A Monograph on the Sub-class Cirripedia · Charles Darwin
TABLE OF CONTENTS.
_Larva, First Stage._ The larvæ in this stage are known, amongst the Balanidæ, in Balanus, Pyrgoma, Coronula, Platylepas, and Chthamalus; and these genera include all the principal forms. Amongst the Verrucidæ they are known in its one genus, Verruca. Amongst the Lepadidæ, in Scalpellum, Ibla, Alcippe, Lepas, Conchoderma, &c.; and in all these genera the larvæ present no important difference--hardly any difference which could be viewed as generic, were these larvæ independent animals,--as may be inferred, chiefly, from Mr. C. S. Bate's descriptions.[51] The abstract given in my former volume was not accompanied by any illustrations, and I have consequently here given (Pl. 29, fig. 8), a view of the larva, in the first stage before moulting, of _Scalpellum vulgare_: the natatory legs are not drawn with accuracy, only the relative position of the several organs having been carefully attended to. I have also had copied from Mr. Bate's memoir, a figure of the larva (Pl. 29, fig. 9) of _Balanus balanoides_, in its first stage, _before_ moulting, with its ventral surface exhibited; and another figure (with a few trifling alterations made after examining specimens most kindly sent me by Mr. Bate) of the larva of _Chthamalus stellatus_ (fig. 10), in its first stage, but _after moulting once_. It should be observed that Mr. Bate has given a drawing of the larva of this latter cirripede, in the first stage, _before moulting_; and it does not differ essentially from that just referred to (fig. 9), of _B. balanoides_, but is rather more fully developed. These drawings suffice to show the character of the larvæ in the first stage, both before and after the first moult, and even after the second moult, throughout the Order of Thoracica. The larvæ sometimes undergo their first moult within the sack of their parent, as I have been informed by Mr. Bate, and as I have observed in Coronula.
In this same larva of _Scalpellum vulgare_, within the great lateral horns just alluded to (fig. 8, _c_), filiform organs, supporting rows of spines, could be distinguished; and these appeared to me to be antennæ. These horns or cases resemble in structure the smaller pair just described; they arise from the ventral surface, and can hardly, therefore, be considered as prolongations of the carapace. After the first moult (fig. 10, _c_) they are seen to have increased much in length: in some cases they are of considerable length before the first moult, as in Lepas: in the Balanidæ they seem to be generally shorter than in the Lepadidæ; but in _Balanus galeatus_ I found them one third of the entire length of the animal. Whilst within the egg, these horns are adpressed laterally to the body, and so point posteriorly; afterwards they project rectangularly from the sides, or, as in _Scalpellum vulgare_, are directed somewhat anteriorly. As in the larvæ of all ordinary Crustaceans, as yet known, the antennæ are amongst the earliest developed organs; and as the first pair of natatory legs (Pl. 29, figs.
8-10, _e_) in these Cirripedial larvæ, might so very naturally be thought to be antennæ (as has been remarked to me by Mr. Dana), both from their structure and from their position a little anteriorly to the mouth, I am well aware that to prove my view correct, namely, that these horns are the second pair of antennæ in process of formation, it is not sufficient merely to have seen organs resembling antennæ within them; nor is it sufficient to advance the strictly analogical fact of the first-mentioned pair of antennæ, which in Scalpellum indisputably appear in their earliest condition within an envelope or horn. Further evidence is required, and this is presented in Cryptophialus, in which the lateral horns of the egg-like larva, in its first stage (Pl. 24, fig. 16), can be actually followed step by step until, in the second stage (fig. 17), just before passing into the pupa, the horns are seen to have become larger and more nearly approximated to each other on the ventral surface; and whilst in this condition, I several times dissected out the prehensile antennæ of the future pupa with _every character perfectly recognisable_. Hence I cannot doubt that in the larvæ of Cirripedes the law of development is, that in their very earliest condition, the small first pair of antennæ are enclosed in cases; and that the large second pair remains thus enveloped until the pupal stage. This conclusion, we shall immediately see, is in harmony with the late development of the succeeding appendages or organs of the mouth, which certainly do not appear in the first larval stage, and are not known to appear even till after the final metamorphosis.[53]
