Read Ebook: A Monograph on the Sub-class Cirripedia (Volume 1 of 2) The Lepadidae; Or Pedunculated Cirripedes by Darwin Charles

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Beitr?ge zur Naturgeschichte der Rankenf?sser, s. 16, Tab. i, figs. 3, 4.

Mr. J. D. Dana, who has examined these organs in the larvae of Lepas, informs me in a letter, that in his opinion they "correspond with the inferior antennae, the superior being wanting, as in most Daphnidae." He continues--"I know of no case in which the inferior are obsolete when the superior are developed; but the reverse is often true." In position these antennae certainly correspond to the inferior and central pair of the larva in the first stage, which belong, as it would appear, to the first segment of the body; but judging from the drawing by Burmeister of the larva in the second stage, I am, in some respects, more inclined to consider that they correspond to the larger pair seen within the lateral horns of the carapace in the first stage.

Rev. B. L. King. Annual Report of B. Institution of Cornwall, 1848, p. 55.

M. Dujardin has lately discovered that the "Hypopi are Acari with eight feet, without either mouth or intestine, and which, being deprived of all means of alimentation, fix themselves at will, so as to undergo a final metamorphosis, and they become Gamasi or Uropodi." Here, then, we have an almost exactly analogous case. M. Dujardin asks--"Ought, therefore, the Hypopi to be called larvae, when, under that denomination, have hitherto been comprised animals capable of nourishing themselves?"

Of the internal organs, whilst the Cirripede is still within the larva, I have already mentioned the stomach with its pair of caeca: from the retracted position of the thorax and rudimentary abdomen, and consequently of the anus, compared with these parts in the larva, the alimentary canal is not above half its former length. There is, as yet, no trace of the filaments supposed by some to act as branchiae, at the base of the first pair of cirri. Nor could I perceive a trace of the testes or vesiculae seminales: the penis is represented by a minute, apparently imperforate projection. I have already briefly described the pair of large, gut-formed bodies in the larva, into the anterior ends of which the cement-ducts ran, and evidently derived their slightly opaque, cellular contents. At a very early age, before the young Cirripede can be distinctly made out, the posterior ends of these gut-formed bodies are absorbed, so as not to pass beyond the caeca of the stomach. When the young Cirripede is plainly developed within the larva, these bodies in a relatively reduced condition are still distinct near the caeca, and at the opposite or anterior end , they have branched out into a sheet of delicate inosculating tubes; these could be traced by every stage, until, in the young perfected Cirripede, they filled the peduncle as ordinary ovarian tubes. In the larva, the two gut-formed bodies or incipient ovaria keep of equal thickness from one to the other end, but in the mature Cirripede, the ovarian tubes in the peduncle and the small, glandular, grape-like masses, near the stomach-caeca, are connected only by a delicate tube; this I failed in tracing in specimens in the very immature condition of those now under description.

I have in this abstract treated the metamorphoses at greater length than I should otherwise have done, on account of the great importance of arriving at a correct homological interpretation of the different parts of the mature animal. In Crustacea, according to the ordinary view, there are twenty-one segments; of these I can recognise in the Cirripede, on evidence as good as can generally be obtained, all with the exception of the four terminal abdominal segments; these do not occur in any species known to me, in any stage of its development. If that part of the larva in front of the mouth, bearing the eyes, the prehensile antennae, and in an earlier stage two pair of antennae, be formed, as is admitted in all other Crustacea, of three segments, then beyond a doubt, from the absolute correspondence of every part, and even every coloured mark, the peduncle of the Lepadidae is likewise thus formed. The peduncle being filled by the branching ovarian tubes is no objection to this view, for I am informed on the high authority of Mr. J. D. Dana, that this is the case with the cephalo-thorax in some true Crustaceans, for instance, in Sapphirina. To proceed, the mouth, formed of mandibles, maxillae, and outer maxillae, correspond with the fourth, fifth, and sixth segments of the archetype Crustacean. Posteriorly to the mouth, we come, in the larva, to a rather wide interspace without any apparent articulation or organ, and then to the thorax, formed of six segments, bearing the six pair of limbs, of which the first pair differs slightly from the others. The thorax is succeeded by three small segments, differently shaped, with the posterior one alone bearing appendages; these segments, I cannot doubt, from their appearance alone, and from their apparent function of steering the body, are abdominal segments. If this latter view be correct, the thoracic segments are the six posterior ones of the normal seven segments, and there must be two segments missing between the outer maxillae and first thoracic pair of legs, which latter on this view springs from the ninth segment. Now, in a very singular Cirripede, already alluded to under the name of Proteolepas, the two missing segments are present, the mouth being actually succeeded by eight segments, and these by the three usual abdominal segments,--every segment in the body being as distinct as in an Annelid: hence in Proteolepas, adding the three segments for the mouth and three for the carapace, we have altogether seventeen segments, which, as I stated, is the full number ever observed in any Cirripede, the four missing ones being abdominal, and, I presume, the four terminal segments. That the cavity in which the thorax is lodged, in the larva and therefore in the mature Cirripede, is simply formed by the backward production of the carapace, does not require any discussion. The valves have no homological signification.

