attached to the lower end of the orifice (a), and serve apparently to draw up the orifice from within the fissure-like cavity, in which it lies lodged: others, again, extend transversely on both sides, close beneath the inner tunic of the sack, a little beyond the line whence the ovigerous fræna or branchiæ arise. These transverse muscles lie within the longitudinal muscles, and therefore are quite different from the exterior transverse muscles, which are situated more towards the carinal portion of the peduncle and capitulum, and which are common to most Lepadidæ. The internal transverse muscles, and those running to the lower end of the orifice, are peculiar, but we shall hereafter meet with them even more developed in Cryptophialus.

Along the medial carinal line there is, between the two layers of corium, the usual circulatory channel. On each side of this line, on the inside of the sack, there are generally some slight irregular swellings, and sometimes a large extent of the inner surface is irregularly carunculated with little knobs. The sack (e in fig. 5) extends down almost to the basal point of the peduncle (d), more especially when the ovarian cæca are not gorged with ova.

Branchiæ, or ovigerous Fræna.—Within the sack, on each side of the body, rising not far from the ends of the adductor muscle (b), there is a large fillet or fold; the two occupy so exactly the position of the ovigerous fræna that I cannot doubt such is their nature, though, as happens in the case of some species of Pollicipes, they are destitute of their proper glands, and so do not serve for the attachment of the ovigerous lamellæ; this attachment probably is not required, owing to the protected situation which the lamellæ hold in the sack, under the animal’s body, and over the ovarian cæca. From the unusually large size of these so-called fræna, I cannot doubt that they serve as branchiæ, equally well with the plicated folds of membrane, believed to be homologous with the fræna, in the Balanidæ, which have by every one been considered as branchiæ. The fræna are broad and truncated at their upper ends; their margins are sinuous, and their outer surfaces papillose; they run longitudinally down the sack, narrowing as they extend, almost to the basal point of the peduncle, and hence are of considerable length; they are hidden in the section (fig. 5) by the medial, somewhat protuberant mass (c) of ovarian cæca, and partly by the (i) prosoma.

Body.—The body is constructed on the usual type, and indeed does not differ greatly from that of Ibla. The labrum is very large, its lower or basal margin is separated by an unusual space (capable of being contracted or folded) from the lower end of the orifice of the sack; hence the labrum and whole mouth is placed quite remarkably near the upper (or carinal) end of the orifice. This upper end of the orifice, I may remind the reader, is homologically the posterior end of the general covering or carapace, and all that portion of the whole animal (as the sectional figure, 5, stands) below the lower margin of the labrum, on the rostral or ventral surface, is formed by the three anterior segments of the head. The main part of the body, carrying the mouth, is formed by the great development of that segment of the thorax which bears the first pair of cirri (h), here closely adpressed, as usual, to the sides of the mouth. The lower portion of this segment forms the prosoma (i), and has the characteristic outline, but is not much developed. On each side of the prosoma an oval space of membrane is yellowish and is thickened, and so gives support to this part of the body. The five succeeding thoracic segments, which ought to carry the five succeeding and posterior pairs of cirri, are together of very small size (as in Ibla), in comparison with either the prosoma, or the whole anterior part of the animal. The segment (k) which should have borne the second pair of cirri, is considerably longer than the following segments, and is at the same time less distinct, owing to an oval convex shield of thickened membrane on the sides, not extending the whole length of the segment, thus causing two transverse creases, which, when the thorax is contracted, appear like two additional segments. Had this segment borne cirri, they would have stood, as in Ibla, at a considerable distance from the first pair. The segments (l, m) which should have borne the third and fourth pairs of cirri are like each other, except that the former is rather the longest. The membrane covering all the thorax is surprisingly thin; and at the articulations, which are straight and transverse, is deeply folded, so that the thorax must be highly extensible, to a degree which I have not seen equalled in any Cirripede except in the males of Scalpellum vulgare and ornatum. The thorax is represented as somewhat extended in fig. 5. This part of the thorax is amply furnished with striated and striæ-less muscles for its retraction and protrusion, and for lateral movements. The segment (m) which should have borne the fourth pair of cirri, at first sight falsely appears like the terminal segment of the thorax: in one monstrous specimen it bore a single cirrus, showing (if there had been any doubt) that it was a true segment. The three terminal pairs of articulated appendages, form together a brush; they consist of the fifth and sixth pairs of cirri and of the caudal appendages: my reasons for considering the last-named organs as of caudal origin will be given hereafter. A moderately careful inspection, especially of the ventral surface, will show that the fifth pair of cirri are borne on a small segment (n, fig. 5, but plainer in fig. 13), which is quite distinct from, but partially concealed by, that which ought to bear (and did bear in the monstrous case) the fourth pair of cirri: this segment is oblique, and cannot be traced distinctly all round the dorsal surface. The segment (o, fig. 13) bearing the sixth pair is much less distinct, and can only be seen by a longitudinal section, or when the cirri are a little separated, but it certainly exists,[144] as is likewise shown by the presence of small apodemes dipping in amongst the muscles, between this and the last segment. The posterior or caudal appendages are closely approximated; they are not separated by any fold from the sixth thoracic segment; but appear as if they were articulated on the dorsal surface of the sixth pair of cirri, in exactly the manner usual in the other Lepadidæ. The segments bearing the fifth and sixth pairs of cirri are highly oblique to the preceding segments, and consequently the cirri, which they support, instead of projecting inwards, lie like a brush in a line with the longitudinal axis of the main part of the thorax. The membrane forming the two small oblique terminal segments of the thorax is strengthened by irregularly shaped plates of thicker and yellowish membrane.

