The Mandibles ([Pl. X], [figs. 1-5]) have from two to ten strong teeth in a single row; where the number exceeds five, several of the teeth are small; the inferior angle is generally pectinated with fine spines; in Lithotrya ([fig. 2]), the interspaces between the teeth are also pectinated. In the same individual there is not unfrequently one tooth, more or less, on opposite sides of the mouth. Internally, the mandibles are furnished on their outer and inner sides with several ligamentous apodemes, in Lithotrya roughened with points ([Pl. X], [fig. 2]), for the attachment of the muscles; of these ([fig. 1]), there is a chief depressor and elevator, attached at their lower ends to near the basal fold of the mouth, and a lateral muscle, attached to the broad basal end of the palpi, and serving, apparently, to oppose the edge of mandible to mandible. The Maxillæ in the different genera ([Pl. X], [figs. 9 to 15]) differ considerably in outline; they are generally about half the size of the mandibles; at the upper corner, there are always two or three spines larger than the others, and often separated from them by a notch; the rest of the spinose edge is straight, or irregular, or step-formed, or with the lowest part projecting, or with one or two narrow prominences bearing fine spines. All these spines, quite differently from the teeth of the mandibles, are articulated on the edge of the organ, and stand in a double row. At a point corresponding with the upper articulation of the mandibles, a long, thin, narrow, rigid apodeme, projects inwards ([fig. 10]), and running down nearly parallel to the thin, outer, flexible membrane of the mouth, is attached to the corium, and thus serves as a support to the whole organ. This apodeme is embedded in muscles ([Pl. X], [fig. 10]); there are other large muscles attached to the inner side of the organ, and again others running laterally towards the mandibles. The apodeme, of course, is moulted with the integuments of the mouth. The Outer Maxillæ ([Pl. X], [figs. 16, 17]) serve as a lower lip; they are thicker than the other trophi; they have their inner surfaces clothed with spines, sometimes divided into an upper and lower group, and occasionally separated by a deep notch: there are often long bristles outside. They are furnished with at least two muscles; in sessile Cirripedes I have seen that they are capable of a rapid to and fro movement, and I have no doubt that their function is to brush any small creature, caught by the cirri, towards the maxillæ, which are well adapted to aid in securing the prey, and to hand it over to the mandibles, by them to be forced down the œsophagus. On the exterior face of the outer maxillæ, above a trace of an upper articulation, either two small orifices or two large tubular projections can always be discovered; and these, as will presently be mentioned, I believe to be olfactory organs.
Cirri.—The five posterior pair are seated close to each other and equidistant; the first pair is generally seated at a little distance, and sometimes at a considerable distance from the second pair. The first pair is the shortest; the others, proceeding backwards, increase gradually in length. The rami of each pair are either equal in length or slightly unequal: those of the first pair are oftenest unequal. The number of segments in the posterior cirri is sometimes very great; in one species of Alepas, there were above sixty segments in one ramus, the other ramus being in this unique case ([Pl. X], [fig. 28]) small and rudimentary. The pedicels consist of two segments, a lower, longer, and upper short one ([fig. 18, c, d.]) In the usual arrangement of the spines on the segments of the three posterior pair of cirri, there are ([figs. 26, 27]) from three to six pair of long spines on the anterior face, with generally some minute spines (occasionally forming a tuft) intermediate between them: on the dorsal surface, in the uppermost part of each segment, there is a tuft of short spines generally mingled with some longer, finer ones: on the inner side of each segment, on the upper rim, there are generally a few extremely minute and short spines. From the increase of these latter and of the intermediate spines, the antero-lateral faces of the segments of the first cirrus, and of the lower segments of the anterior ramus of the second cirrus ([Pl. X], [fig. 25]), are almost always thickly paved with brush-like masses of spines. The lower segments of the anterior ramus of the third cirrus is generally, though not always, thus paved: these paved segments are much broader than the others. The posterior rami of the second and third cirri are often in some slight degree paved, though in other cases they resemble the three posterior pair of cirri. The two segments of the pedicels have bristles on their anterior faces, essentially arranged on the same plan as on the segments of the rami: the bristles are generally not so symmetrically arranged on the pedicels of the second and third cirri, as on the three posterior pair. There are some exceptions to the foregoing general rules: in the posterior cirri of [Alepas cornuta], there is only one pair of long spines to each segment ([fig. 28]); in [Dichelaspis Lowei], there are eight pair; in [Lepas fascicularis], in old specimens, the segments are paved with a triangular brush of spines; the upper segments in [Pæcilasma eburnea] support small oblong brushes; and, lastly, in [Pæcilasma fissa] ([fig. 29]), and crassa, the spines form a single circle round each segment, interrupted on the two sides. These spines are often doubly serrated or plumose: many of them on the protuberant segments of the first three pair of cirri, are sometimes coarsely and doubly pectinated.
