Chapter 4

I will now proceed to a general description of the different parts and organs in the Lepadidæ. The Capitulum is usually much flattened, but sometimes broadly oval in section. It is generally formed of five or more valves, connected together by very narrow or broad strips of membrane; sometimes the valves are rudimental or absent, when the whole consists of membrane. When the valves are numerous, and they occasionally exceed a hundred in number, they are arranged in whorls, with each valve generally so placed as to cover the interval between the two valves above. Of all the valves, the scuta are the most persistent; then come the terga, and then the carina; the rostrum and latera occur only in Scalpellum and Pollicipes, and in a rudimentary condition in Lithotrya, and, perhaps, in the fossil genus Loricula. The valves are formed sometimes of chitine (as in Ibla and Alepas), but usually of shell, which varies from transparency to entire opacity. The shell is generally white, occasionally reddish or purple; exteriorly, the valves are covered by more or less persistent, generally yellow, strong membrane. The scuta and terga are always considerably larger than the other valves: in the different genera the valves differ so much in shape that little can be predicated of them in common; even the direction of their lines of growth differs,--thus, in Lepas and some allied genera, the chief growth of the scuta and of the carina is upwards, whereas in Pollicipes and Lithotrya, it is entirely downwards; in Oxynaspis, and some species of Scalpellum, it is both upwards and downwards. Even in the same species, there is often very considerable variation in the exact shape of the valves, more especially of the terga. The adductor muscle is always attached to a point not far from the middle of the scuta, and it generally has a pit for its attachment. In several genera, namely, Pæcilasma, Dichelaspis, Conchoderma, and Alepas, the scuta show a tendency to be bilobed or trilobed. The valves are placed either at some distance from each other, or close together; but their growing margins very rarely overlap each other, though this is sometimes the case with their upper, free, tile-like apices; in a few species the scuta and terga are articulated together, or united by a fold. The membrane connecting the valves, where they do not touch each other, is like that forming the peduncle, and is sometimes brilliantly coloured crimson-red; generally, it appears blueish-gray, from the corium being seen through. Small pointed spines, connected with the underlying corium by tubuli, are not unfrequently articulated on this membrane: the tubuli, however, are often present where there are no spines. To allow of the growth of the capitulum, the membrane between the valves splits at each period of exuviation, when a new strip of membrane is formed beneath, connected on each side with a fresh layer of shell,--the old and outer slips of membrane disintegrating and disappearing: when there are many valves, the line of splitting is singularly complicated. This membrane consists of chitine,[13] and is composed of numerous fine laminæ. After the valves have been placed in acid, a residue, very different in bulk in different genera, is left, also composed of successive laminæ of chitine. It appears to me that each single lamina of calcified chitine, composing the shell, must once have been continuous with a non-calcified lamina in the membrane connecting the several valves: at the line where this change in calcification supervenes, the chitine generally assumes some colour, and becomes much harder and more persistent; and as the whole valve is formed of component laminæ thus edged (the once continuous laminæ of non-calcified chitine connecting the valves, having disintegrated and disappeared) the surfaces of the valves are generally left covered by a persistent membrane, constituted of these edgings: this membrane has been called the epidermis. In some genera, as in Lepas, this so-called epidermis is seldom preserved, excepting on the last zone of growth: in Scalpellum and Pollicipes it usually covers the whole valves. It appears to me that the laminæ of chitine, and of calcified chitine composing the valves, are both formed not by secretion, but by the metamorphosis of an outer layer of corium into these substances.

