I. (fœm.) valvarum marginibus lateralibus, et superficie interiore, cæruleis: pedunculi spinis plerumque annulis cæruleo-fuscis.
Fem. — Valves coloured, along the lateral margins and on the upper interior surface, blue: spines on the peduncle, generally ringed with blueish-brown.
Caudal appendages barely exceeding in length the pedicels of the sixth cirrus: rami of the first cirrus unequal in length by about two segments.
Male, — with scarcely a vestige of a capitulum: maxillæ with fewer spines than in the female.
Hab. — Philippine Archipelago, Island of Guimavas; invariably attached to the peduncle of Pollicipes mitella, in groups of two or three together; Mus. Cuming. Tavoy, British Burmah Empire; Mus. A. Gould of Boston.
FEMALE.
The capitulum is formed of four valves, but is hardly distinct from the peduncle. The latter includes, in its wide upper part, the animal’s body. The valves, namely, a pair of scuta and terga, are composed of an extremely hard, horny substance, or properly chitine, and do not contain any calcareous matter; they are extremely flat or thin, and both pairs project freely, like curved horns, to a considerable height above the sack enclosing the body: the terga project about twice as much as the scuta, and their flat apices generally diverge a little. The tips of the valves are frequently broken off; their surfaces are plainly marked or ribbed by the layers of growth, which are wide apart. The bases of the valves externally are hidden by the long spines of the peduncle.
Scuta. — These are shorter and broader than the terga; their internal (Pl. IV, fig. 8 b´) growing or corium-covered surfaces are slightly concave, triangular, with the basal margin longer than the other margins and slightly excised in the middle: there is no depression for the strong adductor muscle: the internal surface of the free horn-like portion, has a small central fold (formed by an oblique crest) running from the summit of the triangular growing surface to the tip of the valve: in perfect specimens, the growing and the free horn-like portions (the latter represented much too long in fig. 8 a´ and b´) are about equal in length: the basal portion of one side of the scutum overlaps the tergum.
Terga. — The internal glowing surface (fig. 8 b´) is almost diamond-shaped, and less in area than the sputa: external surface rounded; internal surface of the free horn-like portion, slightly concave.
Colour and Structure of Valves. — The external surfaces of the scuta and terga are yellow along the middle, plainly marked by zones of growth, and finely ribbed longitudinally: the internal surfaces and sides of the horns of the two valves, are coloured fine blue or purple; in the terga, however, the internal surface is mottled with yellow. In some specimens, especially in one from Tavoy, each zone of growth was only very narrowly edged with blue. When a thin layer is removed from one of the valves, the dark blue or rather purple appears by transmitted light a beautiful pale blue; and it is a very singular fact, that this blue portion is permanently turned by very gentle into a fiery red; the same singular effect is produced by muriatic and acetic acids. This blue part is much harder than the yellow; the latter exhibits, under a high power, a folded structure, and is penetrated by a few tubuli, whereas the harder blue portion has a cellular or scaled appearance. The spines of the peduncle exhibit, in a smaller degree, similar phenomena.
Peduncle. — This, as already remarked, cannot be distinctly separated from the capitulum; it is much compressed; it is composed of unusually thin and delicate membrane, transversely wrinkled and thickly clothed with long cylindrical horns or spines of chitine. These horns (fig. 8 c´) are not the analogues of the spines which are articulated on the external membranes of many Pedunculated and Sessile Cirripedes, but of the calcified scales on the peduncle of Scalpellum and Pollicipes; for they pass through the membrane (the underlying corium being marked by their bases) and are persistent, being added to, like the valves, during each successive period of growth. Their bases are concave, so that a section of the layers of growth exhibits a series of pointed cones, one within another. Each spine is nearly cylindrical, irregularly curled, and nodose or slightly enlarged at intervals: the apex smooth and pointed; the exterior surface longitudinally and finely ribbed, like the valves. The spines increase irregularly in size from the bottom to the top of the peduncle, those at the carinal and rostral ends being generally the longest; they point upwards and hide the bases of the valves. They are not arranged symmetrically, and new ones are formed over all parts of the peduncle. They are formed of the same substance as the valves, and do not contain any calcareous matter. These horns are yellowish, generally ringed with pale and dark blueish brown, which on pressure becomes slightly opalescent with pale blue and fiery red: sometimes only the upper horns are thus ringed, and in rare instances all are simply yellowish. The muscles of the peduncle run up to the bases of the four valves.
