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Prehensile Antennae.—I examined these in the larvae presently to be mentioned, and therefore they were in better condition than in the mature animal when cemented. Their total length, measured along the outside, from the basal articulation to the end of the disc, is 32/6000ths or 33/6000ths of an inch—that is, one third longer than in I. Cumingii; whilst the hoof-like disc itself is 8/6000ths, or only 1/6000th of an inch longer than this same part in I. Cumingii: the apex of the disc is downy, or bears some excessively minute spines. The ultimate segment has its end irregularly rounded, with the spines obscurely divided into two groups, the outer group consisting of two or three longer and thinner spines, and the inner group of, as I believe, five rather shorter spines: the longer spines equal in length the whole ultimate segment. I could not perceive that they were plumose, as in many other genera. A single, rather thicker and long spine, pointing backwards, is attached to the under side of the disc, nearly opposite to the point where the ultimate segment is articulated on the upper convex surface. Another single, curved spine is attached on the outer side of the basal segment, near its distal end.
Development of the Male.—In the specimen before alluded to, which included two males, one of these was only the 30/1000ths of an inch in length, and therefore between one fifth and one sixth of the size of the mature male. It had, probably, undergone only one exuviation since its metamorphosis, for the larva is nearly as long, namely, 25/1000ths of an inch. In this young male, the mouth formed one third of the entire length: it was attached, not as in every other case to the sack of the hermaphrodite, but low down to the peduncle of the other male.
In the sack with these two males, there were certainly four, I believe five, larvae, which in every main point of structure resembled the larvae of other pedunculated Cirripedes. From the peculiar form of their prehensile antennae, differing in no respect, except in the proportional lengths of the segments, from the same organ in the male I. Cumingii, I can feel no doubt that these were the larvae of the male I. quadrivalvis;—for a moment's reflection will show how excessively improbable it is, that several larvae of some other Cirripede, and that a Cirripede intimately allied to the parasitic male Ibla, should have forced themselves, without any apparent object, into the sack of the hermaphrodite Ibla. The larvae, though not yet attached, were on the point of attachment, so that the single eye of the mature animal could be distinctly seen, lying near to the two great compound eyes of the larva. We have also just seen, that one male quite recently here had undergone its metamorphosis. The larvae are 25/1000ths of an inch in length, and rather more than 10/1000ths in width in the widest part: they are boat-shaped, the dorsal edge forming the keel of the boat; the anterior end is only a little blunter than the posterior end; the quasi-bivalve carapace is smooth. All the essential points of structure in the larvae of other Cirripedes at this stage, could be distinctly here seen,—such as the two compound eyes, with the apodemes to which they are attached, and the two oblong sternal plates whence the apodemes spring,—the adductor muscle,—the six natatory legs, with long plumose spines,—the abdomen, with its three small segments and the caudal appendages,—the prehensile antennae already described,—and, lastly, the two little (auditory?) sacks at the antero-sternal edges of the carapace, but not so near the anterior extremity as in Lepas. The four or five larvae, after having undergone in the open sea the several preparatory metamorphoses common to the class, must have voluntarily entered the sack of the hermaphrodite: ultimately would they, on finding two males already attached there, have retired, and sought another individual less well provided; or would they all have remained, and so formed a polyandrous establishment, such as we shall presently see occurs sometimes in Scalpellum? This must remain quite uncertain.
In this same hermaphrodite specimen of I. quadrivalvis, the two ovigerous lamellae contained some hundreds of larvae in the first stage of development, which were liberated from their enveloping membranes by a touch of a needle: they were about the 16/1000ths of an inch in length, and presented all the usual characters of larvae at this period. What a truly wonderful assemblage of beings of the same species, but how marvellously unlike in appearance, did this individual hermaphrodite present! We have the numerous, almost globular larvae, with lateral horns to their carapaces, with their three pair of legs, single eye, probosciformed mouth and long tail:—we have the somewhat larger larvae in the last stage of development, much compressed, boat-formed, with their two great compound eyes, curious prehensile antennae, closed rudimentary mouth and six natatory legs so different from those in the first stage:—we have the two attached males, with their bodies reduced almost to a mouth placed on the summit of a peduncle, with a minute, apparently single eye shining through the integuments, without any carapace or capitulum, and with the thorax as well as the legs or cirri rudimentary and functionless:—lastly, we have the hermaphrodite, with all its complicated organisation, its thorax supporting six pairs of multi-articulated two-armed cirri, and its well-developed capitulum furnished with horny valves, surrounding this wonderful assemblage of beings. Unquestionably, without a rigid examination, these four forms would have been ranked in different families, if not orders, of the articulated kingdom.
Concluding Remarks.—If the creature which I have considered as the male of Ibla Cumingii be really so, and the evidence formerly given seems to me amply conclusive, then the animal just described, from its close affinity in every point of structure with the former, assuredly is the male of Ibla quadrivalvis. But feeling strongly how improbable it is, that an additional or complemental male should be associated with an hermaphrodite, I will make a few remarks on the only possible hypothesis, if my view be rejected,—namely, that the two parasites considered by me to be exclusively males, are not so, but are independent hermaphrodite Cirripedes, the female organs and ova (which, if present, would have been nearly mature, judging from the presence of spermatozoa in both species) having been overlooked by me in every specimen: and again, that in the animal described as the female I. Cumingii, I have, though minutely dissecting several specimens, and finding far smaller parts, such as the organs of sense and nervous system, entirely overlooked all the conspicuous male organs, though when I came to I. quadrivalvis, and naturally expected to find it likewise exclusively female, a single glance showed me the great probosciformed penis, and by the simplest dissection the vesiculae seminales and testes were exhibited. Such an oversight is scarcely credible; but even if assumed, we have to believe in the extraordinary circumstance of the two parasites being species of an independent genus, not only the very next in alliance to the animals to which they are attached, but in certain most important points, namely, the organs of the mouth, actually deserving a place in the very same genus. Moreover, the two parasites differ from each other, not only in about the same slight degree, but in a corresponding manner, as do the two Iblas to which they are attached; thus the mouths of Ibla quadrivalvis and I. Cumingii are closely similar, (the difference being barely of specific value,) so are the mouths of the two parasites; but the parts are larger in the hermaphrodite I. quadrivalvis, than in I. Cumingii, so are they in the parasites. Again, the most conspicuous character in I. quadrivalvis, is the number of segments in the caudal appendages, far exceeding those in the other species of Ibla, as well as of every other pedunculated Cirripede, and the parasite of this species has articulated spinose appendages, far larger than the barely visible, non-articulated pair in I. Cumingii.
Considering the whole case, there seems no room to doubt the justness of the conclusion arrived at, under the former as well as under the present species, namely, that these little parasites are the males of the two species of Ibla to which they are attached;—wonderful though the fact be, that in one case, the male should pair with an hermaphrodite already provided with efficient male organs. It is to bring this fact prominently forward, that I have called such males, Complemental Males; as they seem to form the complement to the male organs in the hermaphrodite. We look in vain for any, as yet known, analogous facts in the animal kingdom. In the genus Scalpellum, however, next in alliance to Ibla, in which, consequently, if anywhere, we might expect to find such facts, they occur; and until these are fully considered, I hope the conclusions here arrived at, will not be summarily rejected. Although the existence of Hermaphrodites and Males within the limits of the same species, is a new fact amongst animals, it is far from rare in the Vegetable Kingdom: the male flowers, moreover, are sometimes in a rudimentary condition compared to the hermaphrodite flowers, exactly in the same manner as are the male Iblas. If the final cause of the existence of these Complemental Males be asked, no certain answer can be given; the vesiculae seminales in the hermaphrodite of Ibla quadrivalvis, appeared to be of small diameter; but on the other hand, the ova to be impregnated are fewer than in most Cirripedes. No explanation, as we have seen, can be given of the much simpler case of the mere separation of the sexes in Ibla Cumingii: nor can any explanation, I believe, be given of the much more varied arrangement of the parts of fructification in plants of the Linnean class, Polygamia.
Genus—SCALPELLUM. Pls. V, VI.
SCALPELLUM. Leach. Journ. de Physique, t. lxxxv, July, 1817.
LEPAS. Linn. Systema Naturae, 1767.
POLLICIPES. Lamarck. Animaux sans Vertebres, 1818.
POLYLEPAS. De Blainville. Dict. des Sc. Nat., 1824.
SMILIUM (pars generis). Leach. Zoolog. Journal, vol. 2, July, 1825.
CALANTICA (pars generis). J. E. Gray. Annals of Philosophy, vol. x, (new series,) Aug. 1825.
THALIELLA (pars generis). J. E. Gray. Proc. Zoolog. Soc., 1848.
ANATIFA. Quoy et Gaimard. Voyage de l'Astrolabe, 1826-34.
XIPHIDIUM (pars generis). Dixon. Geology of Suffolk, 1850.