[13] Chitine is confined to the Articulata. It was Dr. C. Schmidt (Contributions, &c., being a Physiologico-Chemical investigation: in Taylor’s ‘Scientific Memoirs,’ vol. v), who discovered that the membrane connecting the valves and forming the peduncle, and the tissues of the internal animal, were composed of this substance. But Dr. Schmidt says that the valves in Lepas are composed of 3.09 of albuminates, and 96.81 of incombustible residue; I cannot but think that the existence of the albuminates is an error caused by Dr. Schmidt’s belief that the Cirripedia were intermediate between Crustacea and Mollusca, in the shells of which latter, the animal basis consists of albuminates. For after placing the valves of Lepas and Pollicipes in cold acid, I found that the membrane left could _not be dissolved_ in boiling caustic potash, but could, though slowly, (and without change of colour,) in boiling muriatic acid; and these are the main diagnostic characters of Chitine, compared with albuminous substances. I may add, that Schmidt was also induced to consider the shells of Cirripedia as having the same nature with those of Mollusca, from finding that in the above 96.81 of incombustible matter, 99.3 consisted of carbonate and only 0.7 of phosphate of lime; but Dr. Schmidt’s own analyses prove how extremely variable the proportions of these salts are in the Crustacea, as the following instance shows:--

_Lobster._ _Squilla._

Phosphate of Lime 12.06 47.52 Carbonate of Lime 87.94 52.48

And, therefore, it is not very surprising that Cirripedia should have still less phosphate of lime in their shells, than has a lobster compared with a squilla.

Within the capitulum is the sack, which, together with the upper internal part of the peduncle, encloses the animal’s body. The sack is lined by a most delicate membrane of chitine, under which there is a double layer of corium; this double layer is united together by short, strong, transverse bundles of fibres, branched at both ends:[14] in some genera, the ovarian tubes extend between these two layers. We have seen, under the head of the Metamorphoses, that the delicate tunic lining the sack is simply a duplicature of the thick membrane and valves forming the capitulum, the whole being the posterior portion of the carapace of the larva slightly modified.

[14] I am much indebted to Mr. Inman of Liverpool for having kindly sent me excellent specimens illustrating this structure.

_Peduncle._--Its length varies greatly in different species, and even in the same species, according to the situation occupied by the individual; its lower end is sometimes pointed, but generally only a little narrower than the upper end. In outline, the peduncle is usually flattened, but sometimes quite cylindrical. It is composed of very strong, generally thick, transparent membrane, rarely coloured reddish, and often penetrated by numerous tubuli. The underlying corium is sometimes coloured in longitudinal bands. At each period of growth a new and larger integument is formed under the old one, which gradually disintegrates and disappears; the extreme lower point is often deserted by the corium, and ceases to grow, whilst the whole upper part still continues increasing in diameter: in length the chief addition is made (as is clearly seen in those genera having calcified scales), round the upper margin, at the base of the capitulum. The surface of the membrane is either naked or superficially clothed with minute, pointed, articulated spines, or it is penetrated by calcified scales or styles, (in Ibla alone formed of chitine,) which pass through it to the corium, and are added to at their bases, like the valves, at each period of growth. In Lithotrya alone the scales of the peduncle are moulted together with the connecting membrane. These scales on the peduncle are generally placed symmetrically in whorls, with each scale corresponding with the junctions of two scales, both above and below. Except in _Scalpellum ornatum_ and the fossil _Loricula pulchella_, they are very small compared with the valves of the capitulum. When the scales are symmetrical, new ones are first formed only round the summit of the peduncle, and only those in the few uppermost whorls continue to grow or to be added to at their bases; afterwards membrane is deposited under them. The shelly matter of the scales resembles that of the valves, and the manner of growth is the same; tubuli generally run to and through them from the corium. From the continued enlargement of the membrane of the peduncle, the scales come to stand, in the lower portion, some way apart. In Ibla, new horny styles are formed indifferently in all parts of the peduncle. In some species of Pollicipes, the calcareous styles are not symmetrical or symmetrically arranged; and besides those first formed round the top of the peduncle, there are other and larger ones formed near its base. Lastly, in Lithotrya we have a row of calcareous discs or an irregular, basal cup, formed in the same manner as the valves of the capitulum: in this genus alone (as already stated,) the calcified scales are moulted, and here alone their edges are serrated.

