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Concluding Remarks.—The males from the absence of a mouth (and no doubt of a stomach), must necessarily be short-lived, and, I suppose, are periodically replaced by fresh males.[57] In one instance, the remnants of the two great compound eyes of the larva, could be seen at the end of the pouch, opposite the orifice. The larvae, I conclude, crawl in at the orifice, one side of which is formed, as we have seen, of yielding membrane, and scratch out the dead exuviae of the former occupant: certainly, the males are less firmly attached to their pouches, though some small quantity of cement is excreted, than are other Cirripedes to the objects to which they are attached. The small size of the female, and her valves not being thickly edged with chitine, accounts for the males having pouches specially formed for them, instead of being, as in S. vulgare, laterally imbedded in the chitine-border of the scuta. In hereafter weighing the evidence on the nature of the parasites in Ibla and in Scalpellum, the fact of the valves of the supposed female being here modified for the special purpose of lodging the males, will be seen to be important. If we imagine the male parasites to be extraneous animals, and that by adhering to the sack of the Scalpellum, they injure the corium and thus prevent the growth of the shell over an area exactly corresponding to their own size, and so form for themselves cavities; yet what can be said regarding the preparatory furrows? surely these narrow lines cannot have been produced by the pressure of the much broader parasites. Must we not see in the furrows, the first marking out, if such an expression may be used, of the habitation for the male, which has to be specially formed by the independent laws of growth of the female?
[57] It is possible, though opposed to all analogy, that the females may be short-lived, and breed only once, in which case the males would not have to be periodically replaced.
3. SCALPELLUM RUTILUM. Pl. VI, fig. 2.
S. (Foem. an Herm.) valvis 14 sub-rufis: carinae tecto plano, utrinque crista rotundata instructo; margine basali truncato: lateribus superioribus latitudine duplo longioribus.
(Fem. or Herm.) Capitulum with 14 reddish valves: carina with the roof flat, bordered on each side by a rounded ridge; basal margin truncated: upper latera twice as long as broad.
Mandibles with three teeth: maxillae narrow, bearing only four or five pair of spines: segments of the second and third pair of cirri with one side wholly covered with spines.
MALES, two, lodged in hollows, on the under sides of the scuta; pouch-formed, with four (?) rudimentary valves; no mouth; cirri not prehensile.
Hab. unknown; associated with Dichelaspis orthogonia. British Museum.
FEMALE OR HERMAPHRODITE.
There is only a single specimen in the British Museum, and this had nearly all its valves separated, and many of them in fragments: from its state of decay, I think the specimen must have been dead, when originally collected.
Description.—The capitulum consists of fourteen valves, including from analogy a rostrum.[58] Valves, apparently covered with membrane, bearing some thin spines on the margins; clouded with a fine, though pale, orange tint; surfaces plainly marked with lines of growth.
[58] In my first, and as I thought careful examination of the separated valves (my only materials) of this species, I mistook one of the triangular rostral latera for the rostrum, and hence was unfortunately led into an error in my 'Monograph on the Fossil Lepadidae of Great Britain,' in which I state that the present species has only twelve valves in the capitulum; and I inferred from this, that S. quadratum, S. fossula, &c., had only twelve valves; I still believe this to be correct, but the existence of fourteen valves in S. rutilum and S. ornatum, the recent species to which the above fossils are most closely allied, no doubt is a strong argument in favour of this higher number.
Scuta, elongated, nearly three times as long as broad; apex, pointed; basal margin extremely oblique, forming an acute angle with the occludent margin; the lateral margin is slightly hollowed out, and is separated from the tergal margin by a large rectangular projection or shoulder. The occludent margin is nearly straight; externally, there is a slight ridge running down the middle of the valve, from the apex to the baso-lateral angle; and a second ridge running from the apex to the tergo-lateral angle. The lines of growth do not end abruptly at the tergo-lateral angle, as is the case with S. ornatum and several fossil species, but run up a little way along the tergal margin. The umbo is seated at the uppermost point, and, therefore, the main growth is downwards. There is a large rounded depression for the adductor muscle (a, fig. 2 a'), and higher up, opposite the tergo-lateral angle, there is another hollow (b), for the lodgment of the males; this latter is of nearly the same shape as the hollow for the adductor muscle, but rather more conspicuous than it. From the appearance of the under surface of the scuta, it might readily have been thought, that there had been two adductor muscles.
Terga, of large size, longer than the scuta, flat, triangular, with the whole inferior part much produced and spear-like. A portion of the apex, must have projected freely above the sack.
Carina (Pl. VI, fig. 2 b'), simply bowed (i. e., not rectangularly bent), with the umbo (and primordial valve) seated at the upper point; rather massive, narrow, only slightly increasing in width from the upper to the lower end; the two sides are flat, and at right angles to the roof, which is bordered on each side by a rather broad, square-topped ridge (see section fig. 2 c'), or the roof may be said to have a square-edged furrow running from the apex to the basal margin, and widening downwards; these two ridges have their lines of growth oblique, and hence have a twisted appearance; the central depressed portion of the basal margin, which is square or truncated, descends lower down than the two ridges. The sides of the valve close to the apex are broad, and consist, as I believe, of intra-parietes, as well as of parietes, but these parts are not separated from each other by ridges, as is commonly the case, more especially with the fossil species. I have described the carina in some detail, on account of its resemblance to that of the cretaceous S. fossula, S. trilineatum, and S. quadricarinatum.
Rostrum, unknown; but one probably existed.
Upper Latera, of large size, elongated, quadrilateral, approaching to diamond-shaped, with the angles rounded, nearly twice as long as broad; almost flat; upper half acuminated, lying between the scuta and terga; the lower half broad, forming a rectangular projection lying between two latera of the lower whorl. The umbo is near the apex, the greater part of the growth being downwards, but the valve is added to a little, round the two sides of the apex; these additions do not take place in the early stages of growth, (as explained under S. vulgare,) and, therefore, they form a depressed rim.
Rostral Latera, almost exactly triangular, curved; basal margin furnished with a just perceptible rim.
Infra-median Latera, quadrilateral, sides unequal in length; the carino-basal margin being the longest; in area not quite twice double the rostral latera; directed obliquely upwards.
Carinal Latera, sub-triangular, produced upwards, with the apex rounded, and the two lateral margins hollowed out; the basal margin exceeds a little in length the basal margin of the rostral latera. The umbones of these two latera are seated at their basal outer angles, so that the growth of the valves is towards each other and upwards. The umbo of the infra-median latus is seated at the baso-rostral angle, and hence the growth is obliquely upwards. The umbones of the rostral latera must have been close together, over the unknown rostrum.
Length of capitulum about 4/10th of an inch.
Peduncle, only small fragments are preserved; the calcified scales are small, closely imbricated, several of them together only equalling in length the basal margin of the rostral latera. Each scale is thin, transversely elongated; basal imbedded portion straight; upper margin rounded.
Mouth.—Labrum with the upper part highly bullate, forming an overhanging projection; palpi apparently small and narrow.
Mandibles, narrow, produced, with three teeth; inferior angle pectinated, as is sometimes the third tooth; the distance between the tips of the first and second teeth equals that between the second tooth and the inferior angle.
Maxillae, extremely narrow, produced, without any notch; spinose edge exactly one third of the length of the mandibles: beneath the two upper great spines there are only three or four pair of spines; on the convex upper margin there are some minute tufts of the smallest hairs.
Outer Maxillae, rounded with the inner margins very sparingly but continuously covered with bristles. I could not ascertain whether the olfactory orifices were tubular.
Cirri.—These consisted, in the one specimen, of merely small fragments. The segments of the posterior cirri are elongated, not protuberant, and support, I believe, five pair of non-serrated spines, and an exterior row of very minute spines: dorsal spines fine and long. Either the second or third cirri, or probably both, are remarkable for having the whole of one side of each segment covered with irregular rows of long spines. Moreover, in the upper segments of these same cirri, between each separate dorsal tuft, there is placed one or two long bristles. The first cirrus appears to have had very broad segments, and these are singular from the spines in the dorsal rows, being extremely long. In some of the cirri, several of the basal segments are soldered together.
Caudal Appendages, lost.
From the state of the specimen, it was quite impossible to ascertain whether the individual here described was an hermaphrodite or female; from the analogy of its nearest congener, S. ornatum, the latter is the most probable; but the genus Ibla shows how the sexes may differ in the most closely-allied forms.
Affinities.—From the hollows on the under sides of the scuta, for the lodgment of the males; from the umbones of the scuta and of the carina being situated on the apices of these valves; and from all the characters of the mouth, S. rutilum is much more closely allied to S. ornatum than to any other species.