(Herm. et Foem.) Valvis 12 ad 15: lateribus verticilli inferioris quatuor vel sex, lineis incrementi plerumque convergentibus: sub-rostrum rarissime adest: pedunculo squamifero, rarissime nudo.
(Herm. and Fem.) Valves 12 to 15 in number: latera of the lower whorl, four or six, with their lines of growth generally directed towards each other: sub-rostrum very rarely present: peduncle squamiferous, most rarely naked.
Filamentary appendages, none: labrum, with the upper part highly bullate: trophi, various: olfactory orifices, more or less prominent: caudal appendages, uniarticulate and spinose, or none.
Males, parasitic at or near the orifice of the sack of the female or of the hermaphrodite: thorax enclosed within a capitulum, furnished with three or four rudimentary valves, or with six perfect valves: peduncle either short and distinct, or confounded with the capitulum: sometimes mouth and stomach absent, and cirri non-prehensile; sometimes mouth and cirri normal.
Generally attached to horny corallines, in the warmer temperate seas over the whole world.
I have felt much doubt in limiting this genus: the six recent species which it contains, differ more from each other than do the species in the previous genera. Mr. Gray has proposed or adopted generic names for four of the species, and a fifth certainly has equal claims to this same rank. These genera have been founded almost exclusively on the number of the valves; and oddly enough, the numbers have generally been given wrongly, namely, in Scalpellum, Calantica, Thaliella, and Xiphidium. Scalpellum blends through S. villosum into Pollicipes; and this latter genus has an equal right with Scalpellum, to be divided into sub-genera, three in number. Hence, no less than eight genera might be made out of the twelve recent species of Scalpellum and Pollicipes, and their formation, in some degree, be justified; but, in my opinion, this inordinate multiplication of genera destroys the main advantages of classification. At one time, I even thought that it would be best to follow Lamarck, and keep the twelve recent species in one genus; but considering the number of fossil species, I believe the more prudent course has been followed, in retaining the two genera Scalpellum and Pollicipes; more especially as I can hardly doubt, that several other species will be hereafter discovered.
Having so lately described in the Memoirs of the Palaeontographical Society, the fossil species, I will not here further allude to them, than to state, that out of the fifteen species therein described, S. magnum comes very close to the recent S. vulgare, and that several Eocene and Cretaceous species, such as S. quadratum, S. fossula, and S. maximum, are allied to S. rutilum and S. ornatum. Scalpellum villosum, a recent species, has stronger claims than any other species to be generically separated; and its habits, in not being attached to horny corallines, are also different, but the identity of its Complemental Male with that of S. Peronii, and its numerous points of resemblance in structure with the other species, have determined me not to separate it. Scalpellum Peronii, villosum, and rostratum, in having a sub-carina,—in the rostrum being pretty well developed,—and in the Complemental Male being pedunculated, and furnished with a functional mouth and prehensile cirri, may be separated from S. vulgare, ornatum and rutilum; but even between these two little groups, S. rostratum is in some respects intermediate, namely, in having three pairs of latera, and more especially in the rudimentary condition of the valves of its Complemental Male, and in the position in which the male is attached to the hermaphrodite. The three species in the second little group, namely, S. vulgare, S. ornatum, and S. rutilum, are more nearly allied to each other in all their characters, especially in the characters drawn from their Males, than are the other three species. S. ornatum and S. rutilum are considerably nearer to each other than any other two of the species. Upon the whole I conclude that the six species must be thrown either into five or into four genera (the first three species making one genus), or all into one genus, and this latter has appeared to me the preferable course. The separation even of Scalpellum and Pollicipes, as already stated, is hardly natural. The fact of these genera having existed from a remote epoch, and having given rise during successive periods to many species now extinct, is probably the cause that the few remaining species are so much more distinct from each other, than is common in the other genera of Lepadidae. Whenever the structure of the whole capitulum in the fossil species is well known, and as soon as more species, recent and fossil, shall have been discovered, then probably the genus Scalpellum will have to be divided into several smaller genera.
Description.—The Capitulum is much compressed, and generally produced upwards; it is formed of from twelve to fifteen valves, which are rather thin, and with the exception of S. ornatum, almost entirely covered by membrane, bearing spines: the valves are seldom locked very closely together. A sub-rostrum exists only in S. villosum, which species leads on to Pollicipes; in S. vulgare the rostrum is rudimentary and hidden. The scuta, terga and carina, are much larger than the other valves: these five valves seem to differ essentially from the others in being at first developed under the form of the so-called primordial valves: the other valves commence by a small indistinct brown spot, very different from the hexagonal tissue of the primordial valves: I saw this very clearly in young specimens of S. vulgare. At first, the scuta, terga and carina, grow exclusively downwards (and permanently so in most fossil species), and therefore the growth of the scuta and carina is in an absolutely opposite direction to what it is in Lepas, Paecilasma and Dichelaspis. After a short period the scuta are added to at their upper ends; the portion thus added, stands at a rather lower level, and projects in a rather different direction from the first-formed part of the valve, giving to it, in some respects, the appearance of having been broken and mended. This structure is common to S. vulgare, S. rostratum and S. Peronii. The upper Latera (except in S. villosum) grow in the same manner, namely, at first exclusively downwards, and then both upwards and downwards. The rostral and carinal latera (with the same exception of S. villosum) have their umbones seated laterally, at opposite ends of the capitulum,—the umbones of the rostral latera being close to the rostrum, and those of the carinal pair close to the carina, and consequently their chief growth is directed towards each other. The carina in all the species, except S. villosum, is either bowed or angularly bent; in the latter case the lower half is parallel to the peduncle, and the upper half, extending far up between the terga, is parallel to their longer axes. In some of the species the carina is added to almost equally at both ends; in S. ornatum it grows but little at the upper end, and to a varying degree in different individuals according to their age; in S. rutilum the umbo is at the apex, and there is consequently no upward growth; lastly, in S. villosum the carina widening much from the apex to the basal margin, grows exclusively downwards, and a portion of the apex projects freely,—characters all common to the carina in the genus Pollicipes. The upper latera occur in all the species; in the lower whorl there are either two or three pair of latera, in the former case the infra-median pair being absent. The latera differ considerably in shape in the different species.
The Peduncle is generally rather short, and, with the exception of S. Peronii, is covered with calcified scales. These scales are generally small, and placed symmetrically in close whorls, in an imbricated order, with each scale corresponding to the interspace between two scales in the whorls above and below. In S. ornatum, the scales are so wide, transversely, that there are only four in each whorl. In S. villosum, the scales are spindle-shaped and arranged somewhat irregularly in transverse rows, not very near to each other. New calcareous scales originate only round the top of the peduncle, and they continue to grow only in the few upper whorls; and as the peduncle itself continues to increase in diameter by the formation of new inner membranous layers and the disintegration of the old outer layers, the calcareous scales come in the lower part of the peduncle to stand further and further apart. In the earliest stage of growth there are no calcareous scales on the peduncle in S. vulgare; they first appear under the carina. Spines are articulated in great numbers on the surface of the peduncle in S. vulgare, S. Peronii, and S. villosum, and very short ones on that of S. rostratum.
Attachment.—All the species, except S. villosum, are attached to horny corallines: the singular means of attachment in S. vulgare will be described under that species, and is probably common to several of the other species. The larva in most, or in all cases, when it proceeds to attach itself, clings head downwards to the branch, and hence the capitulum comes to be placed upwards, with its orifice fronting the branch and the carina outwards. The sucking disc of the prehensile antennae of the larva, in the five species examined, was a little pointed, and in shape resembled the hinder hoof of a mule: this may perhaps be accounted for by the narrowness of the branches of the corallines, to which it has to adhere: a large circular disc, as in Lepas, would have been worse than useless: the ultimate segment in most or all the species, has on its inner side (the segment being supposed to be extended straight forward) a notch or step, bearing, I believe, two spines.
Size and Colour.—Some of the species attain a medium size, others are small. The valves are generally clouded red or pink, but sometimes white.
Mouth.—The various parts vary far more than in any genus hitherto described. The labrum is highly bullate, with the upper part forming a rounded overhanging projection, and with the lower part much produced, so that the mouth is placed far from the adductor scutorum muscle, and consequently the orifice is directed more towards the ventral surface of the thorax than in most other Cirripedes: on the crest of the labrum there are some very small teeth in several of the species, but not in all. The mandibles have either three or four main teeth, generally with either one or two small teeth intermediate between the first and second large teeth, and in the case of S. Peronii, with small teeth between all the larger ones. The maxillae have their edges furnished with many spines, and are either straight or have the inferior part prominent and step-formed. The outer maxillae have the spines on their inner edges either continuous or divided into two groups, of which latter structure we have not hitherto had any very well characterised example. The olfactory orifices are either highly or moderately protuberant.
