7. Anelasma

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_ ANELASMA. Gen. Nov. Pl. IV.

ALEPAS. Lovén. Ofversigt of Kongl. Vetenskaps-Akad. Fördhandlinger: Forsta Argangen. Stockholm, 1844, p. 192, Tab. 3.

Capitulum sine valvis: aperturâ amplâ: pedunculus fimbriatus, sub-globosus, infossus. Capitulum without valves; aperture large; peduncle fimbriated, sub-globular, imbedded. Cirri without spines; outer maxillæ and palpi rudimentary, spineless; mandibles minute, with several small teeth irregularly placed; maxillæ minute, with very minute irregularly scattered spines. No caudal appendages. * * * * * I owe to the great kindness of Professor Steenstrup, an examination of this very curious cirripede, well described and figured by Lovén, who considered it an Alepas. It lives parasitic, with its peduncle imbedded in the skin of sharks, in the North Sea. According to the principles of classification which I have followed, this cirripede cannot possibly remain in Alepas, and must form a new genus; for some time, indeed, I thought that a new family or sub-family ought to have been instituted for its reception; but when I considered that its highly peculiar characters are all negative, as the non-articular, non-spinose structure of the cirri, and that no new or greatly modified functional organ is present, I concluded that it might properly remain amongst the Lepadidæ. We shall, moreover, hereafter see that the male of Ibla, which, of course, must remain in the same family with the female, is, in some analogous respects, even more abnormal than Anelasma. 1. ANELASMA SQUALICOLA. Pl. IV, figs. 1-7. ALEPAS SQUALICOLA. Lovén, ut supra.

North Sea. Parasitic on Squalus.

Capitulum, destitute of valves; oval, much flattened; the double membrane composing it, thin, highly flexible, coloured externally and internally, by the underlying corium, of a blackish purple; aperture, extremely large, extending from the upper end of the capitulum, to close above the peduncle, gaping, and not protecting (in the dead condition) the cirri and mouth. The Peduncle is about half as long as the capitulum, but, according to Lovén, this part varies in length; it is a little narrower than the capitulum; colourless, from being imbedded in the shark's skin; sub-globular; basal end almost hemispherical. Total length of animal 1.3; diameter of peduncle .4 of an inch. The external membrane of the capitulum is not nearly so thick as is usual in other Cirripedes, and is, therefore, unusually flexible. The internal membrane, on the other hand, is very much thicker than is usual, being only a little thinner than the outside coat; this circumstance, as well as the similarity in colour on both sides, is evidently due to the remarkable openness of the sack, and consequent exposure of its inside. The inner membrane, when viewed under a high power, is seen to be covered with the minutest spines; the external membrane is structureless, except that there are a few rows of very minute beads of hard chitine, like those which occur on the capitulum of Conchoderma aurita. Lovén, however, states that there are imbedded in the outer membrane, scattered, minute, dendritic, calcareous particles. Of these, I could see no trace. There is a very thin muscular layer between the two coats, all round the capitulum, and this layer becomes rather thicker round the base, near the peduncle. The adductor muscle, occupying its usual place close below the mouth, is thinner than in any other Cirripede of the same size seen by me; nor does it end so abruptly at each extremity, as is usual: where attached to the outer coat, no impression is left. It is a singular fact, that in this Cirripede alone, the fibres of the adductor, and of the muscles of the cirri, and of the trophi of the mouth, are destitute of transverse striæ; but it is not singular, that the muscles surrounding the capitulum should, also, be destitute of striæ, for this is the case with the muscles which, running up from the peduncle, surround the capitulum in Alepas, and partly surround it in Conchoderma. It must not be inferred from the absence of transverse striæ in the muscular fibres of the adductor and of the cirri and trophi, that they are involuntary, but only that they are in an embryonic condition, for I find in the natatory larva, that all the muscles, with the exception of some connected with the eyes, are similarly destitute, and yet perform voluntary movements.[46]

[46] Dr. C. Schmidt in his Contribution to the comparative Anatomy of the Invertebrate animals, &c., (translated in Taylor's Scientific Memoirs, vol. v, p. 1,) says that in young Crustacea, "we find plain primitive fibres, which subsequently acquire the transversely striated aspect."