We come, now, to the three pairs of natatory legs: the first (Pl. 29, figs. 8-10, _e_) has throughout the order only one ramus, whereas the two succeeding pairs (_f_, _g_) are biramous. I must here remark that the straight and strong, and the curved plumose spines, with which these limbs, after the first moult, become furnished, now appear to me as more probably prehensile, rather than masticatory as I imagined in my former volume. That these spines are important organs to the larvæ I do not doubt. With regard to the homologies of these three pairs of limbs, my first impression was that they were the mandibles and the two pairs of maxillæ in their earliest condition; but I consider this view as quite untenable, for several reasons; viz., the wide interval between their bases and the mouth itself,--the somewhat variable position of the mouth with respect to the legs,--and the position which the latter occupy in the _second_ larval stage.[54] A far more tenable view is that these three pairs of legs are the three pairs of maxillipeds, in their earliest condition, in accordance with the view of M. Joly[55] on the nature of the three very similar pairs of natatory legs in the larva of Caridina, a macrourous Crustacean. But, in Cirripedes, the three pairs of natatory legs, in the larva in the first stage, are apparently the very same as the first three pairs, in the larva in the second stage, and in the pupa. And in the pupa the first three pairs, which certainly correspond with the first three pairs of cirri in the mature animal, seem to me, for reasons presently to be assigned, to be the second, third, and fourth thoracic limbs. Hence I am led to the conclusion that the first pair of legs in the larva in the first stage, are homologically the second thoracic (answering to the third pair of maxillipeds in the higher Crustaceans), and that the two succeeding pairs are the third and fourth thoracic limbs; to be succeeded, in the _pupal_ stage, by the fifth, sixth, and seventh thoracic appendages.
Lastly, behind the natatory legs, on the ventral surface, (Pl. 29, figs. 8, 9, _i_), the body is much produced, and terminates in a horny fork, which, after the first moult (fig. 10, _i_), becomes much elongated. Anteriorly to this fork, on the ventral surface, there is another fork (_l_), and again above this I could distinguish, in _Chthamalus stellatus_, after the first moult, another fork (_m_), or at least a pair of short thick spines. From the structure of the forked abdomen in the known larvæ of the Podophthalmia, I presume that this portion of the body is the abdomen of the young Cirripede, but it is not at all plainly articulated. After the first moult, the posterior end of the carapace (_h_), which is always pointed, becomes much elongated and serrated on both sides;[56] reminding one of the structure of the carapace of the so-called Zoea, or larva of certain Podophthalmia. Situated under this posterior prolongation of the carapace, there is a swelling (_n_, with long hairs on both sides), which apparently lies on the dorsal surface of the spinose and forked abdomen; here, when the larva is compressed, the cellular and oily contents of the body burst forth; and I suspect that this swelling is the anus, for it is known from the researches of Rathke,[57] that the anus in the higher Crustaceans opens during the earliest periods dorsally.
From the presence of eyes and of two pairs of antennæ in the larva, during its earlier stages, the front of the head consists, in accordance with all analogy, of three segments; the mouth, likewise, from being formed of three gnathites (which can be detected by dissection in the pupal state), consists, also in accordance with all analogy, of three segments, making altogether six segments--on the nature of which I apprehend no objection will be raised. In two out of the three orders into which Cirripedes may be divided, the mouth is succeeded, in the adult animal, by eleven most distinct segments; of which the first (_i. e._ the seventh cephalic) differs from the succeeding seven thoracic segments; and these seven again differ from the three abdominal and terminal segments. Hence it must be admitted that, as far as the cephalo-thorax of the archetype Cirripede is concerned, it consists, like that of the archetype Crustacean, of fourteen segments, of which eight succeed the first-named six that form the mouth and front of the head; and that, with the three abdominal segments, there are altogether seventeen segments.