In most of the species the prosoma is little developed, and the first cirrus is placed far from the second. The Cirri are generally but little curled, and have elongated segments, with long, generally serrated spines: the first cirrus varies in proportional length; the second and third cirri have both their rami more thickly clothed with spines than are the three posterior cirri, the spines being generally arranged in three or four longitudinal rows: the cirri, however, of S. villosum in all respects resemble closely the cirri of Pollicipes sertus and P. spinosus.
The Caudal Appendages are uniarticulate, small, and clothed with spines: in S. villosum, however, differently from in all other allied forms, there are no appendages.
The Stomach, in those species which I opened, is destitute of caeca. There are no filamentary appendages.
Generative System. The ova are nearly spherical, and remarkably large, as was stated to be the case in the introductory discussion, in which the larva of S. vulgare, in the first stage of development, was described: the ovigerous fraena are small. The testes are large, but the vesiculae seminales in some of the species extraordinarily small. Scalpellum ornatum, and perhaps S. rutilum, are unisexual; the other species are hermaphrodite, but most or at least some of the individuals, are furnished with Complemental Males. These latter are fully described under each species, so I will here only remark, that S. ornatum, which alone (excepting perhaps S. rutilum) is unisexual, has less claim than the other species to be generically separated: we have seen also, in Ibla, that similar sexual differences occur in two most closely allied species. It is very singular how much more some of the Males and Complemental Males in Scalpellum differ from each other, than do the female and hermaphrodite forms; this seems due to the different stages of embryonic development at which the males have been arrested. In the males, however, of S. rostratum, S. Peronii, and S. villosum, compared one with another, but not with the males of the other species, the parts of the mouth and apparently the cirri, resemble each other more closely, than do the same organs in the hermaphrodites. At the end of this genus I shall give a summary on the highly remarkable sexual relations both in Scalpellum and Ibla.
Distribution.—The species seem distributed over the whole world, but as far as we can trust our present scanty materials, are most common in the warmer temperate regions. The S. vulgare ranges from the Norwegian seas to Naples. Most of the species are inhabitants of deep water.
Affinities.—In the preliminary remarks, we have seen how this genus blends into Pollicipes; and under the head of Oxynaspis, I have shown its close affinity to that genus. If, indeed, we take Pollicipes spinosus, and destroy all but six of the already minute and almost rudimentary latera, we shall, as far as the capitulum is concerned, convert it into a Scalpellum, closely similar to S. villosum. If we take any species of Scalpellum, (excepting S. villosum and S. rutilum,) and destroy all the valves, but the scuta, terga and carina, we shall convert it into an Oxynaspis. Lastly, I have shown under Ibla, that in several most remarkable peculiarities of structure, there is a manifest affinity between Scalpellum and that genus.
Geological History.—Full details on this subject have been given in the Memoirs of the Palaeontographical Society. I will here only state, that the oldest known form of Scalpellum occurs in the Lower Green Sand.
[T SUB-CARINA NULLA.]
1. SCALPELLUM VULGARE. Pl. V, fig. 15.
SCALPELLUM VULGARE. Leach. Encyclop. Brit. Suppl., vol. iii, 1824.
LEPAS SCALPELLUM. Linn. Systema Naturae, 1767.
—— Poli. Test. utriusque Siciliae, Pl. vi., fig. 16. 1795.
POLLICIPES SCALPELLUM. Lamarck. An. sans Vertebres, 1818.
POLYLEPAS VULGARE. De Blainville. Dict. Sc. Nat., Plate, fig. 4. 1824.
SCALPELLUM LAEVE, var. Leach. Zoolog. Journal, vol. ii, p. 215, 1825.
—— SICILIAE, var. Chenu. Illust. Conch. Pl. iv, fig. 9.
SCALPELLUM VULGARE, (et var.) Brown. Illust. of Conch., 1844, Pl. li., figs. 7 to 20.
S. (Herm.) valvis 14, si rostrum paene rudimentale includatur: lateribus superioribus inaequaliter ovatis.
(Herm.) Capitulum with 14 valves, including the rudimentary rostrum: upper latera irregularly oval.
Mandibles, with four or five teeth: maxillae, with the edge straight, bearing numerous spines.
COMPLEMENTAL MALE flask-formed, with four rudimentary valves; no mouth; cirri not prehensile; attached to the occludent margin of the scutum, near the umbo.
Great Britain, Ireland, France, Norway, Naples. Attached to horny corallines, at from twenty to thirty, sometimes even to fifty fathoms in depth, according to Forbes and MacAndrew.
HERMAPHRODITE.
Description.—Capitulum much flattened with the apex produced, of a pale brown colour, sometimes faintly tinted purple, composed of 14 valves, of which the rostrum is rudimentary and barely visible externally; valves thin, white, translucent, smooth, slightly marked by the lines of growth, separated from each other by rather wide interspaces of colourless membrane, which is thickly clothed by small, articulated spines of unequal length. The valves, excepting sometimes their umbones, are also covered with membrane, bearing spines, placed in rows parallel to the lines of growth; the spines are particularly numerous round the orifice of the sack.
Scuta slightly convex, thrice as long as broad; upper part much acuminated; occludent margin almost straight; basal margin nearly at right angles to the occludent margin; the tergal margin is separated from the lateral margin by an angle more or less prominent; a slight curved ridge runs from the umbo to this angle, and this deserves especial notice, inasmuch as it indicates the outline which the valve assumed in its earliest growth, and which is permanently retained in most of the older fossil species. Along the occludent margin, there is a trace of a ledge, developed in a variable degree, and which is noticed only on account of the plainly visible ledge along this same margin, in the allied genus Oxynaspis. The umbo, or centre of calcification, is seated close to the occludent margin, and at about one fourth of the length of the valve from the apex. Internally, (fig. 15, a', Pl. V,) the part above the umbo is flat; and beneath this upper part, there is a large rounded hollow (d) for the adductor muscle: a fold or indentation (a) running downwards from the umbo, extends in a very oblique line across the occludent margin. This fold is of high interest as giving lodgment to the Complemental Males, and will hereafter often be referred to.
Terga, triangular, flat; occludent margin, very slightly arched.
Carina much bent, with the umbo placed at barely one third of the entire length of the valve from the apex. Two very slight ridges can be perceived, one on each side, running from the umbo to the basal margin, and separating the roof from the parietes of the valve; these ridges are of great use in distinguishing the fossil carinae of Scalpellum, from the carinae of Pollicipes. The part above the umbo is formed by the upward production of a marginal slip along each side of the valve, which slips in the fossil species (C in the woodcut, fig. 1, given in the Introduction,) I have designated as the intra-parietes. The lower part of the valve gradually widens from the umbo downwards; internally, the whole is deeply concave, and continuously curved. The angle varies at which the upper and lower portions externally meet each other; but is never less than 135 deg. The upper part of the carina runs up between the terga for three-quarters of their length; the basal margin does not extend down low enough to pass between the carinal latera.
Rostrum, (fig. 15 b', seen externally, and highly magnified,) minute, almost hidden by the enveloping membrane and by the small prominent umbones of the rostral latera; in area equalling about one fourth of the rostral latera; externally pyramidal, with the upper side rather longer than the lower; internally slightly concave, square, with the upper margin and sometimes with the lower margin, slightly hollowed out. Umbo of growth nearly central.
Upper Latera, flat, irregularly oval, with an almost rectangular shoulder under the basal angle of the terga; in area, about one third larger than the largest valve of the lower whorl; the exact degree of elongation of the oval figure varies a little. Umbo seated a little above the central point.
Lower Whorl,—Rostral Latera, nearly twice as long as broad, lying under the basal margins of the scuta: umbo seated over the rostrum; opposite end, towards which the valve widens either sensibly or but little, is either square or rounded; in area, less than any of the other valves, excepting the rostrum; in breadth, equalling either half or one third of the height of the infra-median latera; growth, directed chiefly towards the infra-median latera. The freely-projecting umbo is about one sixth part of the entire length of the valve.
Infra-median Latera, rather larger than the carinal latera; their shape varies from elongated pentagonal with the angles rounded, to oval, with the longer axis directed upwards. The umbo is seated a little above the middle of the basal margin, so that there is some little growth downwards, but the main growth is upwards. The upper point generally stands a little above that of the carinal latera.
Carinal Latera, flat, less in area than the infra-median latera; basal margin nearly straight; carinal margin slightly hollowed out, terminal margin arched and protuberant. The umbones of the two valves almost touch each other under the middle of the carina; main growth towards the infra-median latera and upwards; umbones projecting not above one fifth of the entire length of the valve.