Surface of Attachment. — The cement appears to proceed from only two points. In some specimens, a considerable length of one side of the peduncle was fastened to the surface of attachment, the horns or spines being enveloped in the cement. The prehensile antennæ of the larva will presently be described under the male.
The length of an average specimen, including the peduncle and valves, is about half an inch, and the width across the widest part one fifth of an inch. Mr. Cuming has one specimen an inch in length, but this is owing to the peduncle being unusually tapering. In a specimen kept some years in spirits, the cirri, trophi, caudal appendages, and corium under the membrane between the scuta, were all dark purple; the sack and corium of peduncle clouded with purple, and the prosoma pale-coloured.
The Body (Pl. IV, fig. 8 a´) is small compared with the capitulum and peduncle; it is much flattened; the prosoma is of a very peculiar shape, being square, the sides of equal length, and, in an average-sized specimen, 75/1000th of an inch long. The peculiar shape arises from the great distance between the first and second cirrus — from the mouth being far removed from the adductor scutorum muscle — and lastly, from the lower part of the prosoma being not at all protuberant. The thorax which supports the cirri is also unusually small, plainly articulated, and separated from the prosoma by a deep fold. The thin membrane of the prosoma is studded with some fine, pointed hairs, about 3/400ths in length, and articulated on little circular discs.
Mouth, placed at a considerable distance from the adductor, and directed in an unusual manner towards the ventral surface of the thorax: the trophi are arranged, in a curved line, facing the thorax (see Pl. V, fig. 2, for this part in the male), and therefore less laterally than is usual.
Labrum (Pl. IV, fig. 8 a´ opposite c) highly bullate; the upper part produced into a blunt point: on its crest there are no teeth.
Palpi (fig. 8 a´ opposite d) small, blunt and rounded at their ends; inner margins slightly concave.
Mandibles (Pl. X, fig. 4), with three teeth, of which the first is much larger than the second and third, and distant from them: inferior angle produced and pectinated; upper edges of the second and third teeth finely pectinated.
Maxillæ (Pl. X, fig. 11) small, slightly but distinctly indented by two notches, supporting, besides the three upper great spines, three pairs of moderately long spines and some finer ones: apodeme short, thick.
Outer Maxillæ, unusually pointed, with the inner bristles not very numerous, continuously arranged; externally, the bristles are longer. Olfactory orifices, tubular, projecting, flattened, square on the summit, smooth: they point upwards and obliquely towards each other: they arise more laterally than in the other genera, namely outside the bases of the outer maxillæ, and between them and the inner maxillæ.
Between the bases of the first pair of cirri, there is a conical prominence, clothed with bristles and coloured purple: it projects nearly as high as the top of the lower segment of the pedicel of the first cirrus: it lies over the infra-œsophageal ganglion, and serves, I suspect, to fill up a little interval between the outer maxillæ.
Cirri long, little curved: the first pair (Pl. IV, fig. 8 a´) is situated at an extraordinary distance from the second; hence its basal articulation is on a level with the upper articulation of the pedicel of the second cirrus. In the three posterior cirri, the segments are laterally very flat, with their anterior surfaces not protuberant; each supports three pairs of thin, non-serrated bristles, of which the second pair is much shorter than the upper, and the lowest pair minute; between each pair there is a minute, rectangulary projecting bristle; dorsal tufts consist of two or three spines, of which one is longer than the others. The two bristles forming each pair, are not of equal length; for in the rami of each cirrus, the inner row of bristles is much shorter than the outer; and this seems to be connected with the flatness of the whole animal, and the consequent little power of divergence in the rami of the cirri. The first cirrus is rather short, with the rami unequal in length by about two segments: the anterior ramus is shorter and thicker than the other: segments numerous, each clothed with several rows of bristles. The second cirrus has the anterior ramus thicker and more thickly clothed with spines than the posterior ramus; this latter is rather more thickly clothed with spines than are the three posterior cirri; the third cirrus is in all these respects characterised like the second cirrus, but in a lesser degree. The pedicels of the second and third cirri are thickly and irregularly clothed with spines; in the three posterior pairs, the spines are placed in two regular rows, with some minute intermediate spines.