Mouth.—The mouth is constructed on the strictly normal type of the Family, but is peculiar in every part; it is remarkable from being situated so near the upper (or posterior) end of the capitulum, this being caused by the great length of the labrum, and of the space of body between the latter and the lower end of the orifice. The labrum is a very singular part of the mouth from its vast size and outline: in fig. 8, we have a front view of the mouth, of which the whole upper pointed part consists of the labrum, and h h is the first pair of cirri; in fig. 11, we have a lateral view of the labrum, with the surrounding thin membrane of the body, a—a, still adhering to its edges; h is the first cirrus on the near side; m the mandible, a little distorted in order to show its tooth, marking the position of the transverse crest of the labrum and of the orifice of the œsophagus; b b is the medial longitudinal ridge of the labrum. In the Balaninæ the labrum forms a mere rim to the back of the mouth, consisting of an inner fold running down the œsophagus, and of an outer fold, both close together: in the Lepadidæ the folds are separated, the outer one being swollen or bullate; and here this structure is carried to even a greater extreme than in Ibla and its allies. The distance between the transverse crest over the œsophagus and the blunt projecting point on the summit of the medial ridge, b b, equals twice the longitudinal diameter of the rest of the mouth. The lower margin on each side of the labrum is produced into two projections (fig. 11), the longer one curling round to a point beneath the jaws, with its extremity imbedded as an apodeme. Another very peculiar character in the labrum, prominently noticed by Mr. Hancock, is caused by a longitudinal row, on each side, of closely approximate, long, very finely pointed hairs, which, as already stated, are fronted on the opposed internal surface of the sack by an irregular band of still finer hairs. The surface of the labrum is partially covered by minute toothed scales, and these, seen on the longitudinal medial ridge, b b, give it a finely denticulated structure. At each end of the transverse crest which overhangs the œsophagus, there is a knob, such as occurs on the labrum of every Cirripede. United to these two knobs, which are formed of thick and yellowish membrane, and springing from the adjoining sides of the mandibles, there are two swellings formed of thin membrane (fig. 8), which occupy the exact position of the palpi, and may be considered as these organs in a rudimentary condition and destitute of bristles.