In the order Thoracica, however, which includes all common Cirripedes, both in the pupa and in the mature animal, only six thoracic segments with their appendages, succeed the mouth, two having been lost; and the question arises which are these two, whether the seventh and eighth, or the thirteenth and fourteenth (_i. e._ the two terminal thoracic) segments; for there is no reason to suspect any other segments of having disappeared. In my former volume, I inferred, without sufficiently entering into my reasons, that it was the seventh and eighth, _i. e._ the last cephalic and first thoracic segments, which had disappeared; but I now find that Mr. Dana[59] believes that, in ordinary Crustaceans, the abortion of the segments with their appendages almost always takes place at the posterior end of the cephalo-thorax. Nevertheless, after due deliberation and fresh examination of the pupa, I must retain my former opinion, that it is the last cephalic and first thoracic segments which have either coalesced with the others, or wholly disappeared. In the pupa, the mouth, although functionless, has its place most plainly marked by being slightly prominent, and by the presence of a sort of labrum and of a shrivelled œsophagus, round which latter the gnathites and the new œsophagus of the future young cirripede are in process of formation.
Now between the mouth of the pupa and the first pair of natatory legs, there is a space of membrane, equalling, when stretched out, the three succeeding thoracic segments in length and breadth: this interspace, I conceive, must have some homological signification; here then we have at least an appearance of the abortion of appendages; whereas, at the posterior end of the cephalo-thorax, no such appearance is presented. Moreover this interspace of membrane is divided nearly in the middle by a most conspicuous fold, which, on the view here adopted, would mark the separation of the seventh (cephalic) from the eighth (thoracic) segment; and the interspace and fold are thus simply explained. Lastly, I have shown, in the Introduction (p. 18), that the first and five succeeding pairs of cirri of the mature Cirripede present certain small, but significant, resemblances in structure and in the origin of their nerves, with the outer pair of maxillipeds and with the five pairs of ambulatory legs in the Podophthalmia; which resemblances are all futile, if the cirri belong to the 7th, 8th, 9th, 10th, 11th, and 12th segments of the cephalo-thorax, or those immediately succeeding the mouth; but are full of meaning, if the six pairs of cirri belong, as I believe, to the 9th, 10th, 11th, 12th, 13th, and 14th segments, or the six posterior segments of the cephalo-thorax.
The surface of the carapace in _L. australis_ is lined, as represented in fig. 4: the colour of this species when alive was blue:[60] in _L. fascicularis_ the surface is punctured: in _L. pectinata_ it is marked with curious points of various shapes, often star-shaped, in parts reticulated, and confluent along the dorsal margin, and in parts lined: in _B. balanoides_ it is very obscurely punctured, and in _B. Hameri_ the punctures pass into lines. The whole of what is externally visible consists of the carapace, for this is produced not only backwards, so as to enclose the thorax and abdomen with their appendages, but also forwards, so as to overhang the whole front of the animal; and the prehensile antennæ, in Lepas, Ibla, Balanus, and probably in all the genera, can be retracted within its lower edge. The protection afforded by the carapace to the antennæ is aided by two crests (Pl. 30, fig. 7, _c_) parallel to this lower edge. The whole sternal surface is very narrow (fig. 4), and is likewise protected by the carapace; that is, when the two sides are drawn together by the adductor muscle. The shell, however, when thus drawn together, gapes a little at the two ends, at least in the case of _Lepas australis_.
The adductor muscle, if introduced in fig. 4, would have crossed close anteriorly to the basal margin of the mouth; and in fig. 2, its end on the near side would have been attached under the dark cæca, which enter the upper end of the stomach. The adductor is shaped almost like an hour-glass, and so differs from this muscle in the mature Lepas, in which it is of the same thickness throughout. I may here add that the pupa of _Lepas australis_ could swim very rapidly, and often on one side in a circle; it could walk by the aid of its antennæ, but often fell over; being thus locomotive, and, as we shall immediately see, well provided with senses, it cannot be considered as very lowly organised. [60] I took this species alive in the Southern Atlantic Ocean; and, mistaking it for an independent Crustacean, was much perplexed where to class it. I had overlooked these specimens when publishing my former volume.