Caudal Appendages (Pl. IV, fig. 8 a´, f), multiarticulate, thin, tapering, in one specimen equalling, in another just exceeding, in length the pedicels of the sixth cirrus. In the latter specimen there were thirteen segments, of which the basal segments were broader and shorter than the upper; these latter are slightly constricted round the middle, so that they resemble, in a small degree, an hour-glass. Their upper margins are surrounded by rings of bristles; the terminal segment being surmounted by one or two very fine bristles much longer than the others. The two appendages are closely approximate; each arises from a narrow elongated slip, attached to the side of the pedicel of the sixth cirrus.
Nervous system. — I examined the upper part of the nervous chord, in order to ascertain whether the infra-œsophagean ganglion, which is of a globulo-oblong shape, was far separated from the second ganglion; and this I found to be the case, in accordance with the distance of the first cirrus from the second. I may here remark, that in S. quadrivalvis I discovered the eye, which, though in all probability really double, appeared to be single; it was situated near to the supra-œsophageal ganglion; and this ganglion was situated near to the adductor scutorum muscle, and at a considerable distance from the labrum. The aperture leading into the acoustic (?) sack, is situated much lower down than is usual (Pl. IV, fig. 8 a´), namely, at the length of the pedicel of the first cirrus beneath its basal articulation.
Generative system. — The specimens here described, of which I examined six, are exclusively female; they have no trace of the external, probosciformed penis, or of the two great vesiculæ seminales, or of the testes: on the other hand, the ovarian tubes within the peduncle are developed in the usual manner, and owing to the large size of the ova, are of large diameter, and hence very distinct: I detected, also, the true ovaria at the upper edge of the stomach.
MALE. Plate V, figs. 1-8.
Of the above-described Ibla Cumingii I dissected six specimens, four from the Philippine Archipelago, and two from the Burmah Empire, and none of them, as we have just seen, possessed the probosciformed penis, the vesiculæ seminales, or the testes, so conspicuous in other Cirripedes; on the other hand, all were furnished with the usual branching ovarian tubes and sometimes with ova, and consequently were unquestionably of the female sex. Within each of these specimens there was attached within the sack, in a nearly central line, at the rostral end, (Pl. IV, fig. 8 a´, h, magnified five times,) a flattened, purplish, worm-like little body, projecting about the 1/20th of an inch: in one of the six individuals, there was a second similar little creature attached at the carinal end of the sack. Before giving the reasons which I think conclusively prove that these little animals are the Males of the ordinary form of the Ibla Cumingii, it will be convenient to describe their structure in detail.
I am deeply indebted to the liberality and kindness of Mr. Cuming, in allowing me to cut up four specimens of this new species; and to Dr. Gould, of Boston, U. S., for the examination of the Burmese specimens.
The whole consists of a long, much flattened peduncle, separated from the mouth and thorax by an oblique fold, (Pl. V, fig. 1 h, b), which is conspicuous on the dorsal margin under the cirri, and can be traced with difficulty to the ventral margin. The thorax, itself rudimentary, and supporting rudimentary cirri, is in some individuals, as in the one represented (fig. 1, magnified 32 times), covered by, or received in the oblique fold h, just mentioned: in other individuals the thorax is drawn out, and then the fold shows merely as a notch on the dorsal margin, and the basal articulations of the cirri stand some little way above it. The basal edge of the large, well-developed month can be traced all round, and on the ventral margin (b), is generally marked by a slight notch. The dimensions and proportions vary much: the longest specimen, including the imbedded portion, was 8/100th, and the shortest barely 5/100ths of an inch in length; the width of the widest portion varied from 1 to 2/100ths of an inch: the specimen figured (Pl. IV, fig. 8 a´, and Pl. V, fig. 1,) is a broad, short individual. Generally, the middle of the peduncle is rather wider than the upper part.
Peduncle. — The main part of the animal, as may be seen in the drawing, consists of the peduncle, of which the imbedded portion tapers more or less suddenly in a very variable manner, and is of variable length, — in one specimen being one fourth of the entire length, and in another consisting of a mere minute blunt point. The free upper part of the animal is bent in various directions, in relation to the imbedded portion. The latter passes obliquely through the chitine membrane and corium, lining the sack of the female, and running along amidst the underlying muscles and inosculating fibrous tissue, is attached to them by cement at the extremity. The peduncle is often, but not in the individual represented, much constricted at the point where it passes through the skin of the female, and generally at several other points, especially towards the extremity (see fig. 1); the stages of its deeper and deeper imbedment being thus marked. The constrictions are, I believe, simply due to the continued growth of the male, whilst the hole through the membrane of the female does not yield. The imbedment, which is considerable only when the lower part of the peduncle is almost parallel to the coats of the sack, seems caused by the growth and repeated exuviations of the female; I believe, that the larva attaches itself to the chitine tunic of the sack, and that the cement, by some unknown means, affects the underlying corium, so that this particular portion of the tunic is not moulted with the adjoining integuments, and that the growth of the surrounding parts subsequently causes this portion to be buried deeper and deeper: it is, I believe, in the same way as the end of the peduncle in Conchoderma aurita, sometimes becomes imbedded in the skin of the whale to which it is attached.