Although in the dead state, the aperture of the capitulum seems to be always gaping, yet I have little doubt, that the living animal can fold the flexible membrane, like a mantle, round its thorax and cirri, and thus protect, though feebly compared with most Cirripedes, these organs. I suspect that the mouth is always exposed. Peduncle.--The membrane of the peduncle is thin; the whole surface is sparingly and quite irregularly studded with minute, much-branched filaments (Pl. IV, fig. 3, highly magnified); these are occasionally as much as l/5th of an inch in length; the degree of branching varies much, but is generally highly complex; the ordinary diameter of the branches is about 1/200th of an inch; their tips are rounded, and even a little enlarged, and frequently torn off, as if they had been attached to or buried in the flesh of the shark, in which the whole peduncle is imbedded. These filaments are formed of, and are continuous with the external transparent membrane of the peduncle, and they contain, up to the tips of every sub-branch, a hollow thread of corium, prolonged from the layer internally coating the whole peduncle. In all other Lepadidæ, the peduncle increases in length, chiefly at the summit where joined to the capitulum, and in diameter, throughout nearly its whole length, except close to the base; but, owing to the constant disintegration of the outer surface, the old outside coat does not split in defined lines, like the membrane of the capitulum. In Anelasma, however, owing to the imbedded position of the peduncle, the old outer coats are preserved, the lines in which they have split during continued growth being thus exhibited: those in the uppermost part almost symmetrically surround the peduncle, showing that here, as in other Lepadidæ, has been one regular line of growth; but in the lower part the lines are extremely irregular; and what is almost unique, it appears that the blunt basal end is constantly increasing in length and breadth, and, apparently, at a greater rate than any other part. I judge of this latter fact, from the whole bottom of the peduncle being covered with numerous curved, or nearly circular, lines of natural splitting, the nature of which can be best understood by examining the much-enlarged drawing (Pl. IV, fig. 3) of a small portion (taken by chance) of the membrane of the base, seen from the outside, and bearing some of the simplest branched filaments: other branches, as may be seen, have been cut off. This manner of growth explains the broad, blunt basal termination of the peduncle, so unlike that in other Lepadidæ. New membrane is formed, not continuously as in other cases, under the whole surface of the old membrane, but in irregular patches; thus the portion marked (a) runs under (b), but not under the little circles (c, c), for these are the last-formed portions and underlie the membrane (a) and (b). I do not understand how the splitting of the old membrane is effected; but no doubt it is by the same process by which the membrane of the capitulum in other genera, as in Scalpellum, splits symmetrically between the several valves. In the branched filaments it is particularly difficult to understand their growth, for it is not possible, after examining them, to doubt that they continue to increase, and send off sub-branches, which it would appear probable, penetrate the shark's flesh like roots. I may remark that one, or more commonly two or three branched filaments stand nearly in the centre of each circular line of exuviation or splitting. The branched filaments first commence as mere little pustules, and these appear to be most numerous at the bottom of the peduncle. The final cause of the downward growth of the bottom of the peduncle, is obviously to allow of the animal burying itself in the shark's body, in the same way as Coronula and Tubicinella become imbedded by the downward growth of their parietes in the skin of Cetacea. The only other genus of Lepadidæ, in which the growth of the peduncle is at all analogous, is Lithotrya, in this genus, however, the animal burrows mechanically into soft rock or shells. I looked in vain for cement, or for the cement-glands, (but the specimen was in an extremely unfavorable state for finding the latter) or for the prehensile antennæ of the larva. No doubt this Cirripede at first becomes attached in the same way as others, but after early life, I suspect it is retained in its place, by being so deeply imbedded in the shark's body, and perhaps by the root-like branched filaments. The irregular growth and splitting of the membrane at the base of the peduncle, where the prehensile antennæ of the larva must originally have been situated, would account for not finding them. The inside of the peduncle (fig. 2 g) was gorged, in the specimen examined by me, with immature ova. The innermost muscular layer consists of longitudinal bundles of unusual size, but placed rather far apart from each other; these do not extend to the very base of the peduncle, and at the upper end they curve inwards, almost to the middle of the under side of the diaphragm, separating the peduncle and capitulum. Outside these longitudinal muscles, there are delicate transverse ones, but apparently there are no oblique muscles in the upper part of the peduncle, as in other Lepadidæ; near the bottom, the transverse muscles form a thicker layer with many of the bundles running in oblique lines. Mouth.