The mandibles are simpler than in any other Cirripede; they are minute; they consist of an oblong plate, with only one very strong tooth at the upper end: the face towards the labrum is swollen: beneath the upper free part there is a small, sub-triangular piece of thickened membrane, let in and forming part of the general outer surface of the mouth, and representing the large square plate found in other Cirripedes. The maxillæ (fig. 15) are smaller but broader than the mandibles; they have an upper tooth and a smaller lower one, lying not quite in the same plane with the upper one, but nearer the mandibles. The apodeme (fig. 8, 15) is of remarkable length, extending beneath the basal fold of the mouth: it does not arise from the ridge or outer edge of the maxilla, but a little on one side, from the face directed towards the mandible. Between maxillæ and mandibles there is a very singular prominent fold of membrane (fig. 8, 15), which resembles, but probably falsely, the supposed rudimentary palpus attached to the mandible. Altogether the maxillæ differ considerably from the same part in other Cirripedes. In structure they seem adapted to assume the function of mandibles; but they do not stand directly over the œsophagus. The outer maxillæ (fig. 8) appear like a minute, deeply notched lower lip: each consists of a simple, oblong, rounded plate, with a few small bristles at its upper end. The basal fold of the mouth in front, beneath the outer maxillæ is distinct, and runs in a line with the basal articulation of the first pair of cirri. In the rudimentary palpi, minute and little developed outer maxillæ; and in the inner maxillæ, taking the function of the mandibles, the mouth of Alcippe presents some resemblance with that of Anelasma.

Cirri.—These consist of the first, fifth, and sixth pairs: the other pairs are absent, except in one monstrous specimen, in which there was a fourth cirrus quite like the fifth. First pair, fig. 14, these are seated on each side of the mouth in the usual position. They are formed of very thin and flexible membrane. The pedicel, as usual, consists of two segments, the upper one is short and not very distinct; but when viewed on the inner side can be seen to have the ordinary structure: both segments are destitute of bristles. There are two short rami, being about one third of the length of the pedicel: they are directed either in the line of the pedicel, or more commonly posteriorly, that is towards the other cirri, and therefore in an unusual direction. The anterior ramus is generally rather longer and thinner (as is commonly the case with other Cirripedes) than the posterior ramus; but there is some variation in this respect. On neither ramus is there any trace of the ordinary articulations: both are thickly clothed with fine bristles, which are singular from being thickened in their lower parts, and plumose, like a feather. These cirri have some resemblance, as remarked by Mr. Hancock, to a pair of pincers; but they cannot act as such; they serve, I believe, as brushes. Delicate muscles, transversely striated, enter and are attached within both rami and within both segments of the pedicel, on the usual type, showing that these organs (if there had been any doubt) are truly cirri.

The fifth and sixth pairs of cirri (fig. 13, n′, o′) are almost exactly alike: they are of very small size: each cirrus consists of four segments: the lower or basal segment is broad, with a few minute bristles scattered on its inner surface: the second segment is also broad, but shorter, with a few, generally hooked bristles, in two short irregular rows, in the upper part: these two segments answer to the two segments of the pedicel of ordinary cirri. The third segment is thinner and longer than the second; it bears two or three longitudinal rows of bristles, most of which are neatly hooked at the point; its upper end is surrounded with a circle of bristles. The fourth and terminal segment is short, thin, and simple, with only a few bristles at the apex. These two upper segments are bent a little inwards; they answer to one of the two normal rami of ordinary cirri. The third segment does not stand exactly on the middle of the summit of the second segment,—the posterior corner of the latter being occupied by a very curious, convex, oblong, rather hard (especially in the lower part), protuberant cushion (as called by Mr. Hancock) or button (fig. 9, c′), transversely wrinkled by fine, distinctly crenated ridges. This button presents a considerably different appearance according to the point of view, fig. 9, 10: on one of its sides it projects beyond the outline of the second segment, whence it arises; on the other side it is prolonged, as a smooth ridge, on the top of the second segment, embracing to a certain extent the base of the third segment. On the face opposite to that which has been drawn (fig. 9) as most characteristic, it is seen to be somewhat constricted round its base; this constriction, representing, I believe, an articulation. When viewed directly in front (fig. 10) its outline is oval, passing into shield-shaped. Its longitudinal axis is 3/1000ths of an inch in length; but it varies a little in shape and size. I shall presently assign my reasons for believing that these buttons are the posterior or inner rami of the fifth and sixth pairs of cirri in a rudimentary and much modified condition.