MALE, OR COMPLEMENTAL MALE.
In the concavity or hollow above the depression for the adductor muscle (Pl. VI, fig. 2 a'), I found males, but in so extremely decayed a condition, that they could hardly be examined. On one side, however, I distinctly saw the larval prehensile antennae, with pointed, hoof-like discs; and part of the thorax, with its small limbs and long spines, as in S. vulgare or S. ornatum. I also saw clearly the eye. The four calcified beads or rudimentary valves, I believe, were present; but in removing the specimen, the whole fell to pieces and was lost. The outer integument was covered with rather thick, very minute bristles, each about, 2/10,000th of an inch in length, and therefore only half the length of those on the complemental males of S. vulgare. The cavities for the males are not formed, as in S. ornatum, by the thickening of the internal surface of the valve round a defined space, but by the scutum being externally convex and internally concave down the middle, hollows being thus produced both for the lodgment of the males and for the attachment of the adductor muscle. These hollows are separated from each other by a slight transverse ridge. I do not know at which point of the margin of the valve, the orifice of the male is situated, but I presume close under the apex. In this species, as in S. ornatum, there can be no question that the scuta of the female are specially modified by their own growth for the reception of the males. It must be added that, as it was not possible to ascertain whether the ordinary form of S. rutilum was hermaphrodite or female, so it must remain doubtful whether the parasites are males or complemental males; but the former, I think, is most probable.
[TT SUB-CARINA PRESENTE.]
4. SCALPELLUM ROSTRATUM. Pl. VI, fig. 7.
S. (Herm.) valvis 15: rostro permagno: laterum paribus quatuor: pari superiore pentagono.
(Herm.) Capitulum with 15 valves: rostrum very large: four pair of latera; upper latera pentagonal.
Mandibles with four teeth; maxillae with the inferior angle prominent.
Complemental Male, attached between the mouth and adductor scutorum muscle; pedunculated; capitulum bearing a pair of elongated scuta and a rudimentary carina; mouth and cirri prehensile.
Philippine Archipelago; Island of Bantayan. Attached to a horny coralline: 20 fathoms. Mus. Cuming.
HERMAPHRODITE.
Capitulum, with the upper part narrow and produced.
Valves, 15 in number, placed close together, clouded pale red, covered with membrane, which is thickly clothed with minute points.
Scuta rather small, oval, with the upper end pointed; rather convex; basal and lateral margins blending into each other; the upper produced portion above the umbo is small; there is a deep pit for the adductor muscle, and there is a fold on the occludent margin in the usual position; occludent margin not straight.
Terga large, one third of their own length longer than the scuta; fat, sub-triangular; the three margins are not quite straight; the carinal margin projects a little above the apex of the carina, and the scutal margin is excised to fit the upper part of the scuta.
Carina bowed, internally deeply concave; upper portion above the umbo, about one fourth of the total length, extending between the terga for two thirds of their length, up to the slight prominences on their carinal margins: a ridge separates, on each side, the parietes from the tectum.
Rostrum (fig. 7 a) unusually large, about two thirds of the length of the scuta, and twice as long as the rostral pair of latera; internally concave, externally carinated; outline of the upper portion acutely triangular, of the lower portion rounded; umbo seated at the upper end.
Upper Latera pentagonal, with the apex rounded.
Rostral Latera flat, four-sided, with the basal margin the longest, and the baso-carinal angle produced.
Infra-median Latera nearly equalling in area the upper latera; not descending so low down as the rostral and carinal latera; outline of lower half semi-oval, of upper half rectangular.
Carinal Latera flat, four-sided, with the basal margin the longest, and slightly protuberant; baso-rostral angle produced; whole valve larger than the rostral latus, but closely resembling it in form.
Sub-carina minute, not above one third of the size of the rostral latera, which are the smallest of the other valves; internally deeply concave; externally solid, pyramidal, standing out beyond the surface of the carina, with the umbo at the apex.
The umbones of the four pair of latera are seated a little above the centre in each valve, on the summit of a raised triangular portion; this arises from the valve at first growing only downwards, and when added to at the upper end, the new part forms a ledge at a lower level round the old part, which had already acquired some thickness.
Peduncle, short, about half the length of the capitulum; narrow; thickly clothed with minute, longitudinally elongated, spindle-shaped, calcareous scales or beads, which project but little.
Length of the capitulum, rather under 3/10ths of an inch.
In a Young Specimen, with its capitulum, together with the peduncle, only 1/10th of an inch long, the scuta, terga, and carina are very large in proportion to the valves of the lower whorl. The latter project more, and are externally more pointed, as in the genus Pollicipes. The rostrum is well developed; the infra-median latera, in proportion, are the least of all the valves. The carina is straight and pointed, and not, relatively to the scuta, quite so long. The scuta are rather broader in proportion to their length, which would naturally follow from less having been added to their apices,—these valves at first growing only downwards. The membrane covering and connecting the valves is furnished with long thin spines.
Mouth.—Labrum placed far from the adductor scutorum muscle, with the upper part exceedingly prominent; apparently there are no teeth on the crest. Palpi blunt.
Mandibles, narrow, with four teeth, of which the second is not smaller than the others; inferior angle sharp and produced, barely pectinated.
Maxillae.—Under the two or three great upper spines, there is a tuft of fine bristles; the inferior part of the edge is step-like, and much upraised.
Outer Maxillae, with the inner edge deeply notched, and the bristles arranged in two quite distinct tufts; the bristles on the outer surface are long. Olfactory orifices, thin, tubular, and projecting.
Cirri.—The first pair is placed far from the second; the three posterior pair are long and straight, with their segments much elongated, not protuberant, bearing four or five pair of long spines, with little intermediate tufts of minute spines, and with the minutest spines on the lateral upper edges. Dorsal tufts with one spine extremely long, equalling a segment and a half in length; the others very short. Spines all serrated. First cirrus not very short; rami nearly equal, with the four terminal segments of both tapering; all the basal segments much thicker, and thickly covered with bristles. Second cirrus (as well as the third in a less degree), with the anterior ramus thicker than the posterior ramus, and with all the lower segments in both rami thickly clothed with three or four longitudinal rows of spines.
Caudal Appendages, spinose, uni-articulate; but the specimen was injured, and I could not exactly make out their shape: I believe it was oval, and thickly fringed with fine spines.
Penis, very small, almost rudimentary, narrow, and hairy, scarcely exceeding in length the pedicel of the sixth cirrus.
COMPLEMENTAL MALE. Pl. VI, fig. 5.
Before describing the parasite of the present species, which departs entirely from the character of the males of the three preceding species, it is proper to state that I consider it to be a Complemental Male simply from analogy, as will hereafter be more fully shown at the end of the genus. Had a specimen of the parasite been brought to me without any information, I should have concluded that it was an immature individual of a new genus of pedunculated Cirripedes, remarkable from the rudimentary condition of the valves, and exhibiting, in one important character, namely, in the form of the larval prehensile antennae, an alliance to Scalpellum. Had I been then told that three individuals in a group, had been found attached to S. rostratum, not outside the valves, but to the integument, in a central line, between the labrum and the adductor scutorum muscle, in such a position that when the Scalpellum closed its valves, these parasites were enclosed within the capitulum, my surprise would have been great; for it is very improbable that this singular and unparalleled position was accidental in this one group of specimens, inasmuch as there seems to be a relation between the naked condition of the capitulum of the parasite, and the protection afforded to it by the capitulum of the Scalpellum. It further becomes apparent on reflection, that these minute parasites, though having the appearance of immaturity, can not increase in size, or but little, for if they did grow, and acquired an ordinary size, they would either be killed by the pressure of the scuta of the Scalpellum, or they would destroy the latter, and in doing so soon lose their own support, and thus necessarily perish!