The outer tunic of the peduncle is thin and structureless: in the fold (fig. 1 h) under the cirri, there is a central triangular gusset of still thinner membrane, corresponding in position to the membrane connecting the two terga in the female, and there subjected to much movement. I may here remark, that this fold, in its office of slightly protecting the thorax and in its position, evidently represents the capitulum with its valves, enclosing the whole body of the female. The outer tunic is lined by corium, mottled with purple, and within this there are two layers of striæ-less muscles, transverse and longitudinal, as in all pedunculated Cirripedes. The corium extends some way into the imbedded portion of the peduncle, and consequently, the outer tunic there continues to be added to layer under layer, and as it cannot be periodically moulted, it becomes much thicker than in the upper free part of the animal: the corium, however, does not extend to the extreme point, so that in it growth of all kind ceases.
Antennæ. — The peduncle terminates (Pl. V, fig. 1 e) in the two usual, larval, prehensile antennæ, which it is very difficult to see distinctly; they are tolerably well represented in fig. 5, greatly magnified. Their extreme length, measured from the basal articulation to the tip of the hoof-like disc, is 22/6000ths of an inch, the disc itself being 7/6000ths of an inch. The disc is slightly narrower than the long basal segment, from which it is divided by a broad conspicuous articulation; its lower surface is flat and its upper convex, altogether resembling in shape a mule’s hoof; its apex is fuzzy with the finest down; it bears a narrow ultimate segment, thrown, as usual, on one side; this segment supports on its rounded irregular summit, at least five, I believe, judging from the structure of the same part in the male larva of Ibla quadrivalvis, six or seven spines, longer than the segment itself: one long spine arises from the under side of the disc, near the base of the ultimate segment, and points backward: there is also a single curved spine on the outside, near the distal end of the basal segment. These organs were imbedded in a heart-shaped ball or cylinder of brown, transparent, finely laminated cement, and thus attached to the fibrous tissue of the female. The two cement-ducts (fig. 1 f) were very plain, each about 1/6000th of an inch in diameter, containing the usual inner chord of opaque cellular matter. I traced them at the one end into the prehensile antennæ as far as the disc; and at the other, up the peduncle for about one fourth of its length, where I lost them, and could not discover with certainty any cement glands. I may, however, here mention, that I found in the lower half of the peduncle, numerous, yellowish, transparent, excessively minute, pyramidal bodies, with step-formed sides; of these two or three often cohered by their bases like crystals; I have never seen anything like these in other Cirripedes, but it has occurred to me that they may possibly be connected with the formation of the cement: for in the last larval condition of Lepas, the cement-ducts run up to the gut-formed ovaria, filled at this period with yellowish, grape-like, cellular masses, without the intervention of cement glands, and I can imagine that similar masses, not being developed into functional ovaria, might give rise to the yellow pyramidal bodies.
Mouth. — The mouth is well developed; it is represented as seen vertically from above, in Pl. V, fig. 2, magnified about 60 times; the positions of the cirri and the outline of the thorax are accurately shown by dotted lines; a lateral view is given in fig. 1. In the specimen figured, the longitudinal diameter of the mouth, including the labrum, was 5/400th of an inch. The muscles of the several trophi have transverse striæ, and are the strongest and most conspicuous of any in the body. The labrum is largely bullate, with its summit slightly concave; the trophi are arranged in a remarkable manner, in a semicircular line, so as to be opposed to the labrum rather than to each other: there are no teeth or spines on the crest of the labrum, which overhangs the œsophageal cavity.
The Palpi (fig. 2 b and fig. 3) are very small, dark purple, bluntly pointed, with a few small bristles at the point; they do not extend beyond the knob at each corner of the labrum, which is here present, as in all other Lepadidæ; they are much smaller than in the female, though of a similar shape, and consequently, their points are much further apart: within their bases, the lateral muscles of the mandibles are, as usual, attached; they are represented in fig. 3, as seen from the inside, with the eye on a level with the concave summit of the labrum. The rudimentary condition of the palpi is connected, as remarked under the Anelasma squalicola, with the absence of efficient cirri.