Caudal Appendages.—These (fig. 13, p) are placed close together, being articulated between the bases of the sixth pair of cirri, the lines of junction being internally marked by minute apodemes. They consist of four segments, resembling in every respect those forming the cirri, with the important exception that there is not a vestige of the button on the summit of the second segment; the segments are not so thick as those of the cirri, and the terminal segment is smaller.

Muscles and Functions of the Cirri.—For their size, the cirri and caudal appendages have voluntary muscles of remarkable strength, attached within their basal segments, and springing from the dorsal and ventral surfaces of the so-called third and fourth (l, m) thoracic segments. Other muscles, rising from within the basal segment of each limb, run to the second segment, and from that to the third segment. I could not distinctly make out whether any entered the terminal segment. I have seen no other instance of muscles entering the caudal appendages, but as in the pupa they are so furnished, we here have only an embryonic character preserved. I may remark that the fifth and sixth cirri, consisting of two large basal and two thinner terminal segments, is likewise an embryonic character. From the position of the cirri, the four hard protuberant buttons or cushions, tend to oppose each other at a common point; and the caudal appendages fill up a gap behind, between the cirri of the sixth pair. I at first thought, with Mr. Hancock, that these buttons served to catch the prey; but, reflecting on their convexity and hardness, they appear very badly adapted for this purpose; it would, in fact, be a marvellous feat to secure, in the dark, any moving object between four balls. On the other hand, this very convexity, the hardness, and especially the crenated ridges, and the powerful muscles (which from the first surprised me), are all well explained, if we suppose the prey, being secured by the terminal segments, to be triturated between these four balls: any part which escaped upwards would, moreover, be retained in a sort of cage, formed by the inwardly inflected terminal segments with their hooked spines. This view of the very curious and unparalleled use made of a modified portion, not of the haunch, but of an upper part of the two posterior pairs of thoracic limbs, is in some degree confirmed by finding that Cryptophialus, which has apparently analogous habits, requires its food to be triturated, though in this case it is effected by very different means, namely, by four beautifully toothed discs, with brushes of hairs, developed within the lower end of the œsophagus.

The prey, when caught, would probably at once be carried by the movement of the articulated thorax to the mouth (itself moveable), and being there secured by the mouth in front, the caudal appendages behind, the tips of the cirri above, and the broad pedicels of the first pair on the two sides, it would be triturated by the four crenated buttons, and would then be forced down the œsophagus by the action of the simple jaws. I looked in vain in several specimens for any object within the stomach. I believe, that when the specimens are first taken, all half digested food is ejected by the mouth. Whether we may thus account for the extremely foul condition of the rami of the first cirri in all the many specimens examined by me, I know not; but that these rami, which are thickly clothed with fine plumose hairs, and are furnished with delicate muscles, act as brushes, so as to clean the orifice of the sack, I can hardly doubt.

