Alepas. Sander Rang. Manuel des Mollusques, 1829.
Anatifa. Quoy et Gaimard. Voyage de l’Astrolabe, 1834.
Triton. Lesson. Voyage de la Coquille, 1830.
Cineras. Lesson. Secundum Sander Rang.
Capitulum aut sine valvis, aut scutis corneis, pæne abditis.
Capitulum without valves,[41] or with horny, almost hidden, scuta.
[41] Any one not attending to the characters derived from the softer parts of the Balanidæ and Lepadidæ, might easily confound with Alepas the genus Siphonicella (genus nov.), which, undoubtedly, though having the external appearance of a pedunculated cirripede, belongs to the Balaninæ, and is closely related to Coronula.
Filaments seated beneath the basal articulations of the first pair of cirri; mandibles, with two or three teeth; maxillæ notched, with the lower part irregular, projecting; caudal appendages multi-articulate.
Attached to various living objects, fixed or floating.
Capitulum either entirely destitute of valves, or with transparent horny scuta, not containing any calcareous matter, and almost hidden in membrane. These scuta are formed of a lower and a lateral lobe, placed at above right angles to each other; they are added to by successive layers, and closely resemble in shape the scuta of the Conchoderma aurita. The orifice in A. tubulosa projects so much as to be almost tubular. In A. parasita and A. minuta it does not project, and is either moderately large, or very small in proportion to the length of the capitulum; from contraction it is much wrinkled. The membrane forming the capitulum is smooth and very transparent; it contains very few tubuli, except under certain irregular projections in A. cornuta.
The Peduncle is rather short and narrow; it blends into the capitulum, and is not, in some of the species, separated from it by any distinct line; the surface of attachment is rather wide. Within the peduncle we have the three usual layers of striæ-less muscles; namely, the innermost and longitudinal, which run lower down than the others; the middle and transverse; and, lastly, the exterior, oblique muscles, which cross each other (becoming transparent) on the rostral central line. These several muscles run up from the peduncle and surround the capitulum; from the transparency of the membranes they can be seen from the outside: they are particularly conspicuous round the orifice, which they probably serve to close. There is, in all cases, the usual adductor scutorum muscle (with transverse striæ), which is attached under the horny scuta, where such exist. The fact of the striæ-less muscles of the peduncle surrounding the whole capitulum, has been observed only in one other genus, namely Anelasma. In consequence of this structure, the capitulum must possess considerable powers of contraction.
The antennæ of the larva in the Alepas cornuta and A. minuta have the sucking disc nearly circular, with the spines unusually plain on the distal as well as proximal margin. Basal segment broad, much constricted where united to the disc. The ultimate segment has on the middle of the outer margin, in A. cornuta, two minute spines, which I have not observed in any other cirripede: on the summit there are the usual spines.
Size.—Three of the species are small.
Filamentary Appendages.—These are rather small; there is only one on each side, situated on the posterior margin of a slight swelling, beneath the basal articulation of the first cirrus; and therefore in the position in which the filaments are most constant in Lepas, and where they likewise occur in Conchoderma.
Body.—The prosoma is either pretty well developed or is small, according as the first cirrus is placed near to, or far from the second cirrus.
Mouth.—Labrum moderately bullate, with the lower part more or less produced; crest with blunt, bead-like teeth, and short hairs.
Palpi (Pl. X, fig. 8), acuminated and narrow to an unusual degree.
Mandibles, with two or three teeth, and the inferior angle acuminated; the lateral bristles unusually strong, so as to give the main teeth the appearance of being pectinated.
Maxillæ, widely notched, with three great upper spines; the part beneath the notch projecting, and either straight or irregular.
Outer Maxillæ, with the inner bristles either continuous or divided into two groups: exteriorly there is a smaller or larger prominence, with long bristles. The olfactory orifices are either slightly, or not at all protuberant.
Cirri.—In the three posterior pair, the segments have their bristles arranged in a transverse row, either in the form of a narrow brush, or consisting only of a single pair with two or three minute, intermediate, and lateral marginal spines. The anterior ramus of the second cirrus is thicker, and more thickly clothed with spines than is the posterior ramus: this latter ramus, however, and both rami of the third cirrus, are rather more thickly clothed with spines than are the three posterior pair. The unique case in A. cornuta of the inner rami of the fifth and sixth cirri being rudimentary (Pl. X, fig. 28) will be minutely described under that species.
Caudal Appendages, thin, tapering, multi-articulate, about as long as the pedicels of the sixth cirrus.