The Mandibles (fig. 7) are well developed; they so closely resemble those of the female that it is superfluous to describe them: they are, however, smoother, without any trace of the teeth being pectinated, and with the inferior point smaller: measured in their longer direction, they are 7/2000th of an inch in length, and, therefore, a little less than one third of the size of those of the female. These organs have the usual muscles well developed, and the usual articulations.
The Maxillæ (fig. 8) have a rather rudimentary appearance; yet they have the same size relatively to the mandibles, as in the female, the spinose edge being 3/2000ths of an inch in length. These organs resemble, to a certain extent, those of the female, differing from them in being less prominent, — in the outline being more rounded, with the notches even less distinct, — and in the spines being fewer. The apodeme is short and broad.
The Outer Maxillæ (fig. 6) are pointed, with a small tuft of bristles at the apex; they are much less hairy than in the female, but have nearly the same unusual shape. Outside their bases, and between them and the inner maxillæ, the two well-developed, tubular, flattened, square-topped, olfactory orifices, project in exactly the same remarkable position as in the female; these are not represented in fig. 2, though sometimes they can be very distinctly seen, when the mouth is viewed from vertically above.
Thorax and Cirri. — The thorax is in a rudimentary condition: I did not observe the usual articulations. The whole, as seen from vertically above, is of small size, compared with the mouth; the outline is accurately shown by dotted lines in Tab. 5, fig. 2, together with the positions of the two pair of cirri, the caudal appendages, and anus. The posterior end of the thorax does not rise to the level of the summit of the mouth; and the thorax seems of no service, excepting perhaps as a sort of outer lip to protect the mouth. The cirri are in an extreme state of abortion, and evidently functionless; they are lined with purplish corium, without the vestige of a muscle; they are usually distorted and bent in different directions; they vary in size, and even those on opposite sides of the same individual, sometimes do not correspond, and do not arise from exactly corresponding points of the thorax. There are always two pair of cirri, which, as I conclude from the position of the excretory orifices, answer to the fifth and sixth pair in other Cirripedes. Each cirrus (fig. 4) usually carries only one ramus, placed on a large basal segment, evidently corresponding to the pedicel of a normal cirrus. The posterior are larger than the anterior cirri, which latter spring from points a little lower down on the thorax. In the posterior cirrus figured, the great basal articulation or pedicel, almost equals in length, and much exceeds in thickness, the four segments of the ramus; these segments are furnished on their upper dorsal edges with little brushes of spines, but have not even a trace of the normally larger and far more important anterior spines. In one specimen, the anterior cirrus had a large pedicel, carrying three segments, like those of the posterior pair; but in another specimen, one of the three segments showed traces of being divided into two, thus making four imperfect segments; whilst on the corresponding side of this same individual there were only two ill-formed segments, with their few spines differently arranged. Again, in a third specimen, the great basal segment of the anterior cirrus on one side, bore, exteriorly to the usual ramus, a single segment furnished with bristles, and evidently representing a second ramus; thus showing that the great basal segment certainly answers to a pedicel. I may here add, that on the integuments of these cirri, I observed with a high power, the serrated scale-like appearance common in other Cirripedes. Directly between the bases of the sixth cirrus, there is a very minute papillus, which, under the highest power, can be seen to consist of two closely approximate, flattened points; these, I have no doubt, are the caudal appendages in an extremely rudimentary condition, for I traced the vesiculæ seminales to this exact spot: close outside these rudimentary points, on a slight swelling, is the anus. It will presently be seen that in the male of the closely allied Ibla quadrivalvis, the nature of these caudal appendages admits of no doubt, for in this species they consist of more than one segment, are spinose, and close under them towards the mouth, there is a perfectly distinct papillus, representing the usual probosciformed penis.
Alimentary Canal. — The œsophagus is very narrow, and of remarkable length; from the orifice under the mandibles, it first runs back (in this respect not well represented in Pl. V, fig. 1,) under the bullate labrum, and then straight down the peduncle, where it terminates in the usual bell-shaped expansion, entering one side of the small globular stomach; the latter, at its lower end, is slightly constricted, and then is rather abruptly upturned. The rectum is of unparalleled length, and extremely narrow; it can be best detected after the dissolution by caustic potash of the softer parts, when its inner coat of chitine can be seen to be continuous, in the ordinary manner, with the outer integuments of the thorax. The anus, as already stated, is seated on a slight swelling, and consists of a small longitudinal slit (f, fig. 2), placed close outside the two very minute caudal appendages.