Peduncle, much flattened, rarely as long as the capitulum, with the upper end nearly as wide as it; the lower end is either blunt, or tapers to a very fine point. The calcareous scales are transversely elongated, and are about four times as wide as high; their internal surfaces are slightly concave, and their external, convex; the two ends are pointed. Viewed internally, the scales approach in shape to rhomboids. There are, in a medium-sized specimen, about twenty scales in each whorl, their tips overlapping each other: the whorls are placed not very near each other and at rather unequal distances, except round the uppermost part, where, being in process of formation, they are packed closely together. The membrane uniting the scales, supports numerous transverse rows of articulated spines, varying from 1/100th to 1/500th of an inch in length, and each furnished with a long sinuous tubulus, 1/10,000th of an inch in diameter, running through the membrane to the underlying corium.
Attachment.—Specimens are attached to various horny corallines, and occasionally to the peduncles of each other.[51] In both cases, supposing the coralline to be erect, the capitulum is placed upwards, with its orifice towards the branch to which it is attached, and consequently with its carina outwards. Where several are crowded in a group, their peduncles often become twisted and their positions irregular, with their orifices facing in any direction. This uniform position is simply the consequence of the larva attaching itself head-downwards, and from the position of the prehensile antennae, necessarily with its sternal surface parallel and close to the branch of the coralline; hence the dorsal surface, which afterwards is converted into the carina, faces outwards. The peduncle, as already stated, often tapers, at its basal extremity, to a sharp point. In very young specimens, for instance in one with a capitulum only 1/20th of an inch in length, the method of attachment is the same as in Lepas and many other genera, namely, by cement proceeding exclusively from the antennae of the larva; but in older and full-grown specimens, instead of the whole bottom of the peduncle becoming flattened and broadly attached, which would be here impossible, the cement is poured out through a straight row of orifices along the rostral edge, thus causing, by an excellent adaptation, a narrow margin to adhere firmly to the thin and cylindrical branches of the coralline. These orifices are represented, magnified seven times, in Pl. IX, fig. 7, in which the lower attached portion of the peduncle is split open and exhibited; they are circular, and stand at regular intervals, in a straight line; the higher orifices are larger, but further apart from each other than the lower ones; in one full-grown specimen, I counted ten of these orifices in a length of exactly a quarter of an inch. At each period of growth, the corium recedes a little from the attached portion of the peduncle; of which portion, the greater part is thus left empty and as incapable of further growth, as are the larval antennae at the extreme point: in the specimen figured, the corium extended a little below the upper orifice. The prehensile antennae, however, I must remark, do not strictly rise from the extreme point of the peduncle, but at a little distance from it, on the rostral surface; this simply ensues from the antennae in the larva, being situated on the sternal surface, close to, but not actually on the front of the head. The two cement glands are seated high up on the sides of the peduncle, and remote from each other; they are small, unusually globular and transparent. The two cement-ducts (fig. 7 a a) proceeding from them, are 3/2000ths of an inch in diameter, and run in a zig-zag line; at the point where they pass through the corium to enter the lower attached portion of the peduncle, they become closely approximated, and partially imbedded in the membrane of the peduncle. Together they run along the rostral edge, giving out through each orifice a little disc of brownish cement, and finally they enter the larval antennae. The peduncle, just above the attached portion, where still lined by corium, no doubt increases in diameter at each period of growth, and must, I presume, become pressed against the almost parallel branch of the coralline. The corium, at this same period, shrinks, or is absorbed, and the two cement-ducts come in contact with, and adhere to, the inner surface of the outer membrane of the peduncle; and then, by a process which I do not understand in this or any other Cirripede, apertures are formed both in the ducts and through the membrane, so that the cement passes through, firmly fastening the outer surface of the peduncle with its calcareous scales and spines, to the coralline.
[51] Mr. Peach, (Transact. Brit. Assoc., 1845, p. 65,) states that this is sometimes the case in Cornwall; and I have seen a similar instance in a fine group from Naples.
The structure of the larval prehensile antennae will be most conveniently described when we come to the Complemental male; and figures (10-12, Pl. V) will be given.
Size and Colours.—Montagu states ('Test. Brit.,' p. 18) that British specimens rarely have a capitulum .62 of an inch in length; I have, however, seen an Irish specimen, .7 long; and several specimens, from the Bay of Naples, .8 long, and including the peduncle, 1.3 in length. The valves in all the specimens are white, and the membrane connecting them either nearly white, or dirty pale yellowish, or purplish-brown. Within the sack the corium under the valves is tinted pale purple, and two very faint bands of the same colour can generally be distinguished running down the two sides of the peduncle. Body, coloured yellowish-white, with the upper segments of the pedicels of the cirri, tinted in front with purple.
Body, much flattened, the prosoma is very little developed; the mouth placed far from the adductor muscle, and is directed in a remarkable manner towards the ventral surface of the thorax: the first pair of cirri stands far separated from the second pair.
Mouth.—Labrum with the upper part highly bullate, forming an overhanging projection equalling the longitudinal axis of the mouth; basal margin much produced; crest with a row of bead-like teeth.
Palpi rather small, with their external margin straight, and internal margin oblique: the bristles on the two palpi just meet each other.
Mandibles, with five or six teeth, with the second, (or second and third, when there are six teeth,) smaller than the others; in two specimens, there were five teeth on one side and six on the other; inferior angle rather broad and strongly pectinated.
Maxillae with the edge nearly straight, without any notch, but with the inferior portion very slightly projecting; there are twelve or thirteen pairs of unequal spines, of which some of the middle ones are rather longer than the others, and almost as long as the two upper great spines.
Outer Maxillae.—On the inner margin the bristles are divided into two separate tufts; exteriorly, near the base, there is a distinct rounded swelling with bristles. The olfactory orifices are highly protuberant, approximate, flattened, scarcely tapering towards their upper ends.
Cirri.—The five posterior pair are elongated, very little curled, with short pedicels; their segments are long, not at all protuberant in front, bearing five or six pairs of long, slightly serrated spines, with a very minute tuft of bristles between each pair, and with some short lateral spines on the inner side of each segment; on the fourth pair of cirri, these lateral spines are considerably developed; dorsal tufts consist of fine spines, with one much longer than the others. First pair short, separated by a wide interval from the second; rami unequal in length, by between two and four segments; longer ramus having nine segments, scarcely half as long as the rami of the second cirrus; shorter ramus with seven segments; in the same individual there were twenty segments in the sixth cirrus. The segments in the shorter ramus of the first cirrus are oblong in a transverse direction, and may be compared to a set of shields placed transversely and strung together; in the longer ramus the segments are longitudinally oblong; in both they are thickly covered with spines. Second cirrus; the anterior ramus is a little broader than the posterior ramus, with the segments bearing about five rows of bristles; fifteen segments in the shorter ramus. Third pair, with the two rami equal in thickness, and with the segments differing very little from those of the posterior cirri, excepting that the serrated spines in the external lateral rows are rather larger. The fourth pair is remarkable by having, on the inner side of the upper edge of each segment, a little tuft of minute smooth spines, flattened, and a little enlarged near their ends, so as to be spear-shaped; I could not see these singular spines on the other cirri. The lower segments of the pedicels of all the cirri, excepting the sixth pair, are remarkable from having their inner edges, in the middle, produced into a considerable, abrupt, rounded projection, irregularly covered with spines.
Caudal Appendages, (Pl. X, fig. 21,) very small, flattened, of nearly the same width throughout; in a medium-sized specimen, only 1/100th of an inch in length; each bears from ten to twenty small bristles placed distantly from each other, of which those on the rounded apex are the longest.
Generative System.—The penis is remarkably acuminated; the vesiculae seminales are unusually small, and enter only for a short distance into the prosoma; the testes are large. The ovarian tubes are of large diameter; the ova are nearly spherical and large, namely, 9/400ths of an inch in diameter; they are not numerous, and lie in single layers in the two lamellae. The ovigerous fraena are well developed, and lie under the scuta; one I measured was 5/100ths of an inch in length and 2/100ths in width; the margin is obliquely truncated and slightly sinuous. This species breeds late in the autumn, and even in mid-winter; I have examined a specimen from Cornwall with ova containing larvae, taken on the 26th of October; again, in another specimen from Belfast, sent to me by Mr. Thompson, taken in January, there were ova in the lamellae, and therefore no doubt impregnated; and on February the 12th I received from Mr. Peach, from Cornwall, specimens so very young that they must have become attached during the first days of the month.
Varieties.—The specimens from near Naples, (which I owe to the kindness of the Rev. F. W. Hope,) are somewhat larger, and differ slightly from those of Britain: they form, I imagine, the S. Siciliae of Chenu. After carefully examining them internally and externally, I think it is quite impossible to consider them specifically distinct, for although in several specimens, the valves were placed a little further apart from each other,—the upper latera a little more elongated,—the carinal latera rather narrower in their upper half,—the infra-median latera rather more rounded,—and, lastly, in the scuta, the tergal margin extended almost in the same line with the lateral margin; nevertheless in other specimens, I could perceive no difference whatever. It is, however, remarkable that in several full-grown Neapolitan specimens there were no Complemental males, whereas I have never seen a single full-grown British specimen without such being present. In some specimens in the British Museum, without any given locality, I have observed considerable variation in the breadth of the carinal and rostral latera.