Homologies.—I have as yet, to a certain extent, assumed that I have correctly named the different parts; and a few remarks on this head may be desirable, considering the absence of certain cirri, the singular condition of the others, the close general resemblance of the cirri and caudal appendages, and the fact of the latter being furnished with muscles. The only cause for any doubt regarding the thoracic segments is the shield of thick membrane on that segment (k, fig. 5), which ought to have borne the second pair of cirri, causing two transverse wrinkles (not distinguishable, however, on the ventral surface), and sometimes making the segment appear as if it consisted of three segments: if it did consist of three, as there can be no doubt about the nature of the first pair of cirri, (not in a more rudimentary condition than in Anelasma) or about the segment whence this first pair arises, the two terminal oblique segments, with their appendages, would be abdominal instead of thoracic: but this is improbable, inasmuch as the abdomen is unusually little developed in the pupa (as presently to be shown), and more especially from the circumstance of a monstrous cirrus, identical in structure with the two succeeding pairs, having been borne on a segment (m), which, in any case must be considered as thoracic, for it is well known how very rarely thoracic and abdominal limbs resemble each other. I cannot myself feel hardly any doubt on the nature of these three pairs of appendages; for, in the first place, the posterior appendages are articulated on and between the bases of the adjoining pair, exactly as the undoubted caudal appendages are articulated in all other members of the family on the sixth pair of cirri or terminal thoracic appendages. Secondly, we see in the male of the allied genus Ibla, the very same appendages preserved as in Alcippe, namely, the caudal, and the fifth and sixth pairs of cirri, which latter, moreover, are generally uniramous. Thirdly and lastly, in the likewise allied Alepas cornuta, we have the posterior rami of these same fifth and sixth pairs of cirri in a rudimentary condition, and resembling in every respect the caudal appendages. Assuming, then, that the several appendages in Alcippe have been rightly denominated, we have to consider the nature of their segments: in all cirripedes, the pedicels of the cirri consist of two segments, of which the lower one (as here) is longer than the upper one, and both (as here) considerably thicker than the segments of the rami: in all cirripedial pupæ, likewise, the thick pedicels of the limbs consist of two segments, and each ramus, also (as here), of two segments: now, with these coincidences, and bearing in mind that in Alcippe the two upper segments do not arise from the exact middle of the summit of the second segment, but from rather its anterior side,—bearing, also, in mind the case just cited of Alepas cornuta with the posterior rami of these very same cirri rudimentary,—we may, I think, safely conclude that here in Alcippe the two lower segments form the pedicel; the two upper segments, the anterior ramus; and that the button-like protuberance is the posterior ramus in a modified condition. As the caudal appendages in none of the Lepadidæ, either in the mature state or in the pupa, have two rami, we can satisfactorily understand the absence of any trace of the button-like protuberance on the top of the second segment.[145]

Alimentary Canal.—The œsophagus runs down from the mouth, beneath and nearly parallel to the straight row of hairs on the two sides of the labrum: it is surrounded by the usual muscles: at the lower end it bends down, and expanding a little, like a bell, enters the stomach. The stomach is of considerable size and fills the main part of the body, bulging out under the mouth, and prolonged as far as about the middle of that segment (l), which ought to have borne the third pair of cirri; here the stomach terminates in a blunt rounded point. The tissue surrounding the stomach, and keeping it in its proper place, can be traced to the posterior end of the thorax, but there is no rectum or anus. I am prepared to assert positively that this is the case,[146] for I made repeated longitudinal sections of the whole thorax in two planes, and I subsequently cleaned the outer tissues with boiling potash, and then, when as transparent as a sheet of glass, I examined every part, and certainly there is no rectum (which in every case is formed of chitine, and so is not acted on by potash) nor an anal orifice. Singular as this fact is, it is not so improbable as it at first appears; inasmuch as I have shown, in my former volume, that the Lepadidæ can reject half digested food by their mouth, and secondly, that the final stage of digestion appears to take place in the upper part of the stomach. In the male of this very species, as we shall immediately see, there certainly is no mouth or stomach, and apparently no rectum or anus; so is it likewise with the males of Scalpellum vulgare and ornatum: in Proteolepas, there is a mouth and an œsophagus, but no stomach, rectum, or anus. There are, I believe, no other known instances, in the whole great class of Crustacea, of the absence of an anus.

The stomach, in Alcippe, is much corrugated, so as to be deeply pitted; but there are no regular cæca. The enveloping hepatic layer is thick, brownish, pulpy, and formed of pellets of cellular matter, not distinctly arranged in lines as is general; there is the usual delicate muscular layer. The stomach was in every case empty, and I did not notice the separated epithelial coat, so generally found in other cirripedes.

Organs of Sense.—I failed in discovering the eye, which I have no doubt exits, as it is conspicuous in the pupa and in the male. The olfactory pouches are seated rather laterally under the maxillæ, as in Ibla. As in this same genus, the acoustic sack is seated remarkably low down (fig. 5), at a very considerable distance beneath the basal articulation of the first cirrus: the orifice is seated on a slight prominence: the acoustic vesicle, I believe, is sub-cylindrical, with irregular projections. I did not make out anything distinctly on the nervous system.