Organ of Sight. — In all the specimens, a little below the fold separating the mouth from the peduncle, and near the abdominal (or rostral) edge, a black ball (c, fig. 1), about 1/1000th of an inch in diameter, is conspicuous. When dissected out, it is somewhat conical in form, and appears to consist of an outer coat, with a layer of pigment-cells of a dark purple colour, surrounding a transparent, rather hard lens, apparently leaving a circular orifice at the summit, and forming a short tube at the base, surrounding what I believe to be a nerve. I was not able to perceive that this eye consisted of two eyes united, which the analogy of other Cirripedes makes me suppose probable, although in the ordinary and hermaphrodite Ibla quadrivalvis, the eye also appeared single. It is seated under the two transparent muscular layers, close upon the upper end of the stomach, and this is the exact position, as stated in the introductory discussion (), in which the eyes of pedunculated Cirripedes are commonly situated.
Generative System. — Within the muscular layer all round the upper part of the peduncle, and surrounding the stomach, there are numerous, little, rather irregular globular balls, with brown granular centres, so closely resembling the testes in other Cirripedes, though of smaller size, that I cannot doubt that this is their nature: they were much plainer, larger, and more numerous in some specimens than in others. The vesiculæ seminales can seldom be made distinctly out; but having cut one specimen transversely across the thorax, they were as plain as could be desired, lying parallel and close to each other above the rectum, (the animal being in the position as drawn,) and therefore in their normal situation. Each had a diameter four times as great as that of the rectum. In this individual the contents seemed (whether from decomposition or state of development, or from my not having used high enough power, I know not,) merely pulpy; but I have since found, in another specimen, masses of the most distinct spermatozoa, with the usual little knots on them, associated with numerous cells, about as large as and resembling those which I have examined in living Cirripedes, and from which I have every reason to believe the spermatozoa are developed. The vesiculæ seminales unite and terminate under the two extremely minute caudal appendages, and here I think I saw an orifice; but there is certainly no projecting, probosciformed penis.
Having dissected the six specimens with the utmost care, and having scrupulously examined the ovaria in other Cirripedes during their early stages of development, even before the exuviation of the larval locomotive organs, and in specimens of smaller size than the male Ibla, I am prepared to assert that there are no ovaria, and that these little creatures are exclusively males. It should be borne in mind, that in some of the specimens there were perfect spermatozoa in the vesiculæ seminales (as likewise in some of the males of I. quadrivalvis), and, therefore, if these individuals had been hermaphrodites, their ova would have been, at this period, well developed, and ready for impregnation: in this state it is almost impossible that they could have been overlooked. Moreover, it is probable that such ova would not have been very small, for the larvæ whence the parasitic males are derived, attain (as might have been inferred from the known dimensions of their prehensile antennæ, and as we shall show actually is the case in I. quadrivalvis,) the size common amongst ordinary Cirripedia.