As we have just seen that the first pair of natatory legs is borne on the ninth segment of the body, so it must be with the first pair of cirri, which consequently correspond to the outer maxillipods of the higher Crustacea, and hence their difference from the five posterior pair, which correspond with the five, ordinary pair of ambulatory legs in these same Crustacea. The part of the body, which I have called the prosoma, that is the protuberant, non-articulated, lower part of the thorax , is a special development, either of the ninth segment, bearing the first pair of cirri, or of the segments corresponding with the organs of the mouth. The three abdominal segments of the larva are represented in the mature Cirripede, in the Order containing the Lepadidae, only by a minute, triangular gusset, let in between the V-shaped tergal arches of the last thoracic segment: in this gusset, small as it is, is seated the anus, and on each side the caudal appendages, often rudimentary and sometimes absent. In another order, I may remark, the cirri, of which there are only three pair, are abdominal.

I feel much confidence, that the homologies here given are correct. The cause of their having been generally overlooked arises, I believe, from the peculiar manner, already described, in which the animal, during its last metamorphosis, is internally almost intersected: even for some little time after discovering that the larval antennae were always embedded in the centre of the surface of attachment, I did not perceive, that this was the anterior end of the whole animal. The accompanying woodcut gives at a glance, a view of the homologies of the external parts: the upper figure is a Stomapod Crustacean, Leucifer of Vaughan Thompson, and the abdomen, which we know becomes in Cirripedes, after the metamorphosis, rudimentary, and therefore does not fairly enter into the comparison, is given only in faint lines: the lower figure is a mature Lepas, with the antennae and eyes, which are actually present in the larva, retained and supposed to have gone on growing. All that we externally see of a Cirripede, whether pedunculated or sessile, is the three anterior segments of the head of a Crustacean, with its anterior end permanently cemented to a surface of attachment, and with its posterior end projecting vertically from it.

CAPITULUM.

I will now proceed to a general description of the different parts and organs in the Lepadidae. 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 , 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, Paecilasma, 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, and is composed of numerous fine laminae. After the valves have been placed in acid, a residue, very different in bulk in different genera, is left, also composed of successive laminae 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 laminae thus edged 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 laminae 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.

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: 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.

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

The peduncle is lined within by three layers of muscles, longitudinal, transverse, and oblique, all destitute of the transverse striae, 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 , 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.

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.

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 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.

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, . 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 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 if not in all the Lepadidae, 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 , 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.

Double eye transverse or longest axis } 126/6000 of both together }

Supra-oesophageal ganglion, } longitudinal axis of } 45/6000

Infra-oesophageal ganglion, } transverse axis of } 120/6000

Infra-oesophageal ganglion, } longitudinal axis of } 114/6000

Proceedings of the Academy of Natural Sciences, Philadelphia. No. i, vol. iv, Jan. 1848.

In all the genera, the double eye is seated deep within the body; it is attached by fibrous tissue to the radiating muscles of the lowest part of the oesophagus, and lies actually on the upper part of the stomach; consequently, a ray of light, to reach the eye, has to pass through the exterior membrane and underlying corium connecting the two scuta, and to penetrate deeply into the body. In living sessile Cirripedes, vision seems confined to the perception of the shadow of an object passing between them and the light; they instantly perceived a hand passed quickly at the distance of several feet between a candle and the basin in which they were placed.