_Acoustic Organs._--Commencing at the anterior end, two small elongated orifices, 10/6000th of an inch in diameter, (_e_, fig. 4, Pl. 30), may be seen; these lead, as described in my former volume, into a sack, with a bag suspended in it, which is provided with a large nerve, and which I believe to be the acoustic vesicle. These orifices occur in the carapace, either in the same position, or a little more posteriorly, in the pupæ of all Cirripedes. In _Balanus balanoides_ they are minute, being only 2/6000th in diameter, but are surrounded with a border: in _Conchoderma virgata_ they are also surrounded by a border: in _Lepas pectinata_, the orifices are 3/6000th of an inch in diameter, and are very singular from being seated on rounded prominences, causing the carapace to have two short, blunt horns in front. In _Lepas australis_, and I believe in the other species, the corium round the acoustic orifices is darkly coloured; and these coloured marks can be distinguished for some little time on the peduncle of the young Cirripede, after the metamorphosis, and after the entire organ, together with the whole pupal carapace and eyes, has been moulted. Knowing the connection in the higher Crustacea, of the acoustic organs and the antennæ, and seeing the very backward position (figs. 2 and 4) of the one great pair of antennæ, I have always imagined that these orifices probably marked the normal position of the anterior pair of antennæ, which, since the earlier larval stages, have disappeared. And I now find[61] that Schödler affirms, that in most, if not in all Daphnidæ, there is a black spot in front of the eye, which is connected with an opening in the basal portion of the anterior antennæ, and he concludes that it is an organ of hearing.
In the Southern Atlantic I took some specimens of the pupa of _Lepas australis_, not yet attached, and therefore with the muscles of the antennæ, not having suffered any of that absorption, which they undergo, as soon as the pupa is permanently cemented to some floating object. In my former volume I noticed a pair of strong muscles, attached to the tips of the middle forks (Pl. 30, fig. 7) of the apodemes, and I now find two pairs attached to the bases of the two outer forks, and directed dorso-anteriorly; and two other pairs, also attached to their bases, but directed dorso-posteriorly, so that altogether there are five pairs of muscles attached to the apodemes; their chief function, I should think, was to draw the antennæ posteriorly and upwards within the carapace; but as the apodemes cannot be moved without the great compound eyes being likewise moved, the muscles probably serve a double purpose. When the pupæ were alive, I noticed that their eyes were constantly kept in a state of vibratory movement. Flexor and extensor muscles are attached at one end to the posterior margin of the basal segment, and at the other end to the second or main segment; other powerful muscles attached to this latter segment, are prolonged by ligaments into the disc. In Cryptophialus I observed that the disc-segment had a movement almost like that of the wrist. Whether any muscles enter the small terminal segment, I know not.
The drawing in Pl. 30, fig. 8, of part of the second segment, of the third or disc segment, and of the fourth or ultimate segment, in _Lepas australis_, is, I think, very accurate. The second segment articulates on the upper or dorsal surface of the disc, and has the articulation on one side constricted and formed of thin and flexible membrane; the little terminal segment, which is turned outwards at right angles, also, articulates on the disc. That the disc forms a true segment is shown clearly in Cryptophialus (Pl. 24, fig. 18), where the articulation with the second segment is not in such close contact. The disc is either circular, as in Lepas, or hoof-shaped, as in Ibla: in _B. balanoides_ the disc is rather hollowed out on the inner side. It has the power of adhering even to so smooth a substance as glass, placed vertically. It is surrounded by a rim of transparent membrane. On the hinder margin some spines arise from the central and more opaque part: in _Lepas australis_, there are no less than seven of these spines (fig. 8): in _Conchoderma virgata_ there are only four, in _Scalpellum_ and _Ibla_ only one. When the disc is placed on the surface of attachment, these spines lie parallel to it.
The middle part of the disc is, almost always, nearly opaque; and in tracing the cement-ducts from within the body of the pupa, or of the young Cirripede, I in many cases traced them as far as this point, but here lost them. From this same obscure central part of the disc, in most, if not in all species of the Lepadidæ, spokes radiate, which sometimes are branched, and are not regular, not always even resembling each other on the opposite sides of the same individual. Round the proper membranous border of the disc, a second one may be observed (fig. 8, _p_), which differs in shape and extent in different specimens: under favorable circumstances, and very high powers, it may be seen to have a reticulated structure, and to be of a very pale brown colour; towards the exterior margin, the reticulations become finer, and are blended together and lost; on the inner margin, the substance forming this membrane may be seen to come out of the spokes. This substance is the cement, which has the power of adhering to whatever substance it grows against; and thus the disc of each antenna becomes cemented down, and soon both the antennæ are surrounded by a common border of cement, which gradually increases, after the metamorphosis, in extent. Occasionally the cement forms little projections, like short spines, on the edges opposite to the orifices of the spokes.