The one full-grown specimen of S. rostratum, in Mr. Cuming's collection, was in a good state of preservation, but dry. The three parasites were attached, as stated, close under the labrum, between it and the adductor muscle. They are constructed like ordinary Cirripedia, and have a mouth, thorax and cirri, enclosed in a capitulum, supported on a peduncle of moderate length and narrow. The entire length of the capitulum and peduncle, as far as could be ascertained in the shrivelled condition of the specimens, was 35/1000ths, and the greatest width of the capitulum 11/1000ths of an inch. Both capitulum and peduncle are hirsute with spines, nearly 1/1000th of an inch in length, mingled with shorter hairs in little rows of three and four together. The figure (5) in Pl. VI is merely a restoration, as accurate as could be made from the much shrivelled specimens. There are only three valves,—namely, an oval carina (a), seated rather high up on the capitulum, in a rudimentary condition and only 1/1000th of an inch in length, and a pair of scuta; these latter consist of a narrow, slightly curved plate, 8/1000ths in length, broadest at the lower end, where the breadth is 2/1000ths of an inch. The prehensile antennae, at the end of the peduncle, have pointed hoof-like discs: I was not able to make out the other parts. It deserves notice, that in the young specimen of the ordinary form of S. rostratum, 1/10th of an inch in length, and therefore only thrice as long as the parasites, all the valves were perfect, and seemed to have followed the ordinary law of development.
Mouth.—The largely bullate labrum is placed far from the adductor, in the same manner as in the hermaphrodite. The mandibles have three large sharp teeth, with the inferior point very sharp and small, so that there is one less tooth than in the hermaphrodite. The maxillae have two or three large upper spines, the others being very thin; I believe the lower part is upraised and step-like, as in the hermaphrodite. The outer maxillae are bilobed in front, with a few short bristles on the outer side near the bottom. I was not able, from the dried state of the specimens, to discover whether the olfactory orifices were tubular. Altogether it was apparent, from this imperfect examination, that there was a close similarity between the mouth of the parasite and of the hermaphrodite.
The Thorax is unusually elongated.
Cirri.—The first pair is very short, and is distant from the second. All have the appearance of immaturity, with their pedicels very long in proportion to their rami; the latter are slightly unequal in length, even in the sixth pair. There appeared to be six segments in the rami of the sixth pair, each segment bearing two or three pair of long spines.
Caudal Appendages, with two or three little spines on their summits.
Penis, short, blunt, thick at the apex, with one or two spines on it. I did not see any ovaria, but this could hardly have been expected in specimens in a dried condition, without they had happened to have been in a gorged condition. Certainly there were no ova.
In the general summary at the end of the genus, I shall give my reasons for believing this parasite to be the Complemental Male of the Scalpellum rostratum.
5. SCALPELLUM PERONII. Pl. VI, fig. 6.
SMILIUM PERONII. J. E. Gray. Annals of Philosoph., new series, tom. x, 1825.
—— —— . . . . . Spicilegia Zoologica, tab. iii, fig. 10, 1830.
ANATIFA OBLIQUA. Quoy et Gaimard. Voyage de l'Astrolabe, Pl. xciii, fig. 16, 1823-1834.
POLLICIPES OBLIQUA. Lamarck. An. sans Vertebres (2d edition).
S. (Herm.) valvis 13: laterum paribus tribus; pari superiore multum elongato: pedunculi squamis calcareis nullis.
(Herm.) Capitulum with 13 valves: three pair of latera; upper latera much elongated: peduncle without calcareous scales.
Mandibles with 10 or 11 unequal teeth: maxillae with the edge nearly straight, bearing numerous spines.
COMPLEMENTAL MALE, attached externally, between the scuta and below the adductor muscle; pedunculated; capitulum formed of six valves, with the carina descending far beneath the basal angle of the terga; mouth and cirri prehensile.
Swan River, Australia, attached to a coralline; Mus. Cuming. Port Western, Bass's Straits, as stated in the Voyage of the Astrolabe. Mus. Brit.
HERMAPHRODITE.
Capitulum formed of 13 valves; namely, two scuta, two terga, a carina and sub-carina, a rostrum, a pair of upper latera, and two pair of lower latera; these latter valves, with the sub-carina and the rostrum, make a whorl of six pieces. The upper part of the capitulum is, as usual, produced. The upper valves are separated (in specimens which have not been dried) by rather wide interspaces of membrane; they are covered (excepting, generally, their umbones,) by membrane, which in the interspaces is clothed with fine spines. The spines, or the marks where they were once articulated, are visible over nearly the entire surface of the membrane covering the valves. The spines are particularly numerous round the orifice of the sack. The whole capitulum, (in a dried condition), is coloured dull purplish-red, which is only in part due to the underlying corium, for the valves themselves are pale red. After having been long kept in spirits, the whole capitulum becomes colourless. The valves are smooth, faintly marked by lines of growth. The umbones of the lower valves project outwards, giving a denticulated appearance to the base of the capitulum.
Scuta, slightly convex, oblong, breadth about two thirds of the length, almost quadrilateral, with the upper portion produced into a flat projection; this projection is almost spear-shaped, being constricted a little on each side below the apex. There is a deep pit for the adductor muscle. The umbo is near the apex, the part above not being above one fifth of the whole length of the valve. As in S. vulgare, the growth is at first downwards, and subsequently a little upwards and downwards, thus producing the upper, small, spear-like projection, which lies at a lower level than the umbo. There is a fold on the occludent margin.
Terga, large, flat, triangular; carinal margin slightly hollowed out; occludent margin slightly arched, with a small portion protuberant to a variable amount. The apex is slightly curved towards the carina.
Carina, long, internally deeply concave, angularly bent, the lower portion slightly longer and wider than the upper part; the two halves meet each other at about an angle of 135 deg.; the upper half is parallel to the longer axis of the terga, between which it extends for three fourths of their length. The external surface is rounded, except near the umbo, where the edge is carinated; growth almost equally upwards and downwards; the parietes and tectum are not separated by ridges.
The Sub-carina lies close under the carina, and is placed almost transversely to the longer axis of the capitulum; external surface arched and smooth, the whole having the shape of half of a cone, with the apex a little curved outwards; seen internally, it may be said to be formed of two triangular wings placed at right angles to each other; basal margin straight; in size equalling the carinal latera.
Rostrum, lying almost transversely to the longer axis of the capitulum, under the basal margins of the scuta; in shape (fig. 6 a) closely resembling the sub-carina, but about one third larger than it; larger also than either the rostral or carinal latera; seen externally, appears like a half cone; seen internally, is formed of two triangular wings (with curved edges), placed at right-angles to each other.
Upper Latera, internally flat, oblong, twice as long as broad; upper end square, truncated; upper half rather wider than the lower half; fully twice as large as either of the lower latera. The basal points extend below the basal margins of the scuta. The umbo is placed a little above the centre.
Rostral Latera, minute, scarcely exceeding one third of the size of the carinal latera, and very much less than the rostrum; they are placed transversely under the basal point of the upper latus, or rather between it and the baso-lateral angle of the scutum; basal margin, as seen internally, straight; upper margin arched; rostral angle produced; internally flat; the whole valve is very thick and solid, so that the umbo which lies at the rostral end, projects rectangularly outwards.
Carinal Latera, oblong, nearly quadrilateral, with the upper angle produced; placed obliquely, parallel to the lower half of the upper latera; umbo slightly prominent, seated near the apex, with three rounded ridges proceeding from it; internal surface very slightly concave.
Peduncle and Attachment.—The peduncle is short, not equalling the capitulum in length. The whole surface is most thickly clothed with minute spines, which are not visible when the specimen is dry; I think it probable that they may sometimes all drop off before a new period of exuviation. The peduncle does not (at least in the specimens which I have examined, which were grouped in a bunch) taper at the lower end to a point; and after careful examination, I feel sure that the cement does not debouch from several successively formed orifices, as in S. vulgare and as in some Pollicipes, but only from the two original orifices in the prehensile antennae of the larva. In these latter organs, the sucking disc is hoof-like and pointed, and is narrower than the basal segment. The ultimate segment has on its inner side (supposing this segment stretched straight forwards,) a notch or step bearing at least three spines. The proportions of the different parts differ slightly from those in S. vulgare; but, as I shall hereafter have to give all the measurements, I do not think them worth repeating here. In the one large group of specimens examined by me, in Mr. Cuming's possession, all were attached symmetrically to the coralline, as in the case of S. vulgare, capitulum upwards, and their carinas outwards.
Length of capitulum about three quarters of an inch; width about half an inch; entire length, with peduncle, a little more than one inch.
The Mouth is placed far from the adductor muscle.
Labrum, with its basal margin much produced; upper part highly bullate, forming a rounded projection equalling the longitudinal axis of the rest of the mouth; crest without any teeth.
Palpi, triangular, with the two margins, thickly clothed with bristles; on each side of the mouth, near where the palpi are united to the mandibles, there is a slight, orbicular, shield-like swelling.
The Mandibles (Pl. X, fig. 3) have nine or ten very unequal teeth, with the inferior angle rather broad and pectinated; of these, there are four main teeth, of which the second is always the smallest, and between the four, one or two small teeth are interpolated; so that the total number is either nine or ten, and often varies on the two sides of the same individual, as likewise does the shape of the inferior angle.