I am compelled to differ greatly from the account given by Prof. Steenstrup of the reproductive system in the Cirripedia, in his 'Untersuchungen ?ber das Vorkommen des Hermaphroditismus, ch. v, 1846;--a translation of which I have seen, owing to the great kindness of Mr. Busk. Mr. Goodsir has described what he considers the male of Balanus; but I have seen this same parasitic creature charged with ova, including larvae! From the resemblance of the larvae to the little crustacean described by Mr. Goodsir, in the same paper, as a distinct parasite, I believe the latter to be the male of his so-called male Balanus, and that all belong to the same species, allied to Bopyrus. This genus, as is well known, is parasitic on other crustacea; and it is a rather interesting fact thus to find, that this new parasite which is allied to Bopyrus, in structure, is likewise allied to it in habits, living attached to Cirripedia, a sub-class of the crustacea.

In 'M?ller's Archiv,' 1834, p. 467. I have already several times referred to M. Martin St. Ange's excellent Memoir, read before the Academy of Sciences, and subsequently, in 1835, published separately.

I may here mention, that in all sessile Cirripedes, the ovarian branching tubes lie between the calcareous or membranous basis and the inner basal lining of the sack, and to a certain height upwards round the sack: the true ovaria and the two ducts occupy the same position as in the Lepadidae.

The structure here described is quite conformable with that which we have seen in the larva; in the latter, two gut-formed masses of equal thickness extended from the caeca of the stomach to within the future peduncle, where the cement-ducts entered them, and where, after a short period, they were seen to expand into a mass of ovarian tubes. In the mature Cirripede, the cement-ducts can still be found united to the ovarian tubes in the middle of peduncle; and the cause of the wide separation of the true ovaria and ovarian tubes, can be simply accounted for by the internal, almost complete intersection of the animal, which takes place during the last metamorphosis.

M. Martin St. Ange, describes an orifice under the carina, by which he supposes the ova to enter the sack; this, after repeated and most careful examinations, I venture to affirm does not exist; on the contrary, I have every reason to believe that the ova enter the sack in the following curious manner. Immediately before one of the periods of exuviation, the ova burst forth from the the ovarian tubes in the peduncle and round the sack, and, carried along the open circulatory channels, are collected beneath the chitine-tunic of the sack, in the corium, which is at this period remarkably spongy and full of cavities. The corium then forms or rather resolves itself into the very delicate membrane separately enveloping each ovum, and uniting them together into two lamellae; the corium having thus far retreated, then forms under the lamellae the chitine-tunic of the sack, which will of course be of larger size than the last-formed one, now immediately to be moulted with the other integuments of the body. As soon as this exuviation is effected, the tender ova, united into two lamellae, and adhering, as yet, to the bottom of the sack, are exposed: as the membranes harden, the lamellae become detached from the bottom of the sack, and are attached to the ovigerous fraena. To demonstrate this view, an individual should have been found, with both the old and new chitine tunic of the sack, and with the lamellae lying between them; this, I believe, I have seen, but it was before I understood the full importance of the fact: a great number of specimens would have to be examined in order to succeed again, for the changes connected with exuviation supervene very quickly. I have, however, several times found the ova so loose under the sack, as to be detached with a touch from the ovarian tubes; and I have twice carefully examined specimens, which had just moulted, as shown by even the mandibles being flexible, in which the lamellae had not become united to the fraena, but still adhered to the newly-formed chitine tunic of the sack; in these, the ova were so tender, that they broke into pieces rather than be separated from the membrane of the lamella, itself hardly perfectly developed, for pulpy cellular matter adhered outside some of the ova. These and other facts are quite inexplicable on any other view than that advanced.

The delicate tunic lining the sack, and the integuments of the whole body, are regularly moulted. With these integuments, the membrane lining the oesophagus, the rectum, and the deep olfactory pouches, and the horny apodemes of the maxillae, are all cast together. I have seen a specimen of Lepas, in which, from some morbid adhesion, the old membrane lining one of the olfactory pouches had not been moulted, but remained projecting from the orifice as a brown shrivelled scroll. The new spines on the cirri are formed within the old ones; but as they have to be a little longer than the latter, and as they cannot enter these up to their very points, their basal portions are not thus included, but are formed, running obliquely across the segments of the cirri; and what is curious, these same basal portions are turned inside out, like the fingers of a glove when hastily drawn off. After the exuviation of the old spines, the new spines have their inverted basal portions drawn out from within the segments, and turned outside in, so as to assume their proper positions.