Stomach.—The œsophagus runs in a somewhat sinuous course, and enters the top of the stomach obliquely. There are no cæca. The biliary envelope presents a reticulated structure, instead of the usual longitudinal folds.
Generative System.—The penis is hairy, not very long, and ringed or articulated in an unusually plain manner; the space between each ring being about one fourth of the diameter of the penis: the unarticulated basal portion or support is here remarkably long. The vesiculæ seminales are long, tortuous, and enter the prosoma. The ovarian tubes are of wide diameter: in A. cornuta they surround the whole capitulum. The ovigerous fræna are small, constricted at the base, and square on the free margin, which is studded with minute glandular beads, borne on the finest footstalks.
Range.—Southern shores of England, Mediterranean, Atlantic, West Indies, New Zealand, attached to various objects. A. parasita has been always taken on Medusæ.[42]
[42] It appears that Solander (Dillwyn Des. Cat., vol. i, p. 34) observed a species of this genus adhering to a Medusa on the coast of Brazil. Mr. Cocks informs me that an Alepas, apparently A. parasita, has been cast on shore near Falmouth, attached to a Cyanæa; and that two other specimens adhered to the bottom of a vessel arriving at that port from Odessa.
Affinities.—This genus differs from all, except Anelasma, in the manner in which the striæ-less muscles of the peduncle run up and surround the capitulum, and likewise in the reticulated character of the biliary envelope of the stomach. To Conchoderma, especially to C. aurita, there is manifest affinity in the form of the horny scuta: there is also some affinity to this same genus in the presence of filamentary appendages though here little developed, and in the circular form of the disc of the larval antennæ, and, lastly, in the ovarian tubes in A. cornuta surrounding the capitulum. There is quite as close, if not closer affinity to Ibla, in the following peculiarities,—in the curved œsophagus,—in the general character of the cirri and trophi, with the olfactory orifices in one species in some degree prominent,—in the multi-articulated caudal appendages,—and in the plainly-articulated penis, with its elongated unarticulated support, though both these characters are exaggerated in Ibla. Lastly, the scuta in Ibla, though not at all resembling in shape those of A. cornuta, are formed without calcareous matter; and again, in Ibla, the muscles of the peduncle run up to the bases of the valves, and so almost surround the space in which the animal’s body is lodged.
The four species of Alepas appear to form two little groups; viz. A. parasita and A. minuta on the one hand, and A. cornuta and A. tubulosa on the other.
1. Alepas minuta. Tab. III, fig. 5.
Alepas minuta. Philippi. Enumeratio Mollusc. Siciliæ, 1836, Tab. xii, fig. 23.
——— ——— A. Costa. Esercitazione Accadem., vol. ii, part I, Naples, 1840, Pl. iii, fig. 5 (secundum Guerin in Revue Zoolog., 1841, p. 250.)
——— ——— Chenu. Illust. Conch., Pl. iii, figs. 8-10.
A. aperturâ non prominente, capituli longitudinis vix tertiam partem æquante: scutis corneis, pæne absconditis: longitudine totâ ad quartam unciæ partem.
Orifice not protuberant, one third of the length of the capitulum: scuta horny, almost hidden. Total length quarter of an inch.
Outer maxillæ, with the spines in front continuous; posterior cirri, with several long spines arranged in a transverse row on each segment; caudal appendages longer than the pedicels of the sixth cirrus.
Sicily; attached to a Cidaris:[43] island of Capri (A. Costa).
[43] I am greatly indebted to Professor J. Müller, of Berlin, for kindly lending me specimens.
Capitulum oval, blending insensibly into the peduncle; moderately flattened; composed of thin structureless membrane, with the exception of two horny, almost quite hidden scuta. Orifice situated near the summit, and in a line, which is oblique to the longitudinal axis of the peduncle; much wrinkled; barely one third of the length of the whole capitulum.
The Scuta, consist of yellowish, transparent, horny, laminated chitine, without any calcareous matter; externally covered by the common integument of the capitulum; these valves are placed very near to each other, close under the orifice, and therefore high up on the capitulum; the membrane between them is smooth and unwrinkled; they are formed of two rather acuminated lobes, joining each other at above a right angle; one lobe (the longer one) stretching nearly transversely across the capitulum, the other running down parallel to its rostral margin: in shape and position they resemble the scuta of Conchoderma aurita; and if another lobe had been developed it would have run along the orifice, and then these valves would have resembled the scuta of Conchoderma virgata. In a specimen with a capitulum 2/10ths of an inch long, the scuta from point to point were 1/20th of an inch in length.