Female Generative System.—The animal we have thus far described is exclusively female: when a longitudinal section of the thorax is made, and the stomach removed, it can be most plainly seen that there are no vesiculæ seminales or testes. Mr. Hancock has remarked on the absence of the usual probosciformed penis. The male of Alcippe will be subsequently described in detail. The female organs differ in no respect from those of other members of the family, excepting in so far that the layer or mass formed by the ovarian cæca (c) does not lie transversely to the longitudinal axis of the whole animal, but longitudinally under the horny disc. The ovigerous fræna are largely developed, but serve, as previously stated, as branchiæ, and not for their proper function of giving attachment to the ovigerous lamellæ. The ovigerous lamella is single, and nearly corresponds, in size and shape (as would ensue from the manner of its formation) to the under side of the horny disc. The ova are broadly oval, and rather above 1/100th of an inch in length.

Metamorphoses.—The larva in the first stage has been fully described and figured by Mr. Hancock: it differs in no essential respect from other larvæ of the family. Mr. Hancock overlooked the inferior minute antennæ. With respect to the larvæ in the last stage, or pupa, I obtained several specimens attached to the disc of the female, and which were on the point of being developed into males; and another specimen identical in all respects, but attached independently to the shell of the mollusc, and which, therefore, I have every reason to suppose, would have been developed into a female. In any case these pupæ may be conveniently here described (Pl. 23, fig. 16.) They are .025 of an inch in length; they are of the usual shape, with the anterior end not very blunt and the postero-ventral surface somewhat produced. The whole carapace or shell is very thin and smooth. There are six pairs of thoracic natatory legs, situated far back towards the posterior end of the body; each leg has the usual articulations, and the two rami their usual long but not plumose spines; the presence of the legs deserves notice, considering the rudimentary and modified state of their homologues in the mature animal. The abdomen differs considerably from the same part, as far as I have seen, in other pupæ; it consists of only a single almost globular (fig. 17, q) segment, instead of three segments; and the two caudal appendages (r) are very long, and are composed each of only a single segment (instead of two), carrying at its tip two short spines. There are two purple eyes, 4/3000ths of an inch in diameter, which, after having been dried and then soaked, could be seen to be compound; they are fixed in the usual manner to two rather short apodemes, which latter have their usual origin. But the pupa has a very unusual appearance owing to the presence of a single dark purple eye, half the diameter of the two larger eyes, situated behind and above the latter, and quite disconnected with the apodemes; this is the eye of the mature animal, which, for some reason, is here developed earlier than usual. The prehensile antennæ are remarkable from being seated very close to the anterior extremity: owing to this, the articulation of the second or main segment with the basal segment, is hardly at all oblique. The whole pupa is of exactly the same length as the pupa of Ibla quadrivalvis, and so are the antennæ, (see p. 286 of my volume on the Lepadidæ), viz., 32/6000th of an inch, but the second segment is narrower, (being only 8/6000ths in breadth in the broadest part), and is longer in proportion, for the disc which forms part of the total length is only 4/6000ths in length, whereas in Ibla it was 8/6000ths; the disc is here hoof-shaped, as in Ibla. The ultimate segment is remarkably short and narrow, (being only 3/20000ths in width, and less than half the size of that in Ibla); it carries (I believe) three terminal spines, and is not notched. Altogether the antennæ more nearly resemble those of Ibla than of any other genus in the family. From the position of the antennæ, and from the length of the second segment, the pupa, when cemented by the disc or third segment, to the supporting surface, adheres, with its posterior end almost vertically upwards. With respect to the young cirripede within the pupa, I could only observe that its anterior end was formed into a blunt point.