COMPLEMENTAL MALE. Pl. V, figs. 9-14.
When first dissecting Scalpellum vulgare, I was surprised at the almost constant presence of one or more very minute parasites, on the margins of both scuta, close to the umbones: these are represented, but rendered darker and therefore more conspicuous than in nature, in the drawing, Pl. V, fig. 15, which is three times the natural size. I carelessly dissected one or two specimens, and concluded that they belonged to some new class or order amongst the Articulata; but did not at that time even conjecture, that they were Cirripedes. Many months afterwards, when I had seen in Ibla, that an hermaphrodite could have a complemental male, I remembered that I had been surprised at the small size of the vesiculae seminales in the hermaphrodite S. vulgare, so that I resolved to look with care at these parasites; on doing so, I soon discovered that they were Cirripedes, for I found that they adhered by cement, and were furnished with prehensile antennae, which latter, I observed with astonishment, agreed in every minute character, and in size, with those of S. vulgare: the importance of this agreement will not at present be fully appreciated. I also found, that these parasites were destitute of a mouth and stomach; that consequently they were short-lived, but that they reached maturity; and that all were males. Subsequently the five other species of the genus Scalpellum were found to present more or less closely analogous phenomena. These facts, together with those given under Ibla (and had it not been for this latter genus, I never probably should have even struck on the right track in my investigation,) appear sufficient to justify me, in provisionally considering the truly wonderful parasites of the several species of Scalpellum, as Males and Complemental Males. When these parasites are fully described, will be the proper time to discuss and weigh the evidence on their sexual relations and nature. I will now describe the parasite of S. vulgare.
General Appearance.—Shape, flask-like, compressed (Pl. V, fig. 9, magnified 36 times), with a short neck: the outline is usually symmetrical, but sometimes is a little distorted on the under side. The creature is imbedded more than half its length or depth in the transparent, spine-bearing chitine border of the scutum of the hermaphrodite. Its length, or longer axis, varies from 10 to 11/400ths; its breadth, or transverse axis, is 6 to 7/400ths; and its thickness, for it is much flattened, is only 4/400ths of an inch. On the summit, there is a fimbriated orifice (a), the size of which can rarely be made out quite distinctly, owing to the extreme thinness of the membranous edges. A little way beneath the orifice, there are four little blunt, bristly points (b), generally rather more than the 1/1000th of an inch in length; they are rather variable in size, and seem to be of no functional importance; directly beneath them, there are four little calcareous beads (as may be known by their dissolving with effervescence in any acid, and breaking easily under the needle); these are the 3/2000ths of an inch in their larger external diameter; they are rather deeply imbedded in the outer integument, and taper a little downwards ending in a concave terminal point, into which a minute tubulus enters, like those passing into and through the valves of ordinary Cirripedia: along the axis of imbedment, they are often 4/2000ths of an inch in length. These calcareous beads or rudimental valves are seated in pairs, at the two ends of the flattened animal, so that when the animal is laid on one side, the upper bead in each pair exactly covers and hides the lower one. The outer integument is composed of chitine, as may be inferred from boiling caustic potash having no effect on it; the upper part is thicker than the imbedded portion and is wrinkled transversely; it is covered with minute spines 4/10,000ths of an inch in length, either single or in groups of two and three, (Pl. V, fig. 14.) This outer tunic is lined by corium, sometimes slightly mottled with dull purple; and this by delicate, longitudinal, striae-less muscles, running from the base up to the under edge of the orifice; these longitudinal muscles are crossed, at least, in the upper part, by still finer transverse muscles.
Thorax and Abdomen.—When the external integument is cut open, the thorax (Pl. V, fig. 13) is found lodged within an inner sack or rather tube, extending from near the bottom of the animal, up to the external orifice. The whole thorax is sometimes forced through the orifice, owing perhaps to the action of the spirits of wine and consequent endosmose, and is thus well displayed without dissection. The thorax tapers a little, is much flattened and straight; its length, together with the terminal abdominal lobe, is about 6/400ths of an inch; it is formed of very thin, most finely hirsute membrane, transversely wrinkled and so extensible, that when everted by the internal muscles being seized, it stretches to twice its former length; in this condition, five transverse articulations are displayed. The abdominal lobe is smooth, and cannot be stretched, or turned inside out by pulling the above muscles. On the thorax, corresponding with the interspaces between the five transverse articulations, there are four pair of short limbs, but their bases, I believe, are prolonged across the inner or ventral surface of the thorax, so as almost to touch each other. These limbs, I believe, have no articulations, except, perhaps, where united to the thorax. The anterior or lowest limb, on each side, supports two or sometimes only a single spine; this pair is rather smaller than the second, and is placed a little more distant from it, than are the upper pairs from each other. The second pair differs from the upper two, only in having its three spines a very little shorter. The two upper or posterior pair exactly resemble each other; each has two spines on the summit, and a third seated lower down, on a little notch on the outer side, but with its point on a level with the others. The points of the spines of the two upper limbs, stand on a level with the external spines at the end of the abdomen. All the spines are of excessive tenuity and sharpness; they are straight, long, and not plumose.
The abdominal lobe is square, and from not being wrinkled, has a different appearance from the thorax: on each of the posterior angles, there are three moderately long, very sharp spines, with the tips of the outer pair bent a little inwards; in the middle between them, there are two little spines, and a little below and outside these latter, on the ventral surface, there are two other longer spines with their tips bent inwards; and again, lower down, two other pair, one beneath the other, of short spines. Perhaps, the three pair of spines on the ventral surface, mark the three segments, which are distinct on the abdomen of the larva in the last stage of its development, in Lepas and other genera. In the same way, it is probable that the lateral spine on the notch in each limb, marks the point where, in the larva, there is an articulation. Altogether, there are seven pairs of spines on the abdomen, and eleven pairs on the thoracic limbs.
A little way beneath the lower or anterior pair of limbs, the thorax is abruptly bent, and becomes confluent with the lower internal parts of the whole animal. Here, the very delicate membrane of chitine which lines the sack or tube, extending from the external orifice, can be seen to be continuous, as in all Cirripedes, with the outer tunic of the thorax. Within the thorax, there are some longitudinal muscles, without transverse striae, which, I believe, enter the short limbs, but not the abdomen, as I infer from the latter not being everted when they are pulled. At their lower ends these muscles terminate abruptly, and from being contracted are often a little enlarged. They extend a short way beneath the lower pair of limbs, and are, I suspect, attached to the outer integument of the animal, near the base.
After the most careful dissection of very many specimens, and their examination in many different methods (as by caustic potash, &c.), I can venture positively to assert that there is no vestige of a mouth, or masticatory organs, or stomach: I did not see any anus, but I will not affirm that such does not exist.
In the upper part of the animal, lying under the superficial muscles, and close beneath the upper line of their attachment, I found in all the specimens, an eye, of a pointed oval form, rather less than 11/12,000ths of an inch in diameter, formed of an outer capsule, lined with purple pigment-cells, and surrounding, as it appeared, a lens. The eye is not introduced in fig. 9, for I could not see it, except by dissection, and therefore do not know its exact relative position.
Generative System.—The contents of the animal, between the sack containing the thorax and the outer integuments, and directly under the thorax, varied much in condition: in young and lately attached specimens the whole consisted of a pulpy mass with numerous oil-globules; in other specimens, apparently more mature, there were vast numbers of cells, sometimes cohering in sheets, about 3/10,000ths of an inch in diameter, and having darkish granular centres; these I believe to be the testes, for in a specimen presently to be mentioned, in which the vesicula seminalis was gorged with spermatozoa, I found adhering to its outside, a mass of cells of exactly the same diameter, but now empty and transparent instead of having brownish centres. Lastly, in several other specimens, at the very bottom of the sack-formed animal, there was a brownish, pear-shaped bag, of different sizes in different individuals, and occasionally broader even than the thorax. This bag contained either pulpy matter, or a great mass of spermatozoa. Before being disturbed, these spermatozoa lay parallel to each other in flocks, and they yielded to the needle in a peculiar manner, so that I found (having had experience with these bodies in living Cirripedia) I could almost tell before examination under the compound microscope, whether or not I should see spermatozoa. Many had distinct heads,[52] which were two or three times as broad as the filamentary bodies; the latter when placed between glass were the 1/20,000th of an inch in diameter. I compared these spermatozoa with others taken out of the vesiculae seminales of the individual hermaphrodite S. vulgare, to which the parasite was attached, and could not perceive the slightest difference in them. The brownish pear-shaped bag, or vesicula seminalis, the coat of which seems fibrous, could sometimes be distinctly traced, sending a chord or prolongation far up the thorax: at the end of the abdominal lobe, no doubt there is an orifice; and this, I believe, I once distinguished. Owing to this chord, the bag often adheres to the thorax, when the latter is dissected out of the general integuments; in this condition, I twice clearly made out that it was single: in one other specimen, however, there appeared to be two small vesiculae seminales. By using a condenser and very brilliant light, the outline of the vesicula seminalis could sometimes be distinguished before dissection, at the bottom of the sack-formed animal; and such was the case in the specimen drawn in fig. 9.