--uLovén has not described this part quite accurately, owing to his not having used high enough magnifying powers. He states that the trophi are soft and functionless, which is far from the case. The whole mouth (fig. 2 d), is unusually small; it is, to a certain extent, probosciformed, and being curved a little downwards, projects slightly over the adductor muscle, to which it is closely placed. The labrum does not project more beyond the general surface of the body, than in many other Cirripedes, but the probosciformed structure is caused by the elongation of the surface fronting the thorax. The summit of the mouth stands above the level of the top of the pedicels of the first pair of cirri. The labrum is slightly hollowed out in the middle of its upper margin; it can scarcely be called bullate, in which it differs from all other Lepadidæ; on the other hand, the outer and inner folds of the labrum are not so close together as in Balanus. On each upper corner, there is, as usual, a small rounded prominence, close to which there is a second slight, rounded, spineless swelling; these latter represent the quite rudimentary Palpi. The Mandibles (figs. 4, 5) are more highly developed than the other trophi; they are, however, very minute, the toothed edge being only about 16/1000th of an inch in length, measured in its longest direction; the edge is unusually thick, with the teeth placed rather on one side; this organ, when viewed on the labrum side (fig. 5), shows two large teeth placed low down, with the inferior angle pectinated and broadly truncated; but when viewed on the other or maxillæ side (fig. 4), several large and small teeth, placed alternately and irregularly in pairs, are seen extending along the whole edge. The mandibles are furnished, as usual, with three principal sets of muscles attached to the basal fold of the mouth. The Maxillæ (fig. 7) are still smaller than the mandibles; the spinose edge being only the 1/100th of an inch in length; the edge, instead of being square, and furnished with a double row of long spines, as in all other Cirripedes, is rounded, thick, club-shaped, and with the side facing the mandibles, thinly and irregularly strewed with short, thick, very minute spines; there is a large broad apodeme (a), in the usual place, but it is much more transparent and flexible than common: there are also the usual muscles. In other cirripedes, the mandibles alone seem to force the prey down the oesophagus; but here, the mandibles and maxillæ equally stand over the orifice, and their adjoining spinose faces and edges, seem excellently adapted to force, by their united action, any minute living creature down the passage. The Outer Maxillæ are almost in as rudimentary a condition as the palpi; they are quite spineless; viewed externally, they appear like two smooth, blunt, very minute projecting points; but viewed internally, the membrane forming the supra-oesophageal hollow seems to be united actually to their tips, so that they do not project at all. I was surprised to find that the longitudinal muscles going to these organs were developed, in proportion to the other muscles, quite as fully as in ordinary cirripedes: hence, these two little outer maxillæ, no doubt, serve as an under lip, and possess the usual backward and forward movement. The surface of the probosciformed mouth facing the first pair of cirri, has a deep central longitudinal fold, and rather more than half-way down, a transverse fold; just above this latter fold, and therefore quite below the outer maxillæ themselves, the two olfactory orifices are seated; these are unusually large, and the sack into which they lead, is most unusually large and deep. In this Cirripede, I was first enabled to observe that the membrane lining the sack is tubular, and open at the bottom. Cirri.--There are, as usual, six pair, and not of very small size; they have a shapeless and rudimentary appearance; they are coloured, like the rest of the body, blackish purple: they are quite spineless, and not articulated, but their anterior faces are either obscurely or very plainly lobed, so that in some (for instance in the third pair, Pl. IV, fig. 6), nine or ten prominent steps could be counted, manifestly representing so many segments. The rami are equal in length in the first pair, and slightly unequal in the second and third pair; these two latter are longer than either the first or three posterior pair. There is a small interspace as usual between the first and second pair of cirri. Internally, the cirri are occupied, even up to their tips, by delicate striæ-less muscles. The external membrane of the thorax and limbs, when examined under a very high power, is seen to be covered with minute toothed scales, as in most Cirripedes. The thorax is articulated as usual: the posterior part, however, is smaller, and tapers more suddenly than in other species, and this corresponds with the smaller size and more rudimentary condition, of the three posterior pair of cirri, compared with the anterior pair. The prosoma is hardly at all developed. The orifice (Pl. IV, fig. 2 e) of the acoustic (?) sack, beneath the first cirrus, is unusually large. There are no filamentary appendages. Alimentary Canal.--The membrane lining the oesophagus is unusually thin: it is furnished with the ordinary constrictor muscles, and others radiating from them like spokes of a wheel. The stomach is lined by unusually prominent biliary folds, which in the duodenum are transverse, sending forth, however, short folds at right angles; and these latter, in the proper stomach, become so much developed that the folds appear longitudinal. The rectum extends inwards, about as far as the base of the fourth pair of cirri, but is very short, owing to the little development of the three posterior segments of the thorax. The anus is seated in its usual place, at the dorsal basis of the penis, and is hidden by loose folds of skin; but there are no distinct caudal appendages. The stomach, in the specimen examined, was quite empty. Reproductive Organ.