Caudal Appendages.—These are present ([Pl. X], [figs. 18 to 24]) seated on each side of the anus, in all the genera, except in Conchoderma, Anelasma, and [Scalpellum villosum]; they consist of a very small single segment, destitute of spines in Lepas, and spinose in Pæcilasma, Dichelaspis, Oxynaspis, Scalpellum, and some species of Pollicipes; they consist of several segments in Alepas, Ibla, Lithotrya, and in some species of Pollicipes. In the latter genus, some species have their caudal appendages multiarticulate, though so obscurely articulated, that the passage ([fig. 22]) from several to one segment is seen to be easily effected. When the appendage consists of many articulations, it is generally about as long as the pedicel of the sixth cirrus; but in [Ibla quadrivalvis], it is four times as long. The segments are narrow, slightly flattened, much tapering; each ([fig. 24]) is surmounted by a ring of short spines, which are generally longest on the apex of the terminal segment. I could never trace muscles into these appendages.
Alimentary Canal.—The œsophagus is of considerable length: it is formed of strong, transparent, much folded membrane, continuous with the outer integuments, and moulted with them: it is surrounded by corium, and as already stated, by numerous muscles: at its lower end it expands into a bell, with the edges reflexed, and sometimes sinuous: this bell lies within the stomach, and keeps the upper broad end expanded. According to the less or greater distance of the mouth from the adductor muscle, the œsophagus runs in a more or less parallel course to the abdominal surface between the first and succeeding pairs of cirri, and enters the stomach more or less obliquely. In Ibla alone, it passes exteriorly to, and over the adductor scutorum muscle. The stomach lies in a much curved, almost doubled course; it is often a little constricted where most bent; it is broadest at the upper end, and here, in Lepas and Conchoderma, there are some deep branching cæca; in the latter of these two genera, the whole surface is, in addition, pitted in transverse lines. The stomach is coated by small, opaque, pulpy, slightly arborescent glands, believed to be hepatic; these are arranged in longitudinal lines, in all the genera, except in Alepas, in which they are transverse and reticulated: the whole stomach is thus coated. There is, also, a coating of excessively delicate, longitudinal and transverse muscles without striæ. The rectum varies in length, extending inwards from the anus to between the bases of the second and fifth pair of cirri: it is narrow, and formed of much folded transparent membrane, resembling the œsophagus, continuous with the outer integuments, with which it is periodically moulted. The anus is a small longitudinal slit, in the triangular piece of membrane representing the abdomen, let in between the last thoracic tergal arches, as already mentioned under the head of the Metamorphoses; it lies almost between the caudal appendages, and opens on the dorsal surface. Within the stomach, there can generally be plainly seen, in accordance with the period of digestion when the specimen was taken, a thin, yet strong, perfectly transparent epithelial membrane, not exhibiting under the highest power of the microscope any structure: it enters the branching cæca, and extends from the edge of the bell of the œsophagus to the commencement of the closed rectum, and consequently terminates in a point: it consists of chitine, like the outer integuments of the animal, and by placing the whole body in caustic potash, I have dissolved the outer coats of the stomach, and seen the bag open at its upper end, perfectly preserved, floating in the middle of the body, and full of the debris of the food. In most of the specimens which I have examined, preserved in spirits of wine, this epithelial lining was some little way distant and separate from the coats of the stomach; and hence was thought by M. Martin St. Ange to be a distinct organ, like the closed tube in certain Annelids. Occasionally, I have seen one imperfect epithelial bag or tube within another and later-formed one. Digestion seems to go on at the same rate throughout the whole length of the stomach; if there be any difference, the least digested portions lie in the lower and narrower part. The prey, consisting generally of crustacea, infusoria, minute spiral univalves, and often of the larvæ of Cirripedes, is not triturated: when the nutritious juices have been absorbed, the rejectamenta are cast out through the anus, all kept together in the epithelial bag, which is excluded like a model of the whole stomach, with the exception of that part coated by the bell of the œsophagus. I have sometimes thought that the bag was formed so strong, for the sake of thus carrying out the excrement entire, so as not to befoul the sack. I believe Lepas can throw up food by its œsophagus; at least, I found in one case, many half-digested small Crustaceans in the sack, and others of the same kind in the stomach.