The peduncle is lined within by three layers of muscles, longitudinal, transverse, and oblique, all destitute of the transverse striæ, characteristic of voluntary muscles; they run from the bottom of the peduncle to the base of the capitulum, as in Lepas, or half way up it, as in Conchoderma; in Alepas alone they surround the whole capitulum up to its summit. In Lithotrya there are two little, fan-like, transverse muscles (involuntary), extending from the basal points of the terga to a central line on the under side of the carina. The gentle swaying to and fro movements, and the great power of longitudinal contraction,--movements apparently common, as I infer from facts communicated to me by Mr. Peach, to all the Pedunculata,--are produced by these muscles. The interior of the peduncle is filled up with a great mass of branching ovarian tubes; but in Ibla and Lithotrya, the upper part of the peduncle is occupied by the animal’s body.

_Means of Attachment._--If the peduncle be very carefully removed (Tab. IX, fig. 7 and Tab. I, fig. 6 _b_), from the surface of attachment, quite close to the end, but not at the actual apex, the larval prehensile antennæ can always be found: these have been sufficiently described for our present purpose under the head of the Metamorphoses; but I may add, that the diagnostic differences between them in the several genera are briefly given, for a special purpose, in a discussion on the sexes of Scalpellum at the end of that genus. We have seen in the larva, that the cement-ducts, with their opaque cellular contents, can be traced from within the discs of the antennæ to the anterior or lower ends of the two gut-formed bodies, which it can be demonstrated are the incipient ovaria.

In mature Cirripedes these ducts can be followed, in a slightly sinuous course, along the muscles on each side within the peduncle, till they expand into two small organs, which I have called cement-glands. These glands are found with great difficulty, except in _Conchoderma aurita_, where they are placed on each side under the inner layer of corium, at the bottom of the sack, so as to be just above the top of the peduncle; they resemble in shape a retort, (Pl. IX, fig. 3.). In _Pollicipes mitella_ and _polymerus_ they lie half way down the peduncle, close together, and apparently enclosed within a common membrane; in these two species the broad end of the gland is bent towards the neck of the retort. In Scalpellum the position is the same, but the shape is more globular. In Ibla the structure is more simple, namely, a tube slightly enlarged, running downwards, bent a little upwards, and then resuming its former downward course, the lower portion forming the duct. The gland contains a strongly coherent, pulpy, opaque, cellular mass, like that in the cement-ducts; but in some instances, presently to be mentioned, this cellular mass becomes converted within either the ducts or gland, or within both, into transparent, yellow, tough cement. Generally in Conchoderma, Pollicipes, and Scalpellum, two ovarian tubes, but in one specimen of _Conchoderma aurita_, three tubes, and in Ibla one tube could be seen running into or forming the gland; of the nature of the tubes there could not be the least doubt, for at a little distance from the glands they gave out branches (Pl. IX, fig. 3), containing ova in every state of development. In some specimens as in that figured of _Conchoderma aurita_, the ovarian tube on one side of the gland is larger than on the other, and has rather the appearance of being deeply embedded in the gland than of forming it; but, in other specimens, the two ovarian tubes first formed a little pouch, into which their cellular contents could be clearly seen to enter; and then this pouch expanded into the gland; thus quite removing a doubt which I had sometimes felt, whether the ovarian tube was not simply attached to or embedded in the gland, without any further connection. By dissection the multiple external coats of the gland and ovarian tubes could be seen to be continuous. The cellular contents of the tubes passed into the more opaque cellular contents of the gland, by a layer of transparent, pulpy, pale, yellowish substance. There appeared in several instances to be a relation, between the state of fulness and condition of the contents of the gland, and of the immediately adjoining portions of the ovarian tubes. In one specimen of _Pollicipes mitella_ it was clear that the altered, tough, yellow, transparent, non-cellular contents of the two glands and ducts, had actually invaded for some little distance, the two ovarian tubes which ran into them, thus showing the continuity of the whole. From these facts I conclude, without hesitation, that the gland itself is a part of an ovarian tube specially modified; and further, that the cellular matter, which in the ovarian tubes serves for the development of the ova, is, by the special action of the walls of the gland, changed into the opaquer cellular matter in the ducts, and this again subsequently into that tissue or substance, which cements the Cirripede to its surface of attachment.