[52] I do not understand the development of the spermatozoa in Cirripedia: in a recent Chthamalus and Balanus, I found the greater number had a little filament in front of the head or nodular enlargement, which latter varied in size and in shape from globular to that of a spindle. The filament before the head, also, varied in proportional length; it did not project in exactly the same straight line with the hinder part, and some of the spermatozoa were entirely without this filament in front;—such is the case with the spermatozoa here described.
Although I have dissected, at least, thirty specimens, taken at different times of the year, and from different localities, and when many of the specimens were mature and ready for the impregnation of ova, as clearly shown by the presence of innumerable spermatozoa, I have never seen even a trace of an ovum or ovaria.
Antennae and Attachment.—The prehensile antennae (Pl. V, fig. 10), are seated a little above the very base of the sack-like animal; and this might have been expected from the antennae in the larva, being seated on the ventral surface, not at the very extremity of the head. By a very strong light, they can sometimes just be seen whilst the parasite is attached to the hermaphrodite (the scutum of the latter having been cleaned on the under side), and are thus represented in fig. 9. They are formed of thicker membrane than the general integument of the body: the second segment, or disc, is pointed and hoof-like; when seen in profile (fig. 11), the upper convex surface has a uniform slope with the upper surface of the basal segment; it is furnished with a single backward pointing spine, attached, I believe, on the under side, nearly opposite the articulation of the ultimate segment: at the apex, there are some excessively minute hairs or down. The ultimate segment projects rectangularly outwards as usual, and has on its inner side, rather beneath the middle, a conspicuous notch (fig. 12), which bears two or three long, non-plumose spines; on the summit there are three or four rather shorter spines. On the outside of the great basal segment there is a single spine curving backwards. The importance of the following measurements (in fractions of an inch) will hereafter be seen.
Length of whole organ, from end of disc to the further } margin of the oblique basal articulation } 38-39/6000
Length of whole organ, to the inner margin of the oblique} basal articulation } 1/6000
Breadth of basal segment, measured half-way between the } basal and second articulations,—the limb being viewed } from vertically above } 8/6000
Length of hoof-like disc, measured from the apex to the } middle of the articulation with the basal segment } 9-10/6000
Breadth of ditto 5/6000
Length of ultimate segment 6/6000
Breadth of ultimate segment beneath the notch 7/20000
Breadth of ultimate segment above the notch 5/20000
I did not see the cement-ducts, which, perhaps, was owing to the corium extending from the inside of the whole animal some way into the antennae, thus rendering them rather less transparent than in common Cirripedes. That the ducts and cement-glands exist, is certain, for the antennae in every case were enveloped in a little irregular mass or capsule of the usual, brown, transparent, laminated cement. When several of these parasites were attached close together, the cement ran up between them.
I may here state, that I found on one Scalpellum, three males very lately attached, and not as yet imbedded in the chitine border; they were white, opaque, pulpy, and full of oily globules; the lower part was considerably more pointed, and extended further beyond the prehensile antennae, than in the older and imbedded specimens. There were distinct remnants of two great reddish-brown eyes, showing that in this respect the larvae of the male in their last stage of development, are characterised like the larvae of other Lepadidae. The male larva would, probably, be a little larger than the male itself; but yet compared with the larva in the earliest stage, there can have been unusually little increase of size during the several intermediate metamorphoses; I judge of this from the dimensions of the larva of the hermaphrodite in the first stage, namely, 9/400ths of an inch, exactly the size of some of the smaller males. In the allied genus Ibla, the increase is also less than is usual, namely, from 15/1000ths of an inch, the diameter of the ovum, to only 25/1000ths of an inch, the length of the boat-shaped larva, just before its final metamorphosis.
Habits and Concluding Remarks.—The males are imbedded in the spinose chitine border of the occludent margin of the scuta, exactly over an oblique fold or notch (fig. 15 a a), close by the umbo. This fold has no direct relation to the males, but being present is taken advantage of by them; for it occurs in the young hermaphrodite, before the attachment of the males, and in species of the genus in which the males are attached to other parts. It occurs, also, in fossil species of Pollicipes, and in these it seems caused by the upper inner part of the valve being rendered more and more prominent during growth: in the present species, I suspect, its origin is connected with the formation of a ridge bounding the outer side of the pit for the adductor scutorum muscle: we shall see in the next species, that this fold is of the highest importance in relation to the position of the Males. The transparent chitine border of the scuta is broad, and fills up the fold in the shell, so that the outline of the occludent margin is not affected by it: in the drawing (fig. 9) some of the inner layers of chitine (e e), which dipped into and filled up the fold, have been removed, that the lower part of the animal might be more plainly exhibited. The chitine bears numerous spines of various lengths, which must afford some protection to the males, rudely arranged in lines, parallel to the edge of the valve, indicating the successively-formed layers of chitine; each spine has a fine, tortuous tubulus connecting its base with the underlying corium. The extreme outer edge of the border is thin, forming a kind of lip, close beneath which the delicate tunic lining the sack is attached. During continued growth, the valve is added to in thickness, and so is the chitine border, and likewise in breadth. It appears that the larva of the male must attach itself on the under side of this border, on the edge of the tunic of the sack, and that by the action of the cement, the corium beneath is killed (as I believe always is the case with other parasitic Cirripedia), whereas on both sides, the chitine continues to be added to, so that the male, excepting the upper and always projecting portion, becomes imbedded at first laterally, and ultimately all round: I have seen specimens in several different stages of imbedment. Hence, in old specimens, with a thick and broad chitine border, it might and does come to pass that one male is imbedded (the valve being laid flat) directly beneath another.
I have examined a great number of specimens from various localities, taken at different times of the year,—some dozen specimens from Cornwall,[53] and several from unknown localities in various collections; some from Ireland, from the Shetland Islands, from Norway, and from near Naples. Every one of these specimens, with the exception of some of the Neapolitan ones, had parasitic males attached to them: I must also except very young specimens, on which they never occur. On a Cornish specimen, with a capitulum a little more than one fifth of an inch in length, it may be mentioned as unusual that there were three males. In young specimens there is generally one male on each scutum, but sometimes there are two, and sometimes none on one side. In large old Cornish specimens I have counted on the two sides together, six, seven, and eight males, and in one Irish specimen no less than ten, seven all close together on one valve and three on the other, but I do not suppose that all these were alive at the same time. In the Neapolitan specimens, however, which are the largest that I have seen, there was in no case more than two; and out of seven or eight specimens, four had not any male; so that it would appear there is something in this locality hostile to the development of the parasitic males. I have noticed only one instance (that given in fig. 9) in which the males were imbedded a little way apart; generally they touch each other, and are cemented together: where there are several males, they occur at different levels, as measured from the under or upper surface of the chitine border: in one instance of four males adhering to one valve, I distinctly perceived that the lowest one was white, pulpy, and recently attached; the two above, which were placed close together and between the same laminae of chitine, were mature; and the third still higher up, was dead, empty, transparent, and half decayed: in some other instances, I have found the uppermost parasites dead, and, together with the surrounding chitine, partially worn away.
[53] I am greatly indebted to Mr. Peach for his unwearied kindness in procuring me fresh specimens. Mr. W. Thompson allowed me to dissect one, possessing particular interest, out of his three Irish specimens. Professor Forbes procured me a specimen from the Shetland Islands, and Professor Steenstrup was so kind to take pains to send me some Scandinavian specimens.