Concluding Remarks. — That these animals are true Cirripedes, though having so different an external appearance from others of the class, admits of not the least doubt. The prehensile antennæ, enveloped in cement and including the two cement-ducts, would have been amply sufficient, without other parts — for instance, the mouth, by itself perfectly characteristic with each organ, together with the whole alimentary canal, constructed on the normal plan, — to have proved that they were Cirripedia. Under the head of the closely-allied Ibla quadrivalvis, we shall, moreover, see that the males are developed from larvæ, having every point of structure — the peculiar quasi-bivalve shell, the two compound eyes, the six natatory legs, &c., — characteristic of the Order. But in some respects, the males are in an embryonic condition, though unquestionably mature, as shown by the spermatozoa; — thus, in the thorax and mouth opening throughout their whole width into the cavity of the peduncle, that is, homologically into the anterior part of the head, and in the viscera being there lodged instead of in the thorax and prosoma, there is a manifest resemblance to the larva in its last stage of development: the absence of a probosciformed penis, the spineless peduncle, the food being obtained without the aid of cirri, and the length of the rectum, are likewise embryonic characters. Not only are these males, as just remarked, Cirripedia; but they manifestly belong to the Pedunculated Family. If a specimen had been brought to me to class, without relation to its sexual characters, I should have placed it, without any hesitation, next to the genus Ibla; if the mouth alone had been brought, I should assuredly have placed it actually in the genus Ibla: for let it be observed how nearly all the parts resemble those of Ibla Cumingii, excepting only in size and in being less hairy. The trophi are arranged in the same peculiar position as in the female; the labrum is largely bullate, without teeth on the crest; the palpi, though relatively smaller, are of the same shape; so are the mandibles; the maxillæ are more rounded and less prominent, but have the same exact size relatively to the mandibles; the outer maxillæ have the same, quite peculiar pointed outline, and the olfactory orifices are tubular, and hold the same unusual position. It is most rare to find so close a resemblance in the parts of the mouth, except in very closely allied genera, and often species of the same natural genus differ more. Again, in the long œsophagus and constricted stomach there is a resemblance to Ibla. In the male of Ibla quadrivalvis, the caudal appendages are multi-articulate; now, this is a character confined to four genera, namely, Ibla, Alepas, Pollicipes, and Lithotrya. I may add, that large tubular olfactory orifices are confined to the same genera, together with Scalpellum. Lastly, it particularly deserves notice, that the prehensile antennæ, in having a hoof-like and pointed disc, with a single spine on the heel, much more closely resemble these organs in Scalpellum, certainly the nearest ally of Ibla, than in any other genus; they differ from the antennæ in Scalpellum, only in the ultimate segment not having a notch on one side. These organs, unfortunately for the sake of comparison, were not found in the female and ordinary form of Ibla. The full importance of the above generic resemblance in the antennæ, will hereafter be more clearly seen, when their classificatory value is shown in the final discussion on the sexual relations of Ibla and Scalpellum.
Here, then, we have a pedunculated Cirripede very much nearer in all its essential characters to Ibla than to any other genus, and exclusively of the male sex; and this Cirripede in six specimens, from two distant localities, adhered to an Ibla exclusively of the female sex. May we not, then, safely conclude that these parasites are the males of the Ibla Cumingii? Considering that, in the same class with the Cirripedia, there is a whole family of crustaceans, the Lerneidæ, in which the males, compared with the females to which they cling, differ as much in appearance as in Ibla, and are even relatively smaller, I should not have added another remark, had there not been under the head of the following species, and of the next genus Scalpellum, a class of allied facts to be advanced, which in some respects support the view here taken, but in others are so remarkable and so hard to be believed, that I will call attention to the alternative, if the above view be rejected. The ordinary Ibla Cumingii must have a male, for that it is not an hermaphrodite can hardly be questioned, seeing how easy it always is to detect the male organs of generation; and we must consequently believe in the visits of a locomotive male, though the existence of a locomotive Cirripede is improbable in the highest degree. Again, as the little animal, considered by me to be the male of I. Cumingii, is exclusively a male, (for there were no traces of ova or ovaria, though the spermatozoa were perfect,) we must believe in a locomotive Cirripede of the opposite sex, though the existence in any class of a female visiting a fixed male is unknown: in short, we should have hypothetically to make two locomotive Cirripedes, which, in all probability, would differ as much from their fixed opposite sexes, as does the Cirripede, considered by me to be the male of I. Cumingii, from the ordinary form. This being the case, I conclude that the evidence is amply sufficient to prove that the little parasitic Cirripede here described, is the male of Ibla Cumingii.
It deserves notice, that in the class Crustacea, both in the Lerneidæ and in the Cirripedia, the males more closely resemble the larvæ, than do the females; whereas amongst insects, as in the case of the glow-worm in Coleoptera, and of certain nocturnal Lepidoptera, it is the female which retains an embryonic character, being worm-like or caterpillar-like, without wings. But in all these cases, the male is more locomotive than the female.