As the individuals grow and increase in size, so do the glands and cement-ducts; but it seems often to happen, that when a specimen is immovably attached, the cementing apparatus ceases to act, and the cellular contents of the duct become converted into a thread of transparent tough cement; the investing membrane, also, of the ducts, in Conchoderma sometimes becomes hard and mamillated. I have already alluded to the case of a Pollicipes, in which both glands and ducts, and even a small portion of the two adjoining ovarian tubes, had become thus filled up. As in sessile Cirripedes, at every fresh period of growth a new cement gland is formed, it has occurred to me, that possibly in Pollicipes something similar may take place. In sessile Cirripedes, the old cement-glands are all preserved in a functionless condition, adhering to the membranous or calcareous basis, each new larger one attached to that last formed, and each giving out cement-ducts, which, bifurcating in the most complicated manner, pass outside the shell and thus attach it to some foreign body.

The cement, removed from the outside of a Cirripede, consists of a thin layer of very tough, bright-brown, transparent, laminated substance, exhibiting no structure under the highest powers, or at most a very fine dotted appearance, like a mezzotinto drawing. It is of the nature of chitine; but boiling caustic potash has rather more effect on it than on true chitine; and I think boiling nitric acid rather less effect. In one single instance, namely, in Coronula, the cement comes out of the four orifices of the two bifurcating ducts, in the shape of distinct cells, which, between the whale’s skin and the basal membrane, arrange themselves so as to make a circular, continuous slip of cement; then the cells blend together, and are converted into transparent, structureless cement. Cementing tissue or membrane would, perhaps, have been a more correct title than cement; but, in ordinary cases, its appearance is so little like that of an organised tissue, that I have for this reason, and for brevity-sake, preferred the simple term of Cement.

In the larva the cement always escapes through the prehensile antennæ; and it thus continues to do throughout life in most or all of the species of Lepas, Conchoderma, Dichelaspis and Ibla. In the first two of these genera, the cement escapes from the borders of the lower side of the disc or penultimate segment of the antennæ, and can be there seen radiating out like spokes, which at their ends divide into finer and finer branches, till a uniform sheet of cement is formed, fastening the antennæ and the adjoining part of the peduncle down to the surface of attachment. In _Dichelaspis Warwickii_ and _Scalpellum Peronii_, the cement, or part at least, comes out of the ultimate segment of the antennæ, in the shape of one tube, within another tube of considerable diameter and length. In _Scalpellum vulgare_, and probably in some of the other species, which live attached to corallines, the cement soon ceases to debouch from the antennæ, but instead, bursts through a row of orifices on the rostral margin of the peduncle (Pl. IX, fig. 7), by which means this margin is symmetrically fastened down to the delicate, horny branches of the zoophyte. In Pollicipes, the two cement-ducts, either together or separately (Pl. IX, fig. 2, 2 _a´_), wind about the bottom of the peduncle in the most tortuous course, at each bend pouring out cement through a hole in the membrane of the peduncle. In Ibla the lower part of the peduncle is internally filled by cement, and thus rendered rigid. In _Lepas fascicularis_ a vesicular ball of cement surrounding the peduncle is thus formed (Pl. I, fig. 6), and serves as a float! All these curious, special adaptations are described under the respective genera. How the cement forces its way through the antennæ, and often through apertures in the thick membrane of the peduncle, I do not understand. I do not believe, though some appearances favoured the notion, that the duct itself debouches and divides, at least this is not the case in Coronula, but only that the internal chord of cellular matter thus acts and spreads itself out; nor do I understand how, when the antennæ and immediately adjoining parts are once cemented down, any more cement can escape; yet this must take place, as may be inferred from the breadth of the cemented, terminal portion of the peduncle in Lepas and Conchoderma; and from the often active condition in old individuals of the cementing organs.