Maxillae, with the edge nearly half as long as that of the mandibles, supporting from seventeen to twenty pairs of spines; the upper pair is only slightly larger than the others; a part near the inferior angle projects slightly beyond the rest of the nearly straight edge. The apodeme, at its base or point of origin, is unusually broad and flat.
Outer Maxillae, large and triangular. The inner margin is slightly concave, and continuously covered with short spines. The outer margin is bilobed, as in S. vulgare, with the basal part supporting a great tuft of long bristles, of which the greater number turn outwards, and almost cover the olfactory orifices. The latter are slightly prominent, placed some way apart from each other, with the above-mentioned tufts of bristles between them. All the spines of the trophi are in some degree doubly serrated.
Cirri.—The first pair is seated rather far from the second pair, and the prosoma being little developed, the shape of the body nearly resembles that of S. vulgare. The posterior cirri are elongated, very little curled, with the segments much flattened, not at all protuberant, bearing from five to seven pair of long serrated spines, with a few small spines in an exterior row; between each pair there is a very minute tuft of small bristles; the upper lateral rim of each segment is toothed with small spines; spines of the dorsal tufts, long, serrated. First pair, elongated, having numerous segments, namely, seventeen, whilst the sixth pair in the same individual had only twenty-one segments; rami nearly equal; segments short, nearly cylindrical, thickly clothed with long serrated spines. The second and third pair are nearly equal in length; they have their anterior rami slightly thicker than their posterior rami, both being much more thickly clothed with spines, than are the three posterior pair of cirri. Pedicels, rather short, with their inner edges not forming a projection, as in S. vulgare.
Caudal Appendages (Pl. X, fig. 20), uni-articulate, flat, rounded at their ends and moderately long; clothed most thickly, like brushes, with very fine bristles, which latter are serrated, and are longer than the appendages themselves.
Penis, of small size, narrow, pointed, and thickly clothed with delicate hairs; in length equalling only one fourth of the sixth cirrus.
Ovigerous Fraena, small, semicircular; entire edge thickly covered with glands. Ovarian tubes, within the peduncle, fully developed as usual.
Affinities.—This species differs from all the others in the absence of calcareous scales on the peduncle; but it has no other character which at all justifies its generic separation. In the shape of the scuta and carina it comes nearest to S. vulgare. Taking all the characters together, it is scarcely possible to say to which of the other species it is most closely allied, having close affinities with all. In the entire structure, however, of the Complemental Male, immediately to be described, this species certainly comes nearer to S. villosum than to any other species. I may add, that in S. villosum the latera are almost rudimentary, and therefore tend to disappear, whereas in S. Peronii it is the calcareous scales on the peduncle which have actually disappeared.
COMPLEMENTAL MALE. Pl. VI, fig. 3.
I examined, owing to the great kindness of Mr. Cunning, six dry specimens of the hermaphrodite S. Peronii, from Swan River, and one in spirits from another locality, in the British Museum. Out of these seven specimens, only three appeared to have had parasites attached to them, and these I infer, from reasons to be more fully given at the end of the genus, are Complemental Males. One of the three specimens, however, had two males close together. These parasites were firmly cemented to the integument of the hermaphrodite, in a fold, in a central line between the scuta, a little below (the animal being in the position in which it is figured) the adductor scutorum muscle, and therefore some way below the umbones of these valves. When the scuta are closed, the parasites, from their small size, are enclosed and protected. In every detail of structure, they are obviously pedunculated Cirripedia.
The Capitulum (Pl. VI, fig. 3) has six valves; namely, a pair of scuta and of terga, a carina, and a rostrum, all united by finely-villose membrane, furnished near the orifice with some much longer and thicker spines. The capitulum is truncated in a remarkable manner, the orifice not being, as in the hermaphrodite, in the same line with the peduncle, but almost transverse to it, and therefore almost parallel to the surface of attachment. The largest specimen measured transversely, through the scuta and terga, was 30/1000ths of an inch in breadth; another was only 26/1000ths to 27/1000ths: this latter specimen, measured longitudinally, from the base of the carina to the tips of the terga, was 15/1000ths of an inch. A scutum of the largest specimen was 17/1000ths in length. The scuta and terga are broadly oval, with the primordial valves very plain at their upper ends. I may here mention, that in a central line between the scuta, I observed the apparently single, minute, black eye, as in ordinary Cirripedia.
The Carina is straight, triangular, and internally slightly concave; its basal margin descends far below the basal points of the terga.
The Rostrum is shorter, and internally more concave than the carina: I believe it projects more abruptly outwards than is represented in the figure.
The Peduncle commences some little way below the scuta: it is narrow and very short: it is finely villose: it is lined by delicate transverse striae-less muscles, within which there are the usual stronger, longitudinal muscles. The base is flat and truncated. I examined, and carefully compared, the prehensile antennae with those of the hermaphrodite, and found every part and every measurement the same. The full importance of this identity will hereafter be more fully insisted on. The antennae are represented of their proper proportional size in fig. 3.
Mouth.—The labrum, as in the hermaphrodite, is highly bullate, and far removed from the adductor scutorum muscle. The Palpi are small and triangular, with their blunt apices clothed with a very few scattered bristles.
Mandibles, with only three teeth, and the lower angle minute, slightly pectinated; the first tooth is distant from the second, and larger than it. Width of the whole organ, .0021 of an inch.
Maxillae, bearing only a few spines, furnished with a long apodeme; beneath the upper large pair there is a notch, under which there are two spines of considerable size and a small tuft of fine bristles; width .001 of an inch, and therefore only 1/16th of the size of the same organ in the hermaphrodite: the relative sizes of the maxillae and mandibles are the same in the male and hermaphrodite.
Outer Maxillae blunt, triangular, with a few thinly-scattered bristles on the inner face; those on the outside being longer.
Cirri.—The First pair is far removed from the second; the rami are very short, barely exceeding the pedicel in length; they are formed of only four segments, each bearing a pair of spines; but on the end of the terminal segment, there are three spines, of which the central one is very long. Second pair also short. In the sixth pair there are five or six elongated segments, each bearing three pair of long spines; dorsal tufts large. The cirri are furnished with transversely-striated muscles.
The Caudal Appendages exist as two very minute plates, with a few bristles at their apices.
The Penis is not acuminated, with four bristles at the end; it is short, equalling only the lower segment of the pedicel of the sixth cirrus. In the one specimen preserved in spirits, I unfortunately omitted to search for the vesiculae seminales; I cannot doubt that such existed, but it would have been important to have ascertained whether they contained spermatozoa. I made out, most distinctly, that there was no trace of ovarian tubes within the peduncle; and my assertion may be believed when I state, that I traced the two much finer and more transparent cement-ducts, from the prehensile antennae up to the body of the animal: in Lepas I have repeatedly detected, with ease, the ovarian tubes within the peduncle, before the calcification of the valves had even commenced, and therefore at a much earlier period of growth than in these parasites. Consequently I am prepared to affirm, that these parasites are not females, but that, as far as can be judged, from external organs, they are exclusively males.
Concluding Remarks.—In comparing the capitulum of the hermaphrodite with that of the complemental male (Pl. VI, figs. 6 and 3), we must be struck with the differences in their shape, in the number, relative sizes, and forms of the several valves. It should, however, be borne in mind, that the scuta and carina in the hermaphrodite at first grow exclusively downwards; so that if we remove the upper portions subsequently added, the difference in shape in these valves is not so great as it at first appears. The rostrum in the male is of much larger relative size; whilst of the upper latera there is not a trace, although in the hermaphrodite these valves are larger than the rostrum. The terga, compared with those of the hermaphrodite, differ more essentially than do the other valves; and the manner in which the primordial valves project, shows that from the first commencement of calcification, the lines of growth have followed an unusual course. The great breadth and shortness of the terga is evidently related to the shortening of the whole capitulum, and the transverse position of the orifice; and this shortening of the capitulum, no doubt, is rendered necessary for its reception and protection within the shallow furrow between the scuta of the hermaphrodite. Finally, if we compare the internal parts of the hermaphrodite and male, the differences are considerable, though partly to be accounted for by the youth of the latter: the form and position of the labrum, and the distance between the first and second pair of cirri, is the same in both; but the mandibles and maxillae differ considerably.