Peduncle, much wrinkled, about one third in diameter of the capitulum, and shorter than it; at the base it is generally expanded into two or three finger-like projections. Length of the largest specimen, about one fourth of an inch. Colour, according to A. Costa in the work above cited, “rufo-flava vittatâ;” but after spirits the whole becomes uniformly yellowish.
Filamentary Appendages, situated beneath the basal articulation of the first cirrus, on the posterior edge of the usual enlargement; acuminated, about two thirds of the length of the shorter ramus of the first cirrus.
Prosoma well developed.
Mouth.—On each side there are two slight prominences; one under the mandibles, the other transverse nearer to the adductor muscle.
Labrum, placed near the adductor muscle, with the upper part not more bullate than the lower part; crest with a row of blunt teeth, and many fine bristles growing chiefly outside the teeth; there are many fine bristles on the inner or supra-œsophageal fold of the labrum.
Palpi not nearly touching each other, pointing towards the adductor: much hollowed out on their inner sides, hence narrow and acuminated, with doubly serrated bristles.
Mandibles, with three teeth and the inferior angle ending in a single sharp spine; whole inferior portion narrow; first tooth as far from the second, as the latter from the inferior angle; owing to the presence of short thick spines projecting from the sides of the jaw, the lower edges of the second and third teeth appear pectinated.
Maxillæ, nearly two thirds of the width of the mandibles; beneath the three larger upper spines there is a considerable notch, and the whole lower part is very slightly upraised; edge irregular, with obscure traces of either two projections, or perhaps of four steps.
Outer Maxillæ, with bristles in front continuous; exteriorly there is a slight prominence near each olfactory orifice, with a tuft of long bristles.
Cirri not much elongated; first pair placed not quite close to the second; five posterior cirri nearly equal in length; pedicels long, with irregularly scattered spines,—those on the pedicel of the first cirrus beautifully and conspicuously feathered. The segments of the three posterior pair are not very short or broad; very slightly protuberant, each with a long transverse, crescentic, narrow brush of bristles, which stand two or three deep in the middle, but on the sides are single: dorsal tufts long, and in the upper segments the spines are thick and claw-like. This structure is common to all the cirri. First cirrus with the rami unequal in length by two segments; from the shortness of the pedicel, this cirrus is much shorter than the second, but its rami are about two thirds of the length of those of the second cirrus. Second cirrus (and in a less degree the third cirrus), with the anterior ramus a shade broader than the posterior ramus, and rather more thickly covered with spines than are the three posterior cirri. Fifteen segments in the sixth cirrus; nine in the longer ramus of the first cirrus.
Caudal Appendages, rather longer than the pedicels of the sixth cirrus, composed of seven cylindrical, tapering segments, each with a circle of very fine bristles on its summit.
The acoustic (?) sacks are situated some way below the basal articulations of the first cirrus.
2. Alepas parasita.
Alepas parasita. Sander Rang. Man. des Mollusq., p. 364, Pl. viii, fig. 5, 1829.[44]
Anatifa univalvis. Quoy et Gaimard. Annales des Sciences, Nat., tom. x, p. 234, 1827, Pl. vii, fig. 8.
——— parasita. Quoy et Gaimard. Voyage de l’Astrolabe, Pl. xciii, 1834.
Triton (Alepas) fasciculatus. Lesson. Voyage de la Coquille. Mollusc. Pl. xvi, fig. 6, tom. ii, part I, 1830, p. 442.
[44] M. Sander Rang rejects the specific name “univalvis,” as signifying a generic character, and he has been followed in this by MM. Quoy and Gaimard themselves. This, according to the Rules of the British Association, would hardly have been a sufficient reason, but it appears that A. parasita, like A. minuta, has a pair of horny scuta or valves; and, therefore, the name univalvis is too obviously false to be retained. With respect to the generic name Triton, I fully believe that it was applied by Linnæus to the cast-off exuviæ of sessile Cirripedes.
A. aperturâ non prominente, capituli longitudinis 2/3 æquante: scutis corneis: longitudine totâ ad 2 uncias.
Orifice not protuberant, equalling two thirds of the length of the capitulum: scuta horny. Total length two inches.
Animal unknown.