The larva of the male must have a different instinct from the larva of the hermaphrodite; for the latter attaches itself head downwards to a coralline, whilst the male larva crawling on the scuta of the hermaphrodite, discovers, I presume by eye-sight, the fold in the shell beneath the translucent border of chitine, and there invariably attaches itself. Its object in choosing this particular spot, I believe, simply is that the depth or thickness of the chitine is there greater, and sufficient for its imbedment, which would hardly be the case elsewhere. This parasite has, as we have seen, no mouth or stomach, and indeed, considering its fixed position and the non-prehensile condition of its limbs or cirri, a mouth would have been of no service to it, without it had been extraordinarily elongated. The male must live on the nourishment acquired during its locomotive larval condition; and its life no doubt is short, but yet not very short, as I infer from the depth to which mature specimens are buried in the chitine border. The full development of the spermatozoa consumes, I suppose, some considerable lapse of time. The thorax and limbs, though furnished with muscles, are obviously, as already remarked, of no use for prehension; these parts serve, probably, to defend the little creature, when its eye announces the passing shadow of some enemy, and for this purpose they are well adapted from the extreme sharpness of the spines. The thorax, into which I traced the vesicula seminalis, no doubt also serves for the emission and first direction of the spermatozoa; and hence, perhaps, its singularly extensible structure. I have already remarked, that in specimens preserved in spirits, the thorax is often largely protruded, and bent down at right angles to the orifice. I presume this is caused by endosmose; nevertheless it deserves notice, that it was in these protruded specimens that the vesicula seminalis was most conspicuously gorged with spermatozoa. I suspect the longitudinal and transverse muscles lining the upper part of the outer integuments of the whole animal, can be of little use to the creature, without it be to aid in the protrusion of the thorax, and perhaps in the violent expulsion of the spermatozoa, thus causing them to reach the ovigerous lamellae within the sack of the hermaphrodite. It is also probable, that the action of the cirri of the hermaphrodite, would tend to draw inwards the spermatozoa in the right direction. In one specimen, the spermatozoa in the hermaphrodite and in the male were mature at the same time; in another this was not the case; and as the males, apparently, become attached at all periods of the year, this want of coincidence in maturity must often occur. Can the males retain their spermatozoa, till told by some instinct, that the ova in the sack of the often fecundated hermaphrodite are ready for impregnation; or are the spermatozoa sometimes wasted, as must annually happen with such incalculable quantities of the pollen of many dioecious plants?
This little Cirripede is, in many respects, in a partially embryonic condition. There is no separation between the capitulum and peduncle; there is no mouth; and the thorax, throughout its whole width, opens into the anterior part of the animal: the limbs differ greatly from those both of the mature Cirripede and of the larva, but come closest to the latter: the preservation of the abdomen is a well-marked embryonic character. On the other hand, the four rudimentary calcareous valves, the narrow orifice, the hirsute outer integument, the two muscular layers, the single eye, and male internal organs, are all characteristic of the fully-developed condition. The four little valves, as I believe, represent the scuta and terga, though they are placed considerably below the orifice: the little bristly points have no homological signification, and are absent in the male of the following closely allied species. The four pairs of limbs answer to the four posterior cirri, as may be inferred from their proximity to the abdominal lobe, and from the three posterior pairs closely resembling each other, and differing a little from the first pair; this latter pair corresponds with the third pair in the hermaphrodite form of Scalpellum. If I am right in believing that only a single vesicula seminalis is ordinarily developed in the male, this is a special and singular character.
As stated in the beginning of this description, from the one great fact of the absolute correspondence of the prehensile antennae of the parasite, with those of the hermaphrodite Scalpellum vulgare, together with its fixed condition, its short existence, and exclusively male sex, I have thought myself justified in provisionally considering it as the Complemental Male of the Cirripede to which it is attached; but I hope final judgment will not be passed on this view, until the whole case is summed up at the end of the genus.[54]
[54] I trust, before long, that some naturalist, with more skill than I possess, will examine these parasites on Scalpellum vulgare, which unfortunately is the only species of the genus that can be easily obtained. Fresh specimens, or those preserved in spirits of wine, are necessary. The action of boiling caustic potash is very useful in cleaning the prehensile antennae. If these latter organs are sought in the hermaphrodite for the sake of comparison, young specimens, adhering to clean branches of a coralline, should be procured, and caustic potash used.
2. SCALPELLUM ORNATUM. Pl. VI, fig 1.
THALIELLA ORNATA. J. E. Gray. Proc. Zoolog. Soc., 1848, p. 44, Annulosa, Plate.
S. (Foem.) valvis 14, sub-rufis: lateribus superioribus quadranti-formibus, arcu crena profunda notato.
(Fem.) Capitulum with 14 reddish valves: upper latera quadrant-shaped, with the arched side deeply notched.
Mandibles with three teeth; maxillae narrow, bearing only four or five pair of spines.
MALES, two, lodged in cavities on the under sides of the scuta; pouch-formed, with four unequal, rudimentary valves: no mouth: cirri not prehensile.
Algoa Bay, South Africa. Attached to Sertularia and Plumularia. British Museum.[55]
[55] I am greatly indebted to Mr. Bowerbank for specimens of this extremely interesting species; also to Mr. Morris, to whom Mr. Bowerbank had given some of the original specimens.
FEMALE.
Capitulum oblong, with the upper portion much produced; valves, 14, thick, naked, closely locked together, irregularly clouded with pale crimson; the membrane connecting the valves is not furnished with spines. On most of the valves there are furrows and ridges diverging from the umbones, and the lines of growth are plainly marked: in the valves of the lower whorl, the umbones are slightly protuberant.
Scuta, convex, unusually thick, oblong, quadrilateral, with the occludent margin the longest; lateral margin slightly hollowed out. The umbo (and primordial valve) is situated at the uppermost point of the valve, and consequently the growth is exclusively downwards. On the under side (Pl. VI, figs. 1 b' and 1 c'), in about the middle of the valve, there is a pit (a) for the adductor scutorum muscle, the depth and distinctness of which varies a little; above the pit, and between it and the apex, there is a transverse, oblong, deeper depression (b), within which, the male is lodged. A small portion of the apex of the valve projects over the terga.
Terga, large, nearly equalling the scuta in area, flat and sub-triangular; the scutal margin is not quite straight. The apex of the valve is thick and solid, and must have projected freely for a length equalling one third of the occludent margin.
Carina, laterally broad, angularly bent; slightly widening from the apex to the base; internally, deeply concave. The position of the umbo varies, in young specimens it is seated at the uppermost point, and consequently in such there is no upward growth; in older specimens, from the junction and upward production of that part on each side of the valve, which I have called in fossil specimens the intra-parietes, the valve is added to above the umbo, but to a lesser degree than in S. vulgare. Slight ridges separate the roof from the parietes, and the parietes from the intra-parietes.
Rostrum, minute, narrow, widening a little from the apex downwards, inserted like a wedge between the umbones of the rostral latera, and hardly projecting above their upper margins, so as to be easily overlooked: internally concave.
Upper Latera (fig. 1 a), quadrant-shaped, with a deep square notch cut out of the arched margin, which notch receives the upper point of the carinal latera; the surface of the valve between the notch and the umbo is depressed.[56]
Rostral Latera, small, gradually widening from the umbo to the opposite end, which is obliquely rounded.
Infra-median Latera, approaching to diamond-shaped, placed obliquely to the longer axis of the capitulum; or the upper part may be described as spear-shaped.
[56] The only valve which I have seen at all like this, is a fossil specimen from the Upper Chalk of Scania; this is described in my memoir on the Fossil Lepadidae (Palaeontographical Society), under the name of Scalpellum solidulum (Tab. 1, fig. 8, e, f), and is perhaps erroneously there considered as a carinal latus.
Carinal Latera: these appear as if formed of two valves united together; the upper portion, widening as it ascends in a curved line, terminates in a rounded margin, which enters the deep notch in the upper latera; the other and lower portion is shorter, and terminates in a square margin abutting against the infra-median latera; the umbones of the carinal latera project beyond the line of the carina.
Direction of the Lines of Growth in the Valves.—This should always be carefully observed, on account of the great diversity there is in this respect between the different species, especially when the recent are compared with the older fossil species; moreover one of the chief characters between the genus Scalpellum and Pollicipes, depends on the direction of the lines of growth. In the scuta, terga, rostrum, and upper latera of the present species, the chief growth is downwards; in the carina, in mature specimens, it is both upwards and downwards; in the carinal latera, both upwards and towards the infra-median latera; in the infra-median latera chiefly upwards; and, lastly, in the rostral latera, towards the infra-median latera.
Peduncle, short, not half as long as the capitulum; calcareous scales imbricated as usual, tinged red, almost crescent-shaped, acuminated at both ends, of remarkable length, so that in each whorl there are only four scales: a full-sized scale equals in length one of the rostral latera. The tips of two scales, in one whorl, lie under the middle points of the carina and rostrum; and in the whorl, both above and below, a single much curved scale occupies this same medial position. The peduncle does not seem to have been attached in any definite position to the horny coralline, as is the case with S. vulgare.
Length of capitulum in the largest specimen .2 of an inch.
The Mouth is directed towards the ventral surface of the thorax. The Labrum is far removed from the adductor muscle, with the upper part forming an overhanging projection; I believe there are some very minute bead-like teeth on the crest. Palpi, small, narrow, thinly clothed with bristles.
Mandibles, with three teeth, of which the first is distant from the second; inferior angle not much acuminated, pectinated on both edges.
Maxillae, small, narrow, produced, without any notch, with two large upper spines, of which one is much thicker than the other; on the convex upper margin there are some minute tufts of very small hairs.
Outer Maxillae, with few bristles, arranged in a continuous line on the anterior surface; on the external surface there is a tuft of long bristles. Olfactory orifices situated laterally, forming two flattened, tubular projections.