To put the case as I have before done, if a specimen of one of these parasites had been brought to me to class without any information of its habits,—the downward direction of growth in all the valves, the presence of a rostrum, the villose outer integument, all the details of the prehensile antennae, the form of the animal's body, and the position of the labrum, would have convinced me that, though a quite new genus, it ought to have stood close to Scalpellum, and nearer to it than to Ibla.
6. SCALPELLUM VILLOSUM. Pl. VI, fig. 8.
POLLICIPES VILLOSUS on Plate (TOMENTOSUS in text). Leach. Encyclop. Brit., Suppl., vol. iii, 1824, Pl. lvii.
—— VILLOSUS.[59] G. B. Sowerby. Genera of Shells, Pollicipes, fig. 3, 1826.
CALANTICA HOMII. J. E. Gray. Annals of Phil., vol. x, p. 100, 1825.
[59] As Mr. Sowerby has adopted the name villosus, I have followed him; though as tomentosus is used through some mistake by Leach in the text, both names have equal claims as far as priority is concerned.
In Lamarck, 'Animaux Sans. Vert.,' the P. villosus of Sowerby is made synonymous with Anatifa villosa of Brugiere, which is certainly incorrect, although the A. villosa of this latter author is not positively known.
S. (Herm.) valvis 14: sub-rostro praesente: carina paene recta: laterum paribus tribus; pari superiore triangulo.
(Herm.) Capitulum with 14 valves: sub-rostrum present: carina nearly straight: three pair of latera; upper latera triangular.
Mandibles with four teeth, of which the second is the smallest: maxillae with a projection near the inferior angle: no caudal appendage.
COMPLEMENTAL MALE, attached externally between the scuta, below the adductor muscle; pedunculated; capitulum formed of six valves, with the carina not descending much below the basal angles of the terga: mouth and cirri prehensile.
Eastern Seas[60] (?) attached to shells and rocks. Mus. Brit.; College of Surgeons; Cuming.
[60] No habitat is attached to any of these specimens; but Mr. Sowerby informs me that he has seen specimens attached to the Modiola albicostata of Lamarck, which shell is said by the latter author to be found in the seas of India, Timor, and New Holland.
HERMAPHRODITE.
Capitulum with fourteen valves, consisting of a pair of scuta and of terga, a carina, (which five valves are much larger than the others,) a rostrum, sub-rostrum, sub-carina, and three pair of small latera. All the valves are covered by membrane, as are the calcareous scales on the peduncle; and this membrane everywhere is densely clothed with spines. The upper valves are not very thick; they stand rather close together. The eight valves of the lower whorl are more solid, and are placed far apart; they are small, tending to become rudimentary. None of the valves are added to at their upper ends, in which respect this species differs remarkably from the others of the genus, and approaches in character to Pollicipes.
Scuta, with a deep hollow for the adductor muscle, triangular, with the basal margin elongated, and protuberant.
Terga, large, flat, triangular, basal point blunt, with the carinal margin slightly hollowed out, and the scutal margin protuberant. Apex solid.
Carina, rather longer than the terga, straight, gradually widening from the upper to the basal end, deeply concave. In young specimens the upper part is slightly bowed inwards. Apex solid.
Sub-carina, with the inner surface crescent-shaped; the umbo points transversely outwards; in width it exceeds the largest of the latera.
Rostrum, triangular, internally (fig. 8 a) concave; basal margin slightly hollowed out, and deeply notched; rather less in width than the carina; short, with the umbo pointing upwards and outwards. In young specimens the apex curves a little inwards.
Sub-rostrum, with the inner surface transversely elongated (fig. 8 b), slightly crescent-shaped, about two thirds as wide as the rostrum. The apex points transversely outwards.
Latera, three pair; the middle pair apparently corresponds with the upper latera of the other species of the genus. The two other pair of latera, together with the rostrum and sub-carina, form a whorl. The sub-rostrum lies by itself, a little beneath this whorl. The latera are smaller than the rostrum or the sub-carina. They are placed far distant from each other; their inner surfaces are triangular; their umbones point upwards; the rostral pair is smaller than the other two pair, which are of equal size. The exact position of the rostral latus differed on the two sides of the specimen examined; apparently its normal position is at the baso-lateral angle of the scuta.
Peduncle, wide at the summit, longer than the capitulum; calcified scales small, not arranged very regularly; flattened, spindle-shaped, rather far separated from each other; imbedded in membrane, so that even their summits are rarely uncovered. The surface of the membrane is thickly clothed with spines, which are strong, thick, yellow, pointed, and furnished with large tubuli running to the underlying corium. These spines are arranged in groups of from three or four, to five or six. Besides these larger spines, the whole surface is villose with very minute colourless spines, not above 1/20th of the length of the larger ones. The surface of attachment is broad. This species, not being symmetrically attached to a coralline, the peduncle does not curve, as in most of the other species, towards the rostrum.
The capitulum is above half an inch in length.
Mouth.—The labrum is much produced downwards, but yet the mouth is not very far distant from the adductor muscle: the upper part is bullate, forming a small overhanging point, and in longitudinal diameter equals the rest of the mouth. Palpi blunt.
Mandibles with four teeth, strong, short, thick, the second tooth much smaller than the others; inferior angle broad, pectinated.
Maxillae with a long, rather sinuous edge, which, near the inferior angle, has a narrow projecting point, bearing rather finer spines; there is, also, apparently, a very minute tuft of small spines close under the two large upper spines: there are, altogether, about twenty pair of spines, without counting the smaller ones.
Outer Maxillae, with the inner edge slightly concave, continuously covered with bristles; exteriorly, with a prominence covered with longer bristles. Olfactory orifices prominent, protected by a slight punctured swelling between the bases of the first pair of cirri.
Cirri.—Prosoma moderately developed; first pair of cirri rather far removed from the second pair. The segments of the three posterior pair are not elongated, short, slightly protuberant in front, bearing four or five pairs of strong spines; a little below each pair, there is an intermediate tuft of very fine straight bristles, of which the upper tuft is the largest; on the lateral upper rims there are some short, strong spines; dorsal tufts rather small and thick; spines all more or less serrated, especially on the broad basal segments of the three anterior cirri. Pedicels of the cirri not particularly protuberant in front. First cirrus with rami, slightly unequal in length; not short; basal segments much thicker and more protuberant than the upper segments. Second cirrus; anterior ramus with six or seven basal segments highly protuberant, and crowded with spines; posterior ramus with about six segments, similarly characterised. Third cirrus with the anterior ramus having six, and the posterior ramus five segments, also similarly characterised.
Caudal Appendages absent, there being only a slight swelling on each side of the anus.
The oesophagus runs parallel to the labrum, and enters obliquely the summit of the stomach, which is destitute of caeca: the biliary envelope is longitudinally plicated.
There are no Filamentary Appendages.
Testes large, branched like a stag's horns, attached in a sheet to the ventral surface of the stomach: the vesiculae seminales enter the prosoma, and have their reflexed ends not very blunt. The Penis is rather narrow, with the terminal half plainly ringed, and bearing tufts of fine bristles arranged in circles, one tuft below the other; on the basal half there are only a few scattered minute bristles.
Affinities.—In the downward growth of all the valves, in the presence of a sub-rostrum, in the shape of the scuta, carina, and more especially of the triangular latera, in the form of the peduncle, with its irregularly-scattered calcified scales, in the shape of the animal's body, in the structure both of the mandibles and maxillae, in the arrangement of the spines, both on the anterior and posterior cirri, Scalpellum villosum most closely resembles, or rather is identical with, Pollicipes. Had it not been for the formation of the valves forming the capitulum, and from the presence of Complemental Males, I should have placed this species alongside of Pollicipes spinosus and sertus. In not having caudal appendages, S. villosum differs from all the species of Scalpellum and Pollicipes; but this organ is variable to an unusual degree in Pollicipes.
COMPLEMENTAL MALE. Pl. VI, fig. 4.