Powers of Excavation; Inorganic Deposit of Calcareous Matter; Attachment.—Alcippe, according to Mr. Hancock, attacks only dead shells of the Fusus and Buccinum, and always on their inner sides, especially on the columella. The excavations, in the specimen which I examined, were so numerous as almost to touch, and sometimes to run into each other, the included animal being thus rendered distorted. The orifices are directed with respect to the shell indifferently upwards or downwards. From the shape and size of the cavity corresponding to that of the included animal, there can be no doubt, as stated by Mr. Hancock, that Alcippe forms its own cavity. That the action is mechanical I think may safely be inferred from the whole outer membrane being studded with minute, star-headed points of hard chitine, which rise from halo-like little discs of thickened membrane, which latter are well adapted to allow the underlying adherent muscular layer to act on the points, and thus on the surrounding shell. Consequently the points generally show signs of severe attrition, but they are periodically and often replaced, at each exuviation, by new and much sharper points. There are no points on the permanently attached layers of the horny disc, but it particularly deserves attention, that the renewable membrane always extends beyond the circumference of the disc, and is there most thickly studded with the points. We have met, in Lithotrya, with a precisely analogous fact in the extension of the periodically moulted membrane of the peduncle, furnished with star-headed points of chitine, and in addition with minute calcareous beads (which, however, seem soon worn away), beyond the calcareous discs, by which this cirripede is attached in its cavity. We need not feel much surprise at points of chitine being hard enough to wear away shell, when we consider what work the jaws of insects, likewise formed of chitine, will effect.

With respect to the first commencement of the excavation, the pupa, owing to the position of its prehensile antennæ, fixes itself with its posterior end almost vertically upwards; and the young cirripede, after its metamorphosis, from the greater length of the ventral integuments formed round the eye-apodemes, must be thrown backwards into nearly the position represented in Pl. 22, fig. 12, b. I have not seen a young female at this early age, but I have traced the development of several males, and have found that the lower end of the peduncle, (i. e. what was the anterior end of the pupa), grows at quite a remarkable rate, so as very soon to form a great bag extending beyond the attached prehensile antennæ. Now if we suppose an analogous structure in the female or ordinary Alcippe, and the supposition is quite allowable, we shall almost immediately have the anterior or lower end of the young cirripede, just in advance of its antennæ, pressing against the surface of the shell of the mollusc; and if armed with triturating points, as we have every reason to believe it is, it would wear for itself a cavity. The horny disc on the ventral surface of this protuberant anterior end of the young animal will, we may assume, soon become cemented to the near side of the cavity just supposed to have been excavated. And the whole animal, by further slight changes in direction, namely, by working down more and more obliquely, will take, as shown at (c), its final position. As the whole surface of the animal, with the exception of the horny disc, is provided with triturating points, the animal, when once imbedded, can and does increase its cavity at both ends in length, in depth, and all round the edges of the horny disc,—in short, in every direction excepting directly over the horny disc. I believe, as already explained, that the young Alcippe, (b, diagram), first bores obliquely into the shell; and whatever amount of downward extension the horny disc attains before the young cirripede assumes its proper position, with its ventral surface upwards and parallel to the inner surface of the shell of the mollusc, that amount determines the thickness of the plate of shell hereafter to be left unabraded over the horny disc, as the latter continues to extend in circumference. This plate of shell over the horny disc is so thin, that, as mentioned at the commencement, the colour of the ovaria is seen through; and until I reflected on the following considerations, I was much surprised how the instinct of the animal could so neatly guide it not to grind too deeply, and yet to grind till only a very thin plate of shell was left over its horny disc: these considerations are, that whatever thickness was first given to this plate of shell, when the animal was very young and first assumed its ultimate position, that thickness would in most cases be always retained, owing to the flatness of the disc, and to the membrane armed with triturating points protruding very slightly beyond and above the horny disc, only just enough to wear away the surrounding shell to the thickness necessary to allow of the formation of each new zone of disc; as the disc itself is not armed, it subsequently has no power of wearing away the plate of shell above it. Thus the horny disc, besides giving support and attachment to the peduncle, is of this peculiar service that it seems to guide, (somewhat like the wood-part in a plane), the rasping powers of the lower extreme margin of the peduncle.

I may here observe that certain radiating and often punctured lines, mentioned and figured by Mr. Hancock, which help to render the thin plate of shell over the peduncle conspicuous (fig. 3), are formed by the burrows of an excessively minute annelid, the punctures being apparently the exit orifices: I imagine that these annelids find it difficult to commence their burrows on the smooth surface of the shell, and that they congregate at these particular spots and thence burrow in radiating lines, owing to their having taken advantage of the little cliff-like edges, at the narrow and disused ends of the fissures leading into the cavities occupied by the Alcippe, where alone they would not be disturbed by the action of the cirri, when first they commenced making their little burrows in the shell.