Parasitic on Medusæ, Mediterranean and Atlantic Oceans: south shore of England(?)[45]
I have not seen this species, and have drawn up the above specific character from the Plates and brief descriptions in the Voyages of the Coquille and Astrolabe. M. Lesson thinks that his species differs from that of MM. Quoy and Gaimard; but as the peculiar yellow colour of the capitulum, general shape, short cirri, habits and range, are all common to both, I believe that they are identical. There is, however, one singular difference, namely, that the cirri are coloured bright blue in the Plate in the Voyage of the Astrolabe, and yellowish in that in the Voyage of the Coquille: this possibly may have resulted from the drawing in the latter case having been made from a specimen long kept in spirits.
M. Lesson says that there are seven pair of cirri, from which I infer that this species has a pair of long, articulated, caudal appendages: he asserts that each cirrus has ten segments; the cirri are short and little curled. M. Lesson remarks, that “deux languettes bifurques occupent le bas de l’ouverture ovale:” I can hardly doubt but that these are horny scuta of nearly the same shape as in A. minuta. The whole animal seems to be extremely transparent, and of a “jaune-citron clair.” MM. Quoy and Gaimard, however, remark, that different specimens vary from white to yellow. Entire length two inches, of which the capitulum is fourteen French lines. The peduncle is narrow and short.
[45] See Foot-note, p. 159.
3. Alepas cornuta. Pl. III, fig. 6.
A. aperturâ parvâ, leviter prominente: scutis nullis: capitulo plerumque tribus, parvis, compressis eminentiis secundum carinalem marginem instructo.
Orifice small, slightly protuberant; capitulum without horny scuta; generally with three small flattened projections along the carinal margin.
Outer maxillæ with the inner bristles divided into two groups; segments of the posterior cirri extremely numerous, each with one pair of main spines; inner rami of the fifth and sixth cirri rudimentary.
St. Vincent’s, West Indies, attached to an Antipathes, collected by the Rev. L. Guilding.
Capitulum globular, slightly flattened, smooth, translucent, entirely destitute of valves; orifice slightly projecting or tubular, parallel to the longitudinal axis of the peduncle, with the edges sinuous; it appears more tubular than it really is, from the convexity of the part of the capitulum immediately beneath the orifice. Three small, flexible, horny, irregular prominences project from the carinal margin; one at the bottom of the capitulum; a second about half-way up it; and a third generally close to the orifice; but their positions vary a little, and the prominences vary still more in shape and size, being either rounded and very small, or much flattened and considerably prominent; they are imperforate; in the membrane under them a few tubuli may be seen, which are not elsewhere visible; their summits are roughened with very minute points and beads of chitine; others, still minuter, are scattered over the whole capitulum.
Peduncle short, narrower than the capitulum, into which it insensibly blends; strongly wrinkled; surface of attachment wide; position with respect to the branches of the coralline, various.
Size and Colour.—The largest specimen, including the peduncle, was half an inch in length, and 3/10ths of an inch across the capitulum; colour, after having been long in spirits, brownish-yellow.
Filamentary Appendages, one on each side, short, tapering and pointed; seated on the posterior margin of a slight swelling beneath the basal articulation of the first cirrus; they are about equal in length to the pedicels of this cirrus.
The Mouth is directed abdominally; labrum much produced downwards, so as to be far separated from the adductor muscle; moderately bullate, forming about one third of the longitudinal axis of the entire mouth; upper part forming a slightly overhanging prominence; crest with a row of blunt, bead-like teeth, and externally to them there are numerous curved short bristles.
Palpi (Pl. X, fig. 8,) unusually narrow, a little hollowed out along their inner margins; pointing towards the adductor muscle; thickly covered with doubly serrated bristles.
Mandibles, with either two or three teeth; inferior angle narrow and tooth-like; both sides covered with strong bristles or spines, projecting beyond the toothed edge.
Maxillæ, with two large upper spines, and a third rather distant from them; beneath these, there is a wide notch or hollow; inferior part square, projecting, bearing six pair of moderately long spines, (of which the central one is the longest,) mingled with finer ones.
Outer Maxillæ, with a semicircular outline; the serrated bristles in front are divided into two groups; externally there is a rounded and very considerable projection covered with long bristles. Olfactory orifices slightly prominent, approximate, seated within and just beneath the rounded projections at the base of the maxillæ.
Body.—Prosoma little developed; thorax small.