From the kindness of Professor Owen, Mr. Gray, and Mr. Cuming, I have been enabled to examine six specimens of this species; and on two of them I found Complemental males. They were attached in the same position as in S. Peronii; namely, beneath the adductor muscle, in the fold between the scuta, so as to be protected by the latter when closed. This parasite is six-valved, and has a close general resemblance with that of S. Peronii, but differs in very many points of detail. It is represented of the natural size at a fig. 4. The capitulum is 43/1000ths of an inch, measured across the scuta and terga; and the same measured from the base of the carina to the top of the capitulum; hence it is broader, by a quarter of the above measurement, and considerably higher than the male of S. Peronii. From the capitulum being higher, that is, not so much truncated, the orifice is placed more obliquely. The membrane connecting the valves is finely villose, and is besides furnished with spines, conspicuously thicker and longer than those on the male S. Peronii. The scuta and terga are much more elongated, a scutum being here 35/1000ths of an inch in length. The carina descends only just below the basal points of the terga, instead of far below them. The rostrum is a little broader and more arched than the carina; it is 2/1000ths in length, and therefore more than two thirds of the length of the carina, the latter being 28/1000ths of an inch from the apex to the basal margin. The primordial valves, with the usual hexagonal tissue, are seated on the tips of the scuta, terga, and carina, but not on the rostrum; so that these valves follow the same law of development, as in the ordinary and hermaphrodite form of Scalpellum. The scuta (a, fig. 4, greatly enlarged), the terga (b), and carina (c) of the male, resemble the same valves in the hermaphrodite, much more closely than do these valves in the male and hermaphrodite S. Peronii. The rostrum has not its basal margin hollowed out, and is very much larger relatively to the carina, than in the hermaphrodite. The large relative size of the rostrum in the complemental male both of this species and of S. Peronii, is a remarkable character, which I can in no way account for.
The peduncle is narrow and short, but in a different degree in the two specimens examined. It is naked. The prehensile antennae were not in a good state of preservation: the disc is narrower than the basal segment, and only slightly pointed, in which important respect it differs from the same part in the foregoing species; at its distal end, rather on the inner side, there are two or three spines, apparently in place of the excessively minute hairs, which are found at the same spot in some or in all the other species of Scalpellum, and in Ibla: similar strong spines occur in Pollicipes. Unfortunately, for the sake of comparison, I was not able to find the prehensile antennae in the hermaphrodite S. villosum.
Mouth.—Labrum bullate, with teeth on the crest. Palpi blunt, spinose.
Mandibles, with three teeth; inferior point rather strongly pectinated.
Maxillae, with a considerable notch under the upper pair of large spines; inferior part of the edge not prominent.
Outer Maxillae, with the spines on the inner edge arranged into two groups. Olfactory orifices tubular and prominent, with some long bristles near their bases. In the mandibles having only three teeth, in the maxillae being notched and in the lower part not being prominent, and, lastly, in the bristles on the inner face of the outer maxillae being arranged in two groups, these several organs differ from those in the hermaphrodite.
Cirri.—First pair short, with only three or four segments in each ramus: second cirrus, with the basal segments not very thickly clothed with spines: sixth cirrus with seven segments, not protuberant in front, each bearing four pairs of spines, without intermediate tufts.
Caudal appendages, none. This is an interesting fact, considering that these organs are likewise absent in the hermaphrodite S. villosum,—an absence highly remarkable, and confined to the genus Conchoderma and the one species of Anelasma.
Penis thick, not tapering, rather exceeding in length the pedicel of the sixth cirrus, square at the end, and furnished with some spines. In one specimen, I believe I distinguished the vesiculae seminales: if so, they contained only pulpy matter, and not spermatozoa. There were no ovarian tubes within the peduncle, which was lined by the usual muscles; I traced the two delicate cement-ducts, running from within the antennae close up to the animal's body. Hence in this case, as in that of S. Peronii, I dare positively affirm that ovarian tubes do not occur; for it is out of the question that I could have traced the cement-ducts, and, at the same time, overlooked the far larger and more conspicuous ovarian tubes, into which, moreover, the ducts, had they existed, would have run. Consequently, these parasites are not females; but judging from the probosciformed penis, and from the presence, as I believe, of vesiculae seminales, they are males.
The complemental males of the present species, and of S. Peronii, so closely resemble each other, that what I have stated regarding the affinities of the latter, are here quite applicable. It is singular how much more alike the parts of the mouth and the cirri of these two complemental males are, than the corresponding parts in the two hermaphrodites: this no doubt is due to the two males having been arrested in their development, at a corresponding early period of growth. Several of the characters, by which the hermaphrodite S. villosum so closely approaches, and almost blends into the genus Pollicipes,—such as the thicker cirri, with the intermediate tufts of bristles, the small second tooth of the mandibles, and the little brush-like prominence on the maxillae,—are not in the least apparent in the complemental male.
SUMMARY ON THE NATURE AND RELATIONS OF THE MALES AND COMPLEMENTAL MALES, IN IBLA AND SCALPELLUM.
Had the question been, whether the parasites which I have now described, were simply the males of the Cirripedes to which they are attached, the present summary and discussion would perhaps have been superfluous; but it is so novel a fact, that there should exist in the animal kingdom hermaphrodites, aided in their sexual functions by independent and, as I have called them, Complemental males, that a brief consideration of the evidence already advanced, and of some fresh points, will not be useless. These parasites are confined to the allied genera Ibla and Scalpellum; but they do not occur in Pollicipes,—a genus still more closely allied to Scalpellum; and it deserves notice, that their presence is only occasional in those species of Scalpellum which come nearest to Pollicipes. In the genera Ibla and Scalpellum, the facts present a singular parallelism; in both we have the simpler case of a female, with one or more males of an abnormal structure attached to her; and in both the far more extraordinary case of an hermaphrodite, with similarly attached Complemental males. In the two species of Ibla, the complemental and ordinary males resemble each other, as closely as do the corresponding hermaphrodite and female forms; so it is with two sets of the species of Scalpellum. But the males of Ibla and the males of Scalpellum certainly present no special relations to each other, as might have been expected, had they been distinct parasites independent of the animals to which they are attached, and considering that they are all Cirripedes having the same most unusual habits. On the contrary, it is certain that the animals which I consider to be the males and complemental males of the two species of Ibla, if classed by their own characters, would, from the reasons formerly assigned, form a new genus, nearer to Ibla than to the parasites of Scalpellum: so, again, the assumed males of the three latter species of Scalpellum would form two new genera, both of which would be more closely allied to Scalpellum, than to the parasites of Ibla. With respect to the parasites of the first three species of Scalpellum, they are in such an extraordinarily modified and embryonic condition, that they can hardly be compared with other Cirripedes; but certainly they do not approach the parasites of Ibla, more closely than the parasites of Scalpellum; and in the one important character of the antennae, they are identical both with the parasitic and ordinary forms of Scalpellum. That two sets of parasites having closely similar habits, and belonging to the same sub-class, should be more closely related in their whole organisation to the animals to which they are respectively attached, than to each other, would, if the parasites were really distinct and independent creatures, be a most singular phenomenon; but on the view that they differ only sexually from the Cirripedes on which they are parasitic, this relationship is obviously what might have been expected.
The two species of Ibla differ extremely little from each other, and so, as above remarked, do the two males. In Scalpellum the species differ more from each other, and so do the males. In this latter genus the species may be divided into two groups, the first containing S. vulgare, S. ornatum and S. rutilum, characterised by not having a sub-carina, by the rostrum being small, by the constant presence of four pair of latera, and by the peculiar shape of the carinal latera; the second group is characterised by having a sub-carina and a large rostrum, and may be subdivided into two little groups; viz., S. rostratum having four pairs of latera, and S. Peronii and villosum having only three pairs of latera: now the males, if classed by themselves, would inevitably be divided in exactly the same manner, namely, into two main groups,—the one including the closely similar, sack-formed males of S. vulgare, ornatum, and rutilum, the other the pedunculated males of S. rostratum, Peronii, and villosum; but this latter group would have to be subdivided into two little sub-groups, the one containing the three-valved male of S. rostratum, and the other the six-valved males of S. Peronii and S. villosum. It should not, however, be overlooked, that the two main groups of parasites differ from each other, far more than do the two corresponding groups of species to which they are attached; and, on the other hand, that the parasitic males of S. Peronii and S. villosum resemble each other more closely, than do the two hermaphrodite forms;—but it is very difficult to weigh the value of the differences in the different parts of species.
Besides these general, there are some closer relations between the parasites and the animals to which they are attached; thus the most conspicuous internal character by which Ibla quadrivalvis is distinguished from I. Cumingii, is the length of the caudal appendages and the greater size of the parts of the mouth; in the parasites, we have exactly corresponding differences. Out of the six species of Scalpellum in their ordinary state, S. ornatum is alone quite destitute of spines on the membrane connecting the valves; and had it not been for this circumstance, I should even have used the presence of spines as a generic character; on the other hand, S. villosum, in accordance with its specific name, has larger and more conspicuous spines than any other species. In the parasites we have an exactly parallel case; the parasite of S. ornatum being the only one without spines, and the spines on the parasite of S. villosum being much the largest! This latter species is highly singular in having no caudal appendages, and the parasite is destitute of these same organs, though present inn the parasites of S. rostratum and S. Peronii. Again, S. villosum approaches, in all its characters, very closely to the genus Pollicipes, and the parasite in having prehensile antennae, with the disc but little pointed, and with spines at the further end, departs from Scalpellum and approaches Pollicipes! Will any one believe that these several parallel differences, between the Cirripedial parasites and the Cirripedes to which they are attached, are accidental, and without signification? yet, this must be admitted, if my view of their male sex and mature be rejected.