Circulatory System.—I can add hardly anything to what little has been given by M. Martin St. Ange: like others, I have failed, as yet, in discovering a heart. The whole body is permeated by channels, which have not any proper coat: there is one main channel along the ventral surface of the thorax, dividing and surrounding the mouth, and giving out branches which enter the inner of the two channels in each cirrus: as Burmeister has shown, there are also two channels in the penis. There are two dorso-lateral channels in the prosoma, which are in direct connection with the great main channel, running down the rostral (i. e., ventral) side of the peduncle. This latter main channel branches out in the lower part, and transmits the fluid through the ovarian tubes, whence, I believe, it flows upwards and round the sack, re-entering the body near the sides of the adductor scutorum muscle. The main rostral channel (or artery?) in the uppermost part of the peduncle, has a depending curtain, which, I think, must act as a valve, so as to prevent the circulating fluid regurgitating into the animal’s body during the contractions of the peduncle.
Nervous System and Organs of Sense.—In most of the genera, there are six main ganglia, namely, the supra-œsophageal, and five thoracic ganglia; but in [Pollicipes mitella] there are only four thoracic ganglia. Of these, the first thoracic or infra-œsophageal ganglion is considerably the largest and most massive; it is squarish, or oval, or heart-shaped; it presents no trace of being formed by the union of two lateral ganglia. Two great nerves spring from its under side (A), represented in the woodcut on page 49, by dotted lines, and run straight down amongst the viscera in the prosoma: these nerves are about as large as those forming the collar and those running to the second ganglion; hence, six great nerves meet here, two in front, two behind, and two on the under side. At the anterior end, over the junction with the collar chord, three equal-sized nerves rise on each side, with a fourth, smaller one, outside; these go to the trophi and to the two olfactory sacks. At the posterior end, on each side, a pair of nerves branch out rectangularly, one of which (a,) goes to the first cirrus, and there divides into two branches; of these, the upper runs up the cirrus, and the lower one downwards. The other nerve (b), proceeding on each side from this first thoracic ganglion, runs to the muscles beneath the basal articulation of the first cirrus. The collar surrounding the œsophagus is generally very long, sometimes equalling the whole thoracic chord; at a middle point, a small branch is sent off, and at the anterior end (e, e), close to the supra-œsophageal ganglia, double or treble fine branches run to the true ovaria, lying close to the upper end of the stomach. The four (or only three) other thoracic ganglia, when viewed as transparent bodies, are seen to be solid; but in some of the genera, as in Conchoderma, the outline plainly shows, that each consists of a lateral pair fused together. The second thoracic ganglion (B) is rather small; it is either close to the first, as in Pollicipes mitella and [Lepas fascicularis], or far distant, as in Ibla. The third (C) and fourth are of about the same size with the second: these three ganglia send large branches to the second, third, and fourth pair of cirri: other minute branches spring from their under sides, and from the intermediate double chords. The fifth ganglion is larger and longer than the three preceding ones, and gives off nerves to the fifth and sixth pair of cirri; it is clearly formed by the union of the fifth, with what ought to have formed a sixth ganglion. The two nerves going to the sixth cirrus give off on their inner sides, each a great branch to the penis. In [Pollicipes mitella], in which there are only four instead of five thoracic ganglia, it is evident from the outline and position of the nerves going to the fourth pair of cirri, that the fourth ganglion is fused into the fifth, itself, as we have just seen, normally composed of two consecutive ganglia. In this Pollicipes there is other evidence of concentration in the nervous system, for none of the ganglia show signs of being formed of lateral pairs; the second is close to the first; and the abdominal double chord is in part separated by a mere cleft; lastly, as we shall immediately see, the same remark is applicable to the supra-œsophageal ganglia.