The small terminal segment usually bears on its truncated extremity six spines, some of which are occasionally hooked; in Scalpellum, two spines, rather longer than the others, are borne on a step some way down on the inner side of this segment; but in Lepas, two spines (fig. 8), very much longer than the others, arise from the outer corner of the extremity. These two are very different from the other four borne by this segment, or indeed any other spines on the body; for they are quite flexible, and are furnished with a double row of very long, straight, excessively fine hairs, which seem to be articulated on them--the whole presenting a very beautiful appearance. These spines are of considerable length, and in _Conchoderma virgata_ they even equal in length the whole antenna. I can hardly doubt that these beautiful, plumose, flexible spines, into the thick bases of which the coloured corium could sometimes be seen to enter, serve as feelers. Owing to the facts immediately to be mentioned, I erroneously stated, in my former volume, that there were three long spines.
In three species of Lepas, in _Dichelaspis Warwickii_, and in _Scalpellum Peronii_, after having torn the lately-cemented antennæ from the surface of attachment, I observed proceeding from the end of the terminal segment, from between the above two groups of spines, what appeared to be a long narrow ribbon with its end torn off; and which, in the case of _Lepas australis_, I fancied was one of the plumose, long, flexible spines ripped open. But now that I have examined some of the pupæ of this species before their attachment, I find (as represented in Pl. 30, fig. 8, _v′_) a flattened tube, ending in a blunt point, and having a peculiar ringed structure. I have noticed similar appendages to the antennæ of specimens just attached of _Lepas anatifera_. In the Dichelaspis and the Scalpellum, the tube was very long, and seemed, from its torn appearance, to have been firmly attached to the supporting surface. In both these cases, the tube came out from within another slightly larger tube, which had been broken off close to the extremity of the terminal segment of the antenna. In the case of the _Lepas anatifera_, the tube expanded a little after leaving the antenna.
In the Dichelaspis, it had exactly the same diameter as the cement-duct, which could be clearly distinguished within the two lower segments. From these several facts, and from the peculiar appearance of the tube itself, I believe it to be a tube of cement-tissue which thus, sometimes even before the pupa is attached, independently grows outwards. That the cement-tissue can grow outwards and assume definite forms, we know from the singular case of _Lepas fascicularis_, in which the cement proceeding from several apertures, forms a vesicular float round the peduncle of not only a single individual, but often of a group of specimens: we shall presently find a somewhat analogous fact in the case of Coronula. It is possible that this tube, proceeding from the extremity of the antenna, may be the channel through which cement continues to be poured forth during the continued growth of the above Cirripedes; and the manner in which this is effected, considering how firmly the end of the peduncle is cemented down, has always appeared to me a difficulty.
In those pupæ of _Lepas australis_, which I caught swimming about unattached, it is surprising that the disc should have been edged with cement, and that a tube, similarly formed, should have grown out of the ultimate segment: it shows, I presume, that the cement-tissue will grow out, whether or no the pupa has succeeded in finding a proper object for attachment. Lastly, I have felt some surprise, in two or three instances in observing some dark purple pigment-cells, like those in the corium, within the terminal tube of cement; and likewise within the spokes of cement in the disc: this is the only fact which causes me the least doubt, whether I have rightly determined the nature of the terminal tube, as wholly formed of cement tissue; or whether it may not be covered by an outer integument, itself lined by true corium, coloured purple. Finally, I may add, that, excepting in small details, the prehensile antennæ present no difference throughout the Order: I have minutely examined them in several genera of the Lepadidæ; and in the Balanidæ, I have seen them in Coronula, and in several species of Balanus. In _B. balanoides_ I have examined them carefully; they are smaller and thicker than in Lepas, with the second or main segment bowed outwards, carrying its usual single spine; with the disc excised on its inner margin and apparently without the spoke-like vessels for the cement; and with the ultimate segment proportionably longer, and carrying, I believe, six spines, of which two appeared to be longer and more flexible than the other four shorter and somewhat hooked spines. In _Coronula balænaris_, also, the terminal segment is, proportionably to the others, of large size. Not only throughout the order, but throughout the whole Class, the antennæ are singularly uniform in structure, as will be seen, when the last two orders are described.
Ideas discussed in this chapter
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