The fissure leading into the cavity is required to be broad at the posterior end, in order that the cirri may be there freely exserted out of the sack; and narrow in other parts, to prevent, as it would appear, anything injurious getting in between the animal’s body and the cavity in the shell of the mollusc. As the fissure is increased in length by attrition at the broad posterior end, which end during growth becomes broader and broader, the lower part of the fissure has to be narrowed, and this is effected in a very singular manner, namely, by advantage being taken of the strong tendency, which triturated shell with animal matter, has to set into a solid shelly mass, although constantly agitated.[147] Mr. Hancock noticed this edging of hard shelly matter, and naturally thought it was a secretion. Lines of deposition (Pl. 22, fig. 4, b), parallel to the edges of the furrow can often be perceived in it: its thickness and extension vary much: I have seen it on one side alone of the orifice: it is, of course, never found at the broad end where the process of enlargement goes on. The peculiar worn surface with which it irregularly thins away downwards, on the sides of the cavity, made me (together with the apparent impossibility of such a secretion proceeding from an animal wholly invested by a chitine membrane) suspect it to be inorganic; and this view is certainly correct, for when a fragment is dissolved in acid, a considerable residuum is left of bits of membrane, rubbish, and, in one instance, even of the remnants of a foreign animal, apparently an annelid. We have here all the circumstances favorable for inorganic deposits of this nature, namely, finely triturated shell and chitine or animal matter, produced by the excavation of the chamber, sea-water, and movement.

From the manner of growth of the animal, the fissure leading into the cavity in the shell becomes much longer than the orifice leading into the sack, and to prevent the body being unnecessarily exposed, the upward projection of the disc, already described, is formed under the narrow and disused end of the fissure; moreover, the two rims of the inorganic calcareous deposit sometimes here approach so closely, as almost or actually to touch each other; and between them, as remarked by Mr. Hancock, there is usually a little accumulation of grains of sand. This narrow end of the fissure is generally curled either to the right or left hand; and I can only account for this fact by supposing that, whilst the cirripede is young, and has not a large horny disc attached to the cavity, it cannot keep its body straight during the long-continued boring process.

The animal is attached by its horny disc to the thin shelly roof over the peduncle, and likewise to the under side of the narrow end of the fissure, but is elsewhere quite free. I carefully examined the disc in many specimens, but could not see any cement-ducts: I believe I saw layers of cement at the upper end of the disc, but it is not easy to discriminate between this substance and the yellowish, somewhat disintegrated, layers of the horny disc. The pupa certainly becomes attached by ordinary cement, so that the attachment in early life, at least, is normal. In some full-grown specimens, I found the lower parts of the horny disc attached, along the edges of the layers, to the roof of shell; and as I looked here in vain with the highest powers for cement-ducts, or for cement, it appears to me probable that the rough edges of these layers were united to the roof by a thin layer of the inorganic calcareous deposit. The animal, from its very protected situation, certainly requires to be less firmly cemented than other cirripedes; and even in Lithotrya, which is less deeply imbedded than Alcippe, the cementing apparatus was feebly developed. From the length of the pupal antennæ, cemented by their terminal segments, the position of the young cirripede (Pl. 22, fig. 12) can be changed to a considerable extent, like a ship swinging at her moorings, but in order to assume its final position, the animal must, I think, travel like Lithotrya, but to a much less extent, by a short succession of overlapping horny discs,—the old discs being partially deserted, each new one extending beyond the last-formed one: even in the case of the mature animal, we have seen that, under certain circumstances, it changes, to a certain extent, its position; portions of the old disc being deserted and attached to the roof of a deserted portion of the cavity.

Affinities.—In the preliminary remarks under the Family, I have discussed this subject almost sufficiently: I will here only remark, that the genus, though so abnormal, yet stands naturally between Ibla and Anelasma, having clear affinities, on the one side, through and beyond Anelasma to Alepas; and on the other side, beyond Ibla to Scalpellum, and so to Lithotrya. Moreover, it is very distinctly related to Cryptophialus in the succeeding Order.