Cirri, extremely long, but slightly curled, capable of being protruded so as almost to touch the base of the peduncle or the surface of attachment; segments short, extraordinarily numerous. In the three posterior cirri (excepting the rudimentary rami), each segment supports two long, slightly serrated spines, with two or three minute intermediate ones, and with one or two very short, thick spines on the inner and upper lateral margins: dorsal tufts with only two or three long, fine, unequal spines. All the segments are extremely flat, broad, short, with their anterior faces not protuberant; the greater number of the segments, especially the lower ones, have very obscure articulations, to be seen only with a high power, and these can be capable of little or no movement.
First Cirrus placed far from the second, with the top of its pedicel on a level with the top of the lower segment of the pedicel of the second cirrus; rami short, barely half the length of those of the second cirrus; unequal, the anterior ramus being only two thirds of the length of the posterior one; the shorter ramus contains thirteen inverted-conical segments, with one side rather protuberant; the longer ramus contains twenty-three thinner segments; the segments on both rami are clothed with bristles, arranged in two or three rows, forming narrow transverse brushes.
Second Cirrus, with its pedicel long, and its rami nearly equalling in length those of the sixth pair; the two rami of nearly equal length; the anterior one rather thicker than the posterior one; this posterior ramus has fifty-five segments! The bristles on the second and third cirri are arranged on the same principle as on the three posterior pair; but from an increase in size and number of the little intermediate bristles between the main pairs, and of those on the lateral rims, the segments, especially the basal ones, of the anterior ramus of the second cirrus, are clothed with thin brushes of bristles; these same bristles, on the posterior ramus of the second, and on both rami of the third cirrus, can hardly be said to form brushes, though longer and more numerous than those on the three posterior pair of cirri.
Fifth and Sixth Cirri.—These resemble each other, and have their inner or posterior rami in an almost rudimentary condition. In the sixth cirrus (Pl. X, fig. 28) the outer ramus (a) has actually sixty-three segments, whereas the rudimentary ramus (k) has only eleven, nearly cylindrical segments. These are furnished with extremely minute spines, of which those on the dorsal face are longer than those on the anterior face; the spines on the summit of the terminal segment are the longest; the segments are not half as thick as the normal ones in the outer ramus. The rudimentary ramus is only one seventh part longer than the pedicel which supports both it and the normal ramus. In the fifth cirrus, the rudimentary ramus is rather longer, and has thirteen segments, resembling those in the rudimentary ramus of the sixth. In the fourth cirrus there is no trace of this peculiar structure, the rami being equal in length and strength. The two rudimentary rami on each side are nearly straight, and seem incapable of movement; they project out behind the normal rami, and closely resemble in general appearance, the two caudal appendages; hence this cirripede, at first sight, appears to be six-tailed.
Pedicels of Cirri.—The pedicel of the first pair is very short; that of the second is the longest; those of the posterior cirri decreasing in length. Upper segments short; lower segments in the second, third and fourth cirri, irregularly and rather thickly clothed with bristles, but in the fifth and sixth cirri, there is a regular double row of main spines, with some minute intermediate ones: hence there is a difference, both in the rami and in the pedicels, between the fourth cirrus and the fifth and sixth, and this is a unique case. On the dorsal surface of the pedicel of the second cirrus, there is a tuft of much feathered fine spines.
Caudal Appendages.—Each consists of eight much tapering, very thin segments, furnished with a few short simple spines round their upper margins, and with a longer tuft on the terminal short segment; basal segments twice as thick as the middle ones. In length, these caudal appendages equal the pedicels of the sixth pair of cirri, and are a very little shorter than the rudimentary rami of these same cirri.
General Remarks.—Having examined this species first in the genus, I fully anticipated that the very remarkable character of the inner rami of the fifth and sixth cirri being rudimentary, and serving the same function (if any) with the caudal appendages, would have been generic; but this is not the case, for Alepas cornuta cannot be separated from A. minuta without violating a clear natural affinity.
4. Alepas Tubulosa.
Quoy et Gaimard. Voyage de l’Astrolabe, Pl. xciii, fig. 5, 1834.
A. aperturâ parvâ prominente et tubulosâ: scutis et prominentiis secundùm marginem carinalem, nullis.
Orifice small, tubular, protuberant; capitulum without horny scuta or projections along the carinal margin.
Animal unknown.
New Zealand, Tolaga Bay. Attached to a living Palinurus.
I have given the above brief character from the plate, and imperfect description in the voyage of the Astrolabe. The small and distinctly tubular orifice, and the smooth carinated edge of the globose capitulum, appear sufficiently to distinguish this species from A. cornuta. The colour is stated to have been white with violet tints. Length, two (French) lines.
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 œsophagus; 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-œsophageal 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 œsophagus 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 œsophagus; 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.