--The penis (fig. 2, c) is thick, short (about twice as long as the sixth cirrus), constricted at the base, ringed, spineless, with the terminal aperture large; internally it is well furnished with muscles. The two vesiculæ seminales, appeared to be unusually small; and one was much smaller than the other; they do not (I believe) become united into a common tube, till near the apex of the penis. They were empty; and, I presume, from the state of the ova, that their contents had lately been discharged. The whole thorax was filled with a white, fibrous and cellular mass, consisting perhaps of the testes in their undeveloped state. The individual dissected by me, appeared to have been defective in its last act of reproduction, for there were only two or three ova attached to the frænum on one side, and not very many on the other. The ova are much less elongated than is usual; they are of a remarkable size, namely 22/1000ths of an inch in their longer diameter; the membrane by which they are united into a pair of lamellæ is remarkably strong; the frænum (Pl. IV, fig. 2 f) on each side is large, strong, with rounded edges, pale coloured and hence conspicuous; on the side nearest the body, the whole surface is covered with club-shaped glands, having very short footstalks, and being in total length 5/6000ths of an inch; these glands secrete a reticulated layer of gut-formed fibres, attached to the ovigerous lamellæ. In the specimen described by Lovén, the lamellæ (fig. 1, and fig. 2, b, b) appear to have been very large: and in that examined by myself, the peduncle was gorged with immature ova, showing that the female reproductive powers were ample, though at the foregoing period, only a few eggs had been formed. Habits.--According to Lovén, this species lives imbedded in the skin of Squalus maximus and spinax, in the North Sea: I suspect that it is not closely compressed in its cavity, otherwise, I do not see the use of the two layers of muscles round the whole peduncle; it probably adheres to the sides of the cavity by the tips of the branched, root-like filaments; owing to the flexible nature of the capitulum, this Cirripede can offer little resistance to the water, and, therefore, is little likely to be torn out of its cavity. I have no doubt that it can fold the membrane of the capitulum, like a cloak, round its thorax and cirri; but it certainly can offer far less resistance, than other Cirripedes, to any enemy. This creature must obtain its food, and considering its productiveness much food must be required, in a manner quite different from nearly every other member of its Order. As the whole of the peduncle is imbedded, and as the mouth is probosciformed, with the labrum a little curled over the adductor muscle, I conclude that this Cirripede can reach minute animals crawling by on the surface of the shark's body. It must be borne in mind that the mouth, as in all Cirripedes, has the power of independent movement, and that the mandibles and maxillæ are here beautifully adapted to catch and force down any small living creature into the muscular oesophagus; the rudimentary outer maxillæ, moreover, no doubt have the power of scraping, like a lip, anything towards these prehensile organs. It will hereafter be seen, that the male of Ibla Cumingii, in which the cirri are quite rudimentary, obtains its food in a somewhat analogous manner, though in this case the whole peduncle moves, and not merely a probosciformed mouth: it deserves attention, that in the male Ibla and in Anelasma, in neither of which the cirri are prehensile, the palpi are rudimentary and useless. I am tempted to believe, that the largely developed olfactory sacks, and perhaps, likewise, acoustic (?) sacks, in Anelasma, replace, by giving notice of the proximity of prey, the loss of tactile cirri. It should be remembered that all Cirripedes subsist on animals which happen to swim or float within reach of the cirri; but here it is only those which happen to crawl within reach of the probosciformed mouth. It would, however, be rash to assert that the cirri in Anelasma, considering their muscular though feeble structure, may not be of some slight use, when thrown over the prey, in preventing its escape. Professor Steenstrup informs me that, from late observations, it appears that this animal always adheres to the shark's body in pairs. I regret extremely that I have not been able to examine a pair: that the individual examined by me was bisexual, I can hardly doubt, though the male organs certainly were feebly developed; it appears probable, that the individual described by Lovén was likewise bisexual: but after the facts presently to be revealed regarding the sexes in Ibla and Scalpellum, it is quite possible that the male and female organs may be developed in inverse degrees in different and adjoining individuals. The genus Anelasma is, I think, properly placed between Alepas and Ibla. In several of its characters, such as the absence of calcareous valves, the broad blunt end of the peduncle, the spineless cirri, the small size of the trophi, and more especially the absence of transverse striæ in those muscles, which in mature cirripedes are thus furnished, we see that this genus is in some degree in an embryonic condition. _

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