I have entered on this subject at some length, (and I wish I had space for more illustrations,) from its offering, perhaps, the most curious point in the natural history of the Cirripedia. It is the one chief character of the Sub-class. I am well aware how extremely improbable it must appear, that part of an ovarian tube should be converted into a gland, in which cellular matter is modified, so that instead of aiding in the development of new beings, it forms itself into a tissue or substance, which leaves the body[15] in order to fasten it to a foreign support. But on no other view can the structure, clearly seen by me both in the mature Cirripede and in the larva, be explained, and I feel no hesitation in advancing it. I may here venture to quote the substance of a remark made by Professor Owen, when I communicated to him the foregoing facts, namely, that there was a new problem to solve,--new work to perform,--to attach permanently a crustacean to a foreign body; and that hence no one could, _a priori_, tell by what singular and novel means this would be effected.

[15] The protrusion of the egg-bearing pouches in Cyclops and its kindred genera, outside the body, offers a feeble analogy with what takes place in Cirripedes. Professor Allman (‘Annals of Natural History,’ vol. xx, p. 7,) who has attended to the subject, says that the external egg-bearing pouches are “a portion of the membrane of the true ovaries:” if the membrane of these pouches had been specially made adhesive, the analogy would have been closer.

_Filamentary Appendages._--These have generally been considered to act as branchiæ; they occur at the bases of the first pair of cirri in Lepas, Alepas, Conchoderma, and in three species of Pollicipes: in Conchoderma there are similar appendages attached to the pedicels of the cirri (Pl. IX, fig. 4, _g-k_); and in the above three species of Pollicipes there is a double row of them on the prosoma: their numbers differ in different species (in some there being none) of the same genus, and even in different individuals of the same species; they are entirely absent in the majority of the genera. These facts would indicate that they are not of high functional importance; and they seem so generally occupied by testes (Pl. iv, fig. 5), that I suspect their function is quite as much to give room for the development of these glands, as to serve for respiratory purposes. With the exception of the four above-named genera, the mere surface of the body and of the sack must be sufficient for respiration: in _Conchoderma aurita_ the two great expansions of surface, afforded by the folded, tubular, ear-like projections, aid, as I believe, towards this end.

The shape of the body varies, owing to the greater or less development of the lower part of the prosoma, the greater or less distance of the first from the second pair of cirri, and of the mouth from the adductor scutorum muscle, (Pl. IX, fig. 4, and Pl. IV, 8 _a´_). In all the genera, the body is much flattened. I may here mention a few particulars about the muscular system. One of the largest muscular masses is formed by the adductor scutorum, and by the muscles which surround in a double layer (the fasciæ being oblique to each other) the whole of the upper part of the prosoma. From under the adductor, a pair of delicate muscles runs to the basal edge of the labrum, so as to retract the whole mouth, and two other pair to the integument between the mouth and the adductor, so as to fold it: again, there are other delicate muscles in some (for instance in _Lepas Hillii_) if not in all the Lepadidæ, crossing each other in the most singular loops, and serving apparently to fold the membrane between the occludent edges of the scuta. Within the prosoma there is a strong adductor muscle, running straight from side to side, for the purpose, as it appears, of flattening the body. The thorax, on the dorsal and ventral surfaces, is well furnished with straight and oblique muscles (without striæ), which straighten and curl up this part of the body. The muscles running into the pedicels of the cirri, cross each other on the ventral surface of the thorax; the muscles within the rami are attached to the upper segments of the pedicels. Finally, I may remark that the whole of the body and the cirri are capable of many diversified movements.