One more, and the most important special relation between the parasites and the cirripedes to which they are attached, remains to be noticed, namely that of their prehensile larval antennae. I observed the antennae more or less perfectly in the males of all, and except in S. villosum, in all the species, though so utterly different in general appearance and structure, I found the peculiar, pointed, hoof-like discs, which are confined, I believe, to the genera Ibla and Scalpellum. In the hermaphrodite forms of Scalpellum, I was enabled to examine the antennae only in two species, S. vulgare and S. Peronii, (belonging, fortunately, to the two most distinct sections of the genus,) and after the most careful measurements of every part, I can affirm that, in S. vulgare, the antennae of the male and of the hermaphrodite are identical; but that they differ slightly in the proportional lengths of their segments, and in no other respect, from these same organs in S. Peronii,—in which again the antennae of the male and of the hermaphrodite are identical. The importance of this agreement will be more fully appreciated, if the reader will consider the following table, in which the generic and specific differences of the antennae in the Lepadidae, as far as known to me, are given. These organs are of high functional importance; they serve the larva for crawling, and being furnished with long, sometimes plumose spines, they serve apparently as organs of touch; and lastly, they are indispensable as a means of permanent attachment, being adapted to the different objects, to which the larva adheres. Hence the antennae might, a priori, have been deemed of high importance for classification. They are, moreover, embryonic in their nature; and embryonic parts, as is well known, possess the highest classificatory value. From these considerations, and looking to the actual facts as exhibited in the following table, the improbability that the parasites of S. vulgare and S. Peronii, so utterly different in external structure and habits one from the other, and from the Cirripedes to which they are attached, should yet have absolutely similar prehensile antennae with these Cirripedes, appears to me, on the supposition of the parasites being really independent creatures, and not, as I fully believe, merely in a different state of sexual development, insurmountably great.
The parasites of S. vulgare take advantage of a pre-existing fold on the edge of the scutum, where the chitine border is thicker; and in this respect there is nothing different from what would naturally happen with an independent parasite; but in S. ornatum the case is very different, for here the two scuta are specially modified, before the attachment of the parasites, in a manner which it is impossible to believe can be of any service to the species itself, irrespectively of the lodgment thus afforded for the males. So again in S. rutilum, the shape of the scutum seems adapted for the reception of the male, in a manner which must be attributed to its own growth, and not to the pressure or attachment of a foreign body. Now there is a strong and manifest improbability in an animal being specially modified to favour the parasitism of another, though there are innumerable instances in which parasites take advantage of pre-existing structures in the animals to which they are attached. On the other hand, there is no greater improbability in the female being modified for the attachment of the male, in a class in which all the individuals are attached to some object, than in the mutual organs of copulation being adapted to each other throughout the animal kingdom.
Generic Characters of the larval prehensile ANTENNAE, in the Lepadidae, as far as known from their imperfect state of preservation, and the number of species examined.
Name of Species. Length of, from end of disc to the further margin of the oblique basal articulation: Scale, fractions of the 1/6000ths of an inch. Length of, from end of disc to the inner margin of the basal articulation. Scale same. Width of basal segment, in widest part. Scale same. Disc, length of. Scale same. Disc, width of. Scale same. Ultimate segment, length of. Scale same. Ultimate segment, width of. Scale, fractions of the 1/20,000ths of an inch.
LEPAS: disc large, thin, almost circular, slightly elongated, with several long spines on the hinder margin; end segment with three very long, plumose spines on the upper exterior angle.[61]
L. anatifera (?) 62 20 23 22
L. australis, 111 40 42 39 18 30
L. pectinata, 51 23 16 14 9 16
L. fascicularis, 60 40 22 16 15
DICHELASPIS: disc small, thin, circular, with several spines on the hinder margin; end segment, with two long spines on the upper exterior angle.
D. Warwickii, 54 11 7-8 7-8 6 13-14
CONCHODERMA: disc large, thin, transversely elongated, with several long spines on the hinder margin; end segment, with two excessively long, plumose spines on the upper exterior corner.
C. virgata, 82 40 28 25 35 12 26
C. aurita. 28 40 11 26
ALEPAS: disc small, slightly elongated, with two or more spines on the hinder margin; end segment, with two long spines on the upper inner corner, and four shorter ones on the exterior corner.
A. cornuta, 60 24 14 12 8 20
IBLA (parasitic males of): disc, hoof-like, pointed, elongated, with a single spine on the hinder margin; end segment, with four short spines on the upper exterior corner.
I. Cumingii, 22 7-8 7 3-4 7-8
I. quadrivalvis, 32-33 10 8 5 4 8
SCALPELLUM: disc hoof-like, generally pointed and elongated, with a single spine on the hinder margin; end segment, with a notch on the inner[61] side, bearing two spines, longer than on the exterior corner.
S. vulgare, 39 19 10 10-11 5-6 6 7
S. ornatum, 36 21 10 12
S. Peronii, 30 19 9 6 5 10
POLLICIPES: disc small, hoof-like, not pointed, with a single spine on the hinder margin; end segment, as in Scalpellum.
P. cornucopia, 20 6 6 6 6 8
[61] In the diameter of the disc, the thin membranous border, which is present in the first three genera, is included; but I have some doubts, whether this border be not the first rim of cementing tissue, as all the specimens, of which measurements are here given, had been removed after attachment. In using the terms inner and outer sides of the end segment, it is supposed, that this segment is stretched straight forwards, instead of being bent rectangularly outwards, as in its natural position; and then there can be no doubt which is the inner and outer sides.
It should be observed that the evidence in this summary is of a cumulative nature. If we think it highly, or in some degree probable,—from the ordinary form of Ibla Cumingii having been shown on good evidence to be exclusively female,—from the absence of ova and ovaria in the assumed males of both species of Ibla, at the period when their vesiculae seminales were gorged with spermatozoa,—from the close general resemblance between the parts of the mouth in the parasites and in the Iblas to which they are attached,—from the differences between the two parasites being strictly analogous to the differences between the two species of Ibla,—from the generic character of their prehensile antennae,—and from other such points,—if from these several considerations, we admit that these parasites really are the males of the two species to which they adhere, then in some degree the occurrence of parasitic males in the allied genus Scalpellum is rendered more probable. So the absolute similarity in the antennae of the males and hermaphrodites both in S. vulgare and S. Peronii; and such relations as that of the relative villosity of the several species in this same genus, all in return strengthen the case in Ibla. Again, the six-valved parasites of S. Peronii and S. villosum are so closely similar, that their nature, whatever it may be, must be the same; hence we may add up the evidence derived from the identity of the antennae in the parasite and hermaphrodite S. Peronii, with that from the antennae in the male S. villosum, approaching in character to Pollicipes, to which genus the hermaphrodite is so closely allied; and to this evidence, again, may be added the singular coincident absence of caudal appendages in the male and hermaphrodite S. villosum. If these two six-valved parasites be received as the complemental males of their respective species, no one, probably, will doubt regarding the nature of the parasite of S. rostratum, in which the direct evidence is the weakest; but even in this case, the particular point of attachment, and the state of development of the valves, form a link connecting in some degree, the parasites of the first three species with the last two species of Scalpellum, in accordance with the affinities of the hermaphrodites.
When first examining the parasites of S. rostratum, S. Peronii, and S. villosum, before the weight of the cumulative evidence had struck me, and noting their apparent state of immaturity, it occurred to me that possibly they were the young of their respective species, in their normal state of development, attached to old individuals, as may often be seen in Lepas; this, however, would be a surprising fact, considering that S. rostratum and S. Peronii are ordinarily attached, in a certain definite position, to horny corallines, and considering that the exact points of attachment in these three parasites, (of which I have seen no other instance amongst common Cirripedes,) namely, between the scuta, would inevitably cause their early destruction, either directly or indirectly, by their living supports being destroyed. Nevertheless, I carefully examined a young specimen of S. rostratum only thrice as large as the parasite; and not having very young specimens of S. Peronii and villosum, I procured the young of closely-allied forms, namely, of S. vulgare, (with a capitulum only 4/100th of an inch in length,) and of Pollicipes polymerus, (with a capitulum of less size than that of one of the parasites,) and there was not the least sign of anything abnormal in the development of the valves. In S. vulgare, at a period when the calcified scuta could have been only 1/100th of an inch in length, (and therefore considerably less than the scuta in the parasites,) the upper latera must have been as much as 4/1000ths of an inch in length, and the valves of the lower whorl certainly distinguishable.