The latter (D) alone remain to be described; they present far more diversity in shape than do the thoracic ganglia; they are almost always seen in outline to be laterally distinct, and usually resemble two pears with their tapering ends cut off and united; in a transverse line they are as long as the infra-œsophageal ganglion, but are much less massive. In [Lepas fascicularis] (D), they are pear-shaped; in [Pollicipes mitella] they are globular, and separated by a third globular ganglion, which I believe is the ophthalmic ganglion, presently to be described; in [Pollicipes spinosus], however, the ophthalmic ganglion is, as usual, placed in advance of the supra-œsophageal ganglion, which latter, in this one species, shows no sign of being formed of a lateral pair fused together. In [Alepas cornuta] the supra-œsophageal ganglion consists of two quite distinct ganglia, elongated in the longitudinal axis of the body, and separated from each other by the whole width of the mouth; the chord which unites them is of the same thickness as the rest of the collar. In all the genera, from the front of each of the two supra-œsophageal ganglia, a pair of nerves, (f, f,) united and together as large as the collar nerve, rises, and can be traced running unbranched, in a nearly straight line, for a length equalling the whole rest of the nervous chord, so as to supply the peduncle and the inside of the capitulum or sack. At the inner ends of these two same ganglia, from a central point where they are united, a little central branch runs in front to the adductor scutorum and other adjoining muscles; and still smaller fibrils run behind to the œsophageal muscles.
[Diagram of the anterior portion of the nervous system in Lepas fascicularis.] A. First thoracic or infra-œsophageal ganglion. B. Second thoracic. C. Third thoracic ganglion. D. Supra-œsophageal ganglion. E. The two ophthalmic ganglia. F. Double eye. a. Nerve going to first cirrus; b, to the muscles below the first cirrus; c, to the second cirrus; d, to the third; e, nerves running to the ovaria; f, double nerves supplying the sack and peduncle.
Ophthalmic Ganglia and Eyes.—Owing to Professor Leidy’s[16] discovery of eyes in a Balanus, I was led to look for them in the Lepadidæ. Extending from the front of the two supra-œsophageal ganglia, two chords may be seen in [Lepas fascicularis] (of which a rude diagram is here given), to run into two small, perfectly distinct oval ganglia (E), which are not united by any transverse commissure. From the opposite ends of these two ganglia smaller nerves run, and, bending inwards at right angles, enter, beyond the middle, an elongated (F), almost black, eye, composed of two eyes united together. Although in outline the eye appears single, two lenses can be distinctly seen at the end, directed upwards and towards the ganglia; two pigment-capsules can also be distinguished; these are deep and cup-formed, and of a dark reddish-purple. The following measurements will show the proportions of the parts in a specimen of the [Lepas fascicularis] having a capitulum 4/10ths of an inch in length.