If we look for analogies to the facts here given, we shall find them in the Lerneidæ already alluded to, but in these the males are not permanently attached to the females, only cling, I believe, to them voluntarily. The extraordinary case of the Hectocotyle, originally described as a worm parasitic on certain Cephalopoda, but now shown by Kölliker to be the male of the species to which it is attached, is perhaps more strictly parallel. So again in the entozoic worm, the Heteroura androphora the sexes cohere, but are essentially distinct: “this singular species, however,” according to Professor Owen, “offers the transitional grade to that still more extraordinary Entozoon, the Syngamus trachealis, in which the male is organically blended by its caudal extremity with the female, immediately anterior to the slit-shaped aperture of the vulva. By this union a kind of hermaphroditism is produced; but the male apparatus is furnished with its own peculiar nutrient system; and an individual animal is constituted distinct in every respect, save in its terminal confluence with the body of the female. This condition of animal life, which was conceived by Hunter as within the circle of physiological possibilities, has hitherto been exemplified only in the single species of Entozoon, the discovery of the true nature of which, is due to the sagacity and patient research of Dr. C. Th. Von Siebold.” In Ibla, the males and females are not organically united, but only permanently and immovably attached to each other. We have in this genus the additional singularity of occasionally two males parasitic on one female.
Cyclopædia of Anatomy and Physiology, .
I have used the term parasitic, which perhaps ought strictly to be confined to cases where one creature derives its nutriment from another, inasmuch as the male is invariably and permanently attached to and imbedded in the female, — from its being protected by her capitulum, so that its own capitulum is not developed — and from its feeding on minute animals infesting her sack. The male Ibla must seize its prey, guided probably by its well-developed olfactory organs, through the movement of its long, flexible body, furnished with muscles, and with the mouth seated on the summit. We have already seen one instance of a Cirripede, the Anelasma, obtaining its food without the aid of cirri, by means of its probosciformed, flexible mouth. The eye can serve only to announce to the male when the female opens her valves, allowing occasionally some minute prey to enter. In ordinary Cirripedes the penis is long, articulated, and capable of varied movements, I presume for the purpose of impregnating each separate ovum: the male Ibla has no such organ; and no doubt the whole body, furnished like the penis with longitudinal and transverse muscles, serves the same purpose! I may remark, that it seems surprising that so small a male should secrete sufficient semen to impregnate the ova of the female, but the ova are not nearly so numerous in Ibla as in most genera of Cirripedes; and the smallness of the males in some parasitic Crustacea has already been alluded to. The male must always be younger than the female, for the latter must first grow large enough for the larva of the male to crawl into her sack. Whether the male lives as long as the female I know not, but he certainly lives for a considerable period and increases in size, as shown by the depth to which the end of the peduncle is imbedded. Moreover we shall see, under the next species, that the male is metamorphosed from a larva, not one sixth of its own size.
In the male Ibla, abortion has been carried to an extraordinary and, I should think, almost unparalleled extent. Of the twenty-one segments believed to be normally present in every Crustacean, or of the seventeen known to be present in Cirripedes, the three anterior segments are here well developed, forming the peduncle: the mouth consists as usual of three small segments: the succeeding eight segments are represented by the rudimentary and functionless thorax, supporting only two pair of distorted, rudimentary and functionless cirri: the seven segments of the abdomen have disappeared, with the exception of the excessively minute caudal appendages; so that, of the twenty-one normal segments, fifteen are more or less aborted. The state of the cirri is curious, and may be compared to that of the anthers in a semi-double flower; for they are not simply rudimentary in size and function, but they are monstrous, and generally do not even correspond on opposite sides of the same individual. As males in other classes of the animal kingdom often retain some female characters, so here (though the case is not strictly analogous) the male possesses the cementing apparatus, which homologically is part of an ovarian tube modified.
Certain plants offer a closer, though not perfect, analogy. Thus, in the florets of some compositous flowers, the pistil, besides its proper female functional end, serves to brush the pollen off the anthers; while, in the florets of some other compositæ (see the account of Silphium in ‘Ch. K. Sprengel Das entdeckte Geheimniss der Natur’), the pistil is functionless for its proper end, the flower being exclusively male, but its style is developed, and still serves as a brush. So in the male Ibla, part of the ovaria, in a modified condition, is still present, and serves as a cementing apparatus.
The individuals in every other genus (with the exception of Scalpellum), in the several families, in the three Orders of Cirripedia, are hermaphrodite or bisexual. Why, then, is Ibla unisexual; yet, becoming, in the most paradoxical manner, from its earliest youth, essentially bisexual? Would food have been deficient, and was the seizure of infusoria by another and differently constructed individual, necessary for the support of the male and female organs? The orifice of the sack of the female is unusually narrow; would the presence of testes and vesiculæ seminales have rendered her thorax and prosoma inconveniently thick? Seeing the analogous facts in the six, differently-constructed species of the allied genus Scalpellum, I infer there must be some profounder and more mysterious final cause.