To sum up the evidence on the sex of the parasites, I was not able to discover a vestige of ova or ovaria in the two male Iblas; and I can venture to affirm positively, that the parasites of S. Peronii and S. villosum are not female. On the other hand, in the two male Iblas, I was enabled to demonstrate all the male organs, and I most distinctly saw spermatozoa. In the parasitic complemental male of S. vulgare, I also most plainly saw spermatozoa. In the parasites of S. rostratum, S. Peronii, and S. villosum, the external male organs were present. I may here just allude to the facts given in detail under Ibla, showing that it was hardly possible that I could be mistaken regarding the exclusively female sex of the ordinary form of I. Cumingii, seeing how immediately I perceived all the male organs in the hermaphrodite I. quadrivalvis; and as the parasite contained spermatozoa and no ova, the only possible way to escape from the conclusion that it was the male and I. Cumingii the female of the same species, was to invent two hypothetical creatures, of opposite sexes to the Ibla and its parasite, and which, though Cirripedes, would have to be locomotive! I insisted upon this alternative, because if the parasite of I. Cumingii be the male of that species, then unquestionably we have in I. quadrivalvis a male, complemental to an hermaphrodite,—a conclusion, as we have seen, hardly to be avoided in the genus Scalpellum, even if we trust exclusively to the facts therein exhibited.
With respect to the positions of the parasitic males, in relation to the impregnation of the ova in the females and hermaphrodites, it may be observed that in the two male Iblas, the elongated moveable body seems perfectly adapted for this end; in the males of the first three species of Scalpellum, the spermatozoa, owing to the manner in which the thorax is bent when protruded, would be easily discharged into the sack of the female or hermaphrodite; this would likewise probably happen with the complemental male of S. rostratum, considering its position within the orifice of the capitulum, between the mouth and the adductor scutorum muscle. The males of S. Peronii and villosum being fixed a little way beneath the orifice of the sack, below the adductor muscle, are less favorably situated, but the spermatozoa would probably be drawn into the sack by the ordinary action of the cirri of the hermaphrodite, and therefore would at least have as good a chance of fertilising some of the ova, as the pollen of many dioecious plants, trusted to the wind, has of reaching the stigmas of the female plants. Regarding the final cause, both of the simpler case of the separation of the sexes, notwithstanding that the two individuals, after the metamorphosis of the male, become indissolubly united together, and of the much more singular fact of the existence of Complemental males, I can throw no light; I will only repeat the observation made more than once, that in some of the hermaphrodites, the vesiculae seminales were small, and that in others the probosciformed penis was unusually short and thin.
Viewing the parasitic males, in relation to the structure and appearance of the species to which they belong, they present a singular series. In S. Peronii and S. villosum, the internal organs have the appearance of immaturity; the shape of the capitulum is specially modified for its reception between the scuta of the hermaphrodite, and several of the valves have not been developed. This atrophy of the valves, is carried much further in S. rostratum. In Ibla, many of the parts are embryonic in character, but others mature and perfect; some parts, as the capitulum, thorax, and cirri, are in a quite extraordinary state of atrophy; in fact, the parasitic males of Ibla consist almost exclusively of a mouth, mounted on the summit of the three anterior segments of the 21 normal segments of the archetype crustacean. In the males of the first three species of Scalpellum, some of the characters are embryonic,—as the absence of a mouth, the presence of the abdominal lobe, and the position of the few existing internal organs; other characters, such as the general external form, the four bead-like valves, the narrow orifice, the peculiar thorax and limbs, are special developments. These three latter parasites, certainly, are wonderfully unlike the hermaphrodites or females to which they belong; if classed as independent animals, they would assuredly be placed not in another family, but in another Order. When mature they may be said essentially to be mere bags of spermatozoa.
In looking for analogies to the facts here described, I have already referred to the minute male Lerneidae which cling to their females,—to the worm-like males of certain Cephalopoda, parasitic on the females,—and to certain Entozoons, in which the sexes cohere, or even are organically blended by one extremity of their bodies. The females in certain insects depart in structure, nearly or quite as widely from the Order to which they belong, as do these male parasitic Cirripedes; some of these females, like the males of the first three species of Scalpellum, do not feed, and some, I believe, have their mouths in a rudimentary condition; but in this latter respect, we have, amongst the Rotifera, a closely analogous case in the male of the Asplanchna of Gosse, which was discovered by Mr. Brightwell[62] to be entirely destitute of mouth and stomach, exactly as I find to be the case with the parasitic male of S. vulgare, and doubtless with its two close allies. For any analogy to the existence of males, complemental to hermaphrodites, we must look to the vegetable kingdom.
Finally, the simple fact of the diversity in the sexual relations, displayed within the limits of the general Ibla and Scalpellum, appears to me eminently curious; we have (1st) a female, with a male (or rarely two) permanently attached to her, protected by her, and nourished by any minute animals which may enter her sack; (2d) a female, with successive pairs of short-lived males, destitute of mouth and stomach, inhabiting two pouches formed on the under sides of her valves; (3d) an hermaphrodite, with from one or two, up to five or six similar short-lived males without mouth or stomach, attached to one particular spot on each side of the orifice of the capitulum; and (4th) hermaphrodites, with occasionally one, two, or three males, capable of seizing and devouring their prey in the ordinary Cirripedial method, attached to two different parts of the capitulum, in both cases being protected by the closing of the scuta. As I am summing up the singularity of the phenomena here presented, I will allude to the marvellous assemblage of beings seen by me within the sack of an Ibla quadrivalvis,—namely, an old and young male, both minute, worm-like, destitute of a capitulum, with a great mouth, and rudimentary thorax and limbs, attached to each other and to the hermaphrodite, which latter is utterly different in appearance and structure; secondly, the four or five, free, boat-shaped larvae, with their curious prehensile antennae, two great compound eyes, no mouth, and six natatory legs; and lastly, several hundreds of the larvae in their first stage of development, globular, with horn-shaped projections on their carapaces, minute single eyes, filiformed antennae, probosciformed mouths, and only three pair of natatory legs; what diverse beings, with scarcely anything in common, and yet all belonging to the same species!
[62] 'Annals of Natural History,' vol. ii, (2d series, 1848,) p. 153, Pl. vi. Mr. Dalrymple has published a very interesting paper on the same subject in the 'Philosophical Transactions,' (p. 342,) 1849; and there is another Memoir by Mr. Gosse in the 'Annals of Natural History,' vol. vi, (1850,) p. 18.
Genus—POLLICIPES. Pl. VII.
POLLICIES. Leach. Journal de Physique, tom. lxxxv, Julius, 1817.[63]
LEPAS. Linn. Systema Naturae, 1767.
ANATIFA. Brugiere. Encyclop. Method. (des Vers), 1789.
MITELLA. Oken. Lehrbuch der Naturgeschichte, 1815.
RAMPHIDIONA. Schumacher. Essai d'un Nouveau Syst. &c., 1817 (ante Julium).
POLYLEPAS. De Blainville. Dict. des Sc. Nat., 1824.
CAPITULUM (secundum Klein). J. E. Gray. Annals of Philos., tom. x, new series, Aug. 1825.
[63] This is one of the rare cases in which, after much deliberation, and with the advice of several distinguished naturalists, I have departed from the Rules of the British Association; for it will be seen that Mitella of Oken, and Ramphidiona of Schumacher, are both prior to Pollicipes of Leach; yet, as the latter name has been universally adopted throughout Europe and North America, and has been extensively used in geological works, it appears to me to be as useless as hopeless to attempt any change. It may be observed that the genus Pollicipes was originally proposed by Sir John Hill ('History of Animals,' vol. iii, p. 170), in 1752, but as this was before the discovery of the binomial system, by the Rules it is absolutely excluded as of any authority. In my opinion, under all these circumstances, it would be mere pedantry to go back to Oken's 'Lehrbuch der Naturgeschichte' for the name Mitella,—a work little known, and displaying entire ignorance regarding the Cirripedia.
Valvae ab 18 usque ad 100 et amplius: lateribus verticilli inferioris multis; lineis incrementi deorsum ordinatis: sub-rostrum semper adest: pedunculus squamiferus. |
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