Comparing the pedunculated and sessile Cirripedes, it is, I think, impossible to assign them a higher rank than that of Families. The chief difference between them consists, in the Lepadidae, in the presence of three layers of striae-less muscles, longitudinal, transverse and oblique, continuously surrounding the peduncle, but not specially attached to the scuta and terga; and on the other hand, in the Balanidae, of five longitudinal bundles of voluntary muscles, with transverse striae, fixed to the scuta and terga, and giving them powers of independent movement. In the Lepadidae, the lower valves, or when such are absent, the membranous walls of the capitulum, move with the scuta and terga when opened or shut; and the lower part of the capitulum is separated by a moveable peduncle from the surface of attachment; in the sessile Cirripedes, the lower valves are firmly united together into an immovable ring, fixed immovably on the surface of attachment. I will not compare the softer parts, such as the cirri and trophi, of the Lepadidae with those of the Balanidae, as my examination of this latter family is not fully completed: I will only remark, that there is a very close general resemblance, more especially with the sub-family Chthamalinae.

ANATIFERA. et plerumque Auctorum Anglicorum.

Valves 5, approximate: carina extending up between the terga, terminating downwards in an embedded fork, or in an external disc: scuta sub-triangular, with their umbones at the rostral angle.

Filaments seated beneath the basal articulation of the first cirri; mandibles with five teeth; maxillae step-formed; caudal appendages uniarticulate, smooth.

The prosoma is well developed. The stomach is surrounded in the upper part by a circle of large branching caeca. The generative system is highly developed; the testes coating the whole of the stomach, entering the filamentary appendages and the pedicels of the cirri; the two ovigerous lamellae contain a vast number of ova; they are united to rather large fraena, of which the sinuous margin supports either a continuous row or separate tufts of glands.

ANATIFA vel ANATIFERA vel PENTALASMIS laevis, plerumque auctorum.

I have used, in conformity with botanists, the mark of interjection, to show that I have seen an authentic specimen.

Valves smooth, or delicately striated. Right-hand scutum alone furnished with an internal umbonal tooth: uppermost part of peduncle dark-coloured.

Filaments, two on each side.

Var. . Fig. 1. Scuta and terga with one or more diagonal lines of dark greenish-brown, square, slightly depressed marks.

Var. . Carina strongly barbed.

Extremely common; attached to floating timber, vessels, sea-weed, bottles, &c., and to each other, in the Atlantic Ocean, Mediterranean, West Indies, Indian Ocean, Philippine Archipelago, Sandwich Islands, Bass's Straits, Van Diemen's Land.

Mr. W. Thompson found that 15 specimens, out of about 200, attached to a vessel which came from New Orleans into Belfast, had their carinas barbed.

In museums a vast amount of difference is seen in the colours of this species, caused by the method of preparation: if dried without having been in spirits, and subsequently kept dry, the orange tint round the orifice is preserved; if kept long in spirits, this is quite lost; but sometimes in specimens in spirits the colour of the membrane of peduncle is preserved and rendered pinker. The colours of the sack and animal are either quite discharged or rendered extremely dark. The valves themselves also often become more opaque. In some specimens well preserved in spirits, the sack and cirri were purplish-brown or lead-colour, tinted with dirty green, or orange, or bright yellow, or brick-red.

ANATIFA vel PENTALASMIS LAEVIS plerumque auctorum.

Valves smooth; scuta destitute of internal umbonal teeth; carina standing a little separate from the other valves, with the fork not close to the basal margin of the scuta; uppermost part of peduncle either pale or orange-coloured.

Filaments three on each side.

The chitine membrane at the base of the capitulum, especially at the anterior and posterior ends, is covered with beautiful, little, embedded, yellowish beads, about 3/2000th of an inch in diameter; above this, on each side of the carina, there is a space with similar but smaller little spheres, and still higher up still minuter ones; others occur on different parts of the capitulum; these spaces are seen to be distinctly separated from each other, and present a beautiful appearance under a high power.

Valves approximate, slightly furrowed, especially the terga; right-hand scutum with a strong internal umbonal tooth; left-hand with a small tooth, or mere ridge; occludent margin arched, protuberant: uppermost part of peduncle orange-coloured.

Filaments five or six on each side.

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