A Monograph on the Sub-class Cirripedia · Charles Darwin
ORDER II.--ABDOMINALIA.
The other and lower (homologically anterior) end or lobe is broader, and contains a mass of cellular matter, which, from its close resemblance in appearance and position to similar matter within the male Alcippe, I have no doubt forms the contents of the testis. In one single specimen, I succeeded in isolating a vesicula seminalis of small size, containing perfectly distinct spermatozoa. Across the middle, between the two lobes, close under the outer integument, there is a broad layer of rather strong transverse muscular fasciæ. I did not observe any eye, the presence of which I should have expected from analogy. Internally there is no mouth, thorax, cirri, or other organs, excepting the testis and vesicula seminalis just mentioned, and an immensely elongated probosciformed penis, coiled up and filling the rest of the inside of the sack down to the testis, which latter occupies the whole anterior, and generally lower end of the animal. This penis is plainly articulated, and includes fine transversely-striated muscles: no doubt it can be protruded through the minute orifice, and voluntarily moved about.
Out of a male, 12/1000ths of an inch in length, I dissected a penis, which, when not stretched, measured 50/1000ths of an inch in length; when a portion was pulled between two needles, it could be stretched to apparently three times its former length, and I should think that this organ could be extended by the animal to, perhaps, even the 100/1000ths of an inch,--that is, to between eight and nine times its own entire length! The use of this enormously elongated penis obviously is, that the spermatozoa of these males, which are so extremely small in size, compared to the female, should all be conveyed within the sack, and none be lost. It should be borne in mind, that the whole male, including every part, is scarcely larger than a single ovum, of which sometimes sixty have to be impregnated by only two or three males. In a full-grown female, the distance from one of the attached males to the middle of the orifice leading into the sack, is about the 5/100ths of an inch, equal to the length of the coiled up, not-extended penis: the further distance from the orifice of the sack to an ovum lying at the bottom of the sack, would be almost 10/100ths of an inch, so that the spermatozoa have to pass a distance of 15/100ths of an inch from the testis of the male to the lower ova. I believe two thirds of this distance would be passed safely along the probosciformed penis.
The resemblance between the male of Cryptophialus and of Alcippe is truly surprising; and is the more wonderful, considering the great dissimilarity of their pupæ. Hardly any characters can be pointed out in which these males differ, excepting such as might have been thought of only specific value, namely the relative proportions of the different parts, and mere external shape. The peduncle growing a little after the metamorphosis, in the male of Alcippe, and the prolongation of its capitulum with the included oblique ligamentous fibres, are the greatest differences. Having fully remarked, under Alcippe, on the wonderfully rudimentary condition of these males, destitute as they are of so many parts and organs, I will here say nothing further on these singular creatures, destined to discharge their spermatozoa, die, and be succeeded by a fresh set of short-lived male successors. Order III.--APODA. _Cirripedia, with the carapace reduced to two separate threads, serving for attachment: body consisting of one cephalic, seven thoracic, and three abdominal segments, all destitute of cirri. Mouth suctorial, with the mandibles and maxillæ placed back to back, enclosed in a hood, formed by the union of the labrum and palpi. Metamorphoses unknown._
The characters above given fully justify, I think, the formation of this order; though it contains only one species, the _Proteolepas bivincta_. The mere external appearance (Pl. 25, fig. 7), so wonderfully different from that of every other cirripede, would by itself prompt to this same conclusion. At first sight the Proteolepas, if of fresh-water origin, might even have been mistaken for the larva of some insect, fastened by two threads to its prey. The entire absence of the three anterior segments of the head and therefore of the carapace, or, speaking strictly, the mere rudiment of these parts, forming an envelope to the two cement-ducts,--the absence of a stomach, rectum, and anus,--the entire absence of thoracic and abdominal appendages or cirri,--the absence of a probosciformed penis,--are all negative characters, which might ensue from degradation, so common with parasites; and which might, therefore, have been esteemed of not high classificatory value. But the suctorial mouth, with the palpi and labrum united into a hood, and with the mandibles and maxillæ reversed or turned back to back, so as to be utterly incapable of prehension, is a type of structure not hitherto met with, I believe, in any other animal, and cannot be explained away by degradation.
The formation of the ova within the segments of the body, a peculiarity confined to this one cirripede, evidently results from the non-development of the anterior part of the head, within which the ova are usually formed; but the compound structure of the vesicula seminalis is a peculiarity which cannot be thus explained. Proteolepas has no particular affinity to any other cirripede; it resembles, indeed, Cryptophialus in one important point, but only in one point, namely, in the number of the segments of its body. It is really beautiful to see how the homologies of the archetype cirripede, as deduced from the metamorphoses of other cirripedes, are plainly illustrated during the maturity of this degraded creature, and are demonstrated to be identical with those of the archetype Crustacean. I was at first inclined to rank Proteolepas in one division, and all other cirripedes in another division of equal value; but as it may be inferred from the characters of the prehensile antennæ, that the pupa did not differ much, if at all in any important character, from the pupæ of other cirripedes, I have thought the three orders, which I have instituted, would be the most natural arrangement.
As any one looking at the drawing given of Proteolepas, might very naturally feel inclined to protest against its being ranked as a cirripede, I must reurge the importance of the pupal antennæ being constituted on the common type, for from their structure, by the law of correlation, that of the whole pupa may be inferred; and even still more I must insist on the importance of the one great character of the antennæ being cemented to the surface of attachment by matter proceeding, as we shall see, in a modified state, from the great ovarian sack. The structure, also, of the mouth (to a certain extent), the segmentation of the body, though in appearance so peculiar, the hermaphrodite condition, the single penis, the absence of oviducts, all accord with, and taken together demonstrate, its cirripedial nature. PROTEOLEPAS BIVINCTA. Pl. 24, 25, figs. 1-7. _Hab._--Parasitic within the sack of the _Alepas cornuta_, from St. Vincent's, West Indies, Brit. Mus.
_General Appearance._[153]--The entire animal, as already remarked, curiously resembles, at the first glance, the larva of some insect. It is rounded, but somewhat compressed, and tapers gently towards the posterior end. It lies curved in an arc, the ventral surface being concave, and the dorsal convex, but a little flattened dorsally at the anterior and blunter end. Its length, if straightened, would rather exceed one fifth of an inch. The body consists of eleven segments, which, excepting the three terminal, are conspicuously plain. The first segment is surmounted by a rather small mouth, which any one would, assuredly, at first consider as the entire head, though he would in vain search here for eyes, antennæ, or other parts of the three anterior cephalic segments. On the dorsal surface, low down on the second segment of the body, two, quite flexible, thin, but strong, flattened threads arise, which terminate in a pair of prehensile antennæ, having the usual cirripedial structure. From the penultimate or disc segment of these antennæ, cement has been excreted, by which the antennæ are firmly cemented low down to the rostral end of the sack of the cirripede, the _Alepas cornuta_, on which it is parasitic: hence Proteolepas lies with its back downwards, and with its ventral concave surface fitting the convex body of the Alepas: its mouth lies under the middle of the soft prosoma of the latter cirripede, which I cannot doubt that it lacerates and sucks.
This compounded organ is singularly small compared with the palpi and labrum: it is narrow, being about 