Cirri.—First pair placed not far from the second; the three posterior pair not very long, with their segments elongated, not protuberant, bearing four pair of non-serrated spines, with a single short bristle between each pair; dorsal tufts small, with one spine longer than the others. First cirrus rather short, segments not very broad; second cirrus with the rami nearly equal in length, anterior ramus rather thicker than the posterior ramus, with three longitudinal rows of spines.
Caudal Appendages.—These are minute, rather broad, not half as long as the lower segments of the pedicels of the sixth cirrus, with four very long spines at the tip.
Penis.—There is no trace of a probosciformed penis in the four specimens examined; and as this organ is present in every ordinary cirripede, with the exception of Ibla Cumingii which we know to be exclusively female, so we may infer with some confidence that the form here described is female, although it is impossible in specimens once dried to demonstrate the absence of the vesiculae seminales and testes.
Affinities.—This is a very distinct species; it is, however, much more nearly related to S. rutilum, than to any other species; and next to this, to S. vulgare; from this latter species it chiefly differs in the large scales of the peduncle, in the scuta not being added to at their upper ends, and in the membrane covering and connecting the valves being spineless; but there is a greater difference in the trophi and in the cirri. The peduncle of S. ornatum presents some resemblance to that of the singular cretaceous genus, Loricula.
MALE.
All the specimens, as already stated, were dry, but in an excellent state of preservation, so that after having been soaked in spirits, they could be minutely examined. In the four which I opened, I found, in a transverse pouch on the under side of each scutum, a male lodged; in a fifth dead and bleached specimen, the cavities in the shell for the reception of the males, were present; and in a sixth young specimen, also dead, cavities were in process of formation. As compared with plants, the relation of the sexes in this species may be briefly given, by saying that it belongs to the class Diandria monogynia. I will first describe the males themselves, and then the cavities in the shell of the female. The males differ in every point of detail, from the complemental males of S. vulgare, but yet present so close a general resemblance, that a comparative description will be most convenient.
The general shape of the whole animal is rather more elongated, and I suspect flatter, but this latter point could not be positively ascertained in dry specimens. The entire length is greater, being in the largest specimen 13/400 (instead of at most 11/400), and the width, 7/400 of an inch. The orifice is not fimbriated; the four bristly points over the calcareous beads are absent. The whole outer integument is much thinner, owing evidently to its protected position, and is not covered by little bristles, but with an extremely high power, minute points arranged in transverse lines can be distinguished. The calcareous beads, or rudimentary valves, are thin and regularly oval. It is remarkable that in all the specimens, two on one side were smaller than the two on the other side,—the smaller beads being 16/6000, and the larger, 22/6000 of an inch in diameter; therefore more than twice the size of one of the beads in S. vulgare, which are only 9/6000 externally in diameter. From the position of the eye, close to one margin, near the upper end of the flattened animal, and from the manner in which the little limbs and spines lay between two of the beads at the opposite end, it was manifest that these latter, one large and one small, corresponded with the terga of the other cirripedes, and that the other two, near the eye, answered to the scuta. The valves being of unequal sizes on the right and left-hand sides of the animal, is probably connected with one side being pressed against the hard, shelly valve of the female; in the same way as the valves in certain Paecilasmas; are smaller and flatter on the side nearest to the crustacean to which they are attached. The eye, in being slightly notched on the upper and lower edge, shows signs of really consisting of two eyes, which I believe is always normally the case; it is rather larger, in the proportion of 13 to 11, being 13/12,000 of an inch in diameter, than in S. vulgare; and from the almost perfect transparency of the integuments, is far more conspicuous than in that species. Hence when the valves of the female are opened, the black little eye is the first part of the male which catches the attention. No vestige of a mouth could be discovered.
Thorax and Abdomen.—The thorax, as in S. vulgare, is highly extensible, and when stretched exhibits the same five transverse folds or articulations; when contracted, it is broader, so that even the truncated end of the abdomen is wider than the lower (properly anterior) end of the thorax in S. vulgare. Its thin outer integument is studded with excessively minute points in transverse rows. The four pair of limbs are longer than in S. vulgare, but the spines on them much shorter and thicker; each limb (including the first) supports three spines, of which one is seated on a notch low down on the outside, and is longer than the other two; of these two, the one on the same side with the notch, is a little longer than the other. The spines on the first and second pair of limbs are considerably shorter than those on the third pair, and these latter, are a little shorter than those on the fourth or posterior pair. Hence, the spines on the thoracic limbs, compared with those of S. vulgare, present considerable differences, both in their relative and absolute dimensions. The abdominal lobe is in proportion rather shorter; its end is less abruptly truncated, and supports a row of, I believe, six moderately long, and basally thick spines; these spines are not so long as those surmounting the fourth pair of limbs. On both lateral margins of the abdomen, rather on the ventral face, there is a row of, I believe, seven long spines, but it is very difficult to count the spines in specimens which have been once dried. I was able to distinguish that the two lower pair of spines on the ventral surface, are seated a little way one below and within the other, as in S. vulgare. The abdominal spines altogether form quite a brush, and there are certainly several more than in S. vulgare, and those on the two sides are much longer.
Antennae.—The disc is hoof-like, with the upper surface forming a straight line with the upper edge of the basal segment; the apex is pointed and clothed with some fine down; there is a single spine pointing backwards, which rises from the lower flat surface. The ultimate segment was hidden in laminae of cement; and I was not able to make out its structure. There is a single spine on the outer edge of the basal segment, in the usual position. The entire length of the limb, measured from the end of the disc to the further margin of the basal articulation, is 36/6000ths of an inch; measured to the inner margin, it is; 21/6000ths of an inch; the disc itself is 12/6000ths of an inch long; these measurements differ a little both absolutely ad proportionally, compared with those of the antennae of S. vulgare.
Cavities in the Scuta of the Female for the reception of the Males.—These extend nearly parallel to the tergal margin, transversely across the valves, for three fourths of their width; they are seated above the depression for the adductor muscle, and are more conspicuous than it; they are deep and well defined, and each exactly contains one male. The males are placed with their orifices in a little notch in the occludent margin, and their prehensile antennae at the further end. The distance to which the cavities extend across the valve, and their distance from the upper or tergal margin, varies a little, but chiefly in accordance with the age of the specimens; for the valve continues to increase in width, whilst the size of the cavity remains the same. The occludent margin of the scutum in the largest female, was .1 of an inch in length; of another, in which there was a fully developed cavity, .084; of a third, in which there was no cavity, only a slight concavity, with a preparatory impression, the length of the occludent margin was .062. The larger and smaller of these three valves, are drawn of their proper proportional sizes, in Pl. VI, figs. 1 b', 1 c'. The preparatory impression (fig. 1 c', b), consists of a narrow, not quite straight, extremely slight furrow, of slightly irregular width, bordered on each side by a very minute ridge, which is distinctly continuous with the inner edge of the occludent margin, both above and below the cavity. The furrow appears to have been formed by calcareous matter not having been deposited along this line, during the thickening or growth of the internal surface of the valve: I suspect, that it originates at a single period of growth, for I could see no signs of successively-formed transverse lines. I believe that it is strictly homologous with the fold, over which the complemental male is attached in S. vulgare, but carried, for a special purpose, much further across the valve and rectangularly inwards, for in structure and position both are identical. In comparing the internal views of the scuta in S. vulgare and S. ornatum (Pl. V, fig. 15 a', and Pl. VI, fig. 1 c'), it must be borne in mind, that the latter should be compared, as clearly shown by the lines of growth, with that portion alone of the scutum in S. vulgare, which lies under the curved ridge connecting the umbo and tergo-lateral angle. The deep cavity in which the male is lodged, is formed subsequently to the preparatory furrow, simply by the gradual thickening of the surrounding surface of the valve, more especially of a ridge just above the pit for the adductor muscle, and of another broad ridge just beneath the tergal margin. The deepest part of the cavity lies parallel to the tergal margin along the upper side, and here, in the older valves, the preparatory furrow can by care be distinctly traced. In conformity with the shape of the cavity, the orifice or notch in the occludent margin of the scutum, is situated at the point where the preparatory furrow sweeps round and enters. I believe that the cavity is lined by membrane, and that between the cavity and the body of the female, there is a complex membranous layer,—a pouch or bag being thus formed. An imaginary section of this pouch (with the thickness of all the parts extremely exaggerated and in a reversed position) is given in Pl. VI, fig. 1 d': a is the shell; x the cavity, converted, as I believe, into a pouch by, firstly, the delicate tunic (c) lining the sack of the female; secondly, a double layer (d) of corium; and, thirdly, by a special, rather thick membranous layer (b), which thinning out round the cavity coats only part of the under surface of the scutum. This latter membrane I have not seen in any other Cirripede, and I believe it is nothing but the tissue, here not calcified, which, in a calcified condition, ordinarily forms the valves. On this view, the males may be said to be lodged in pouches, formed in the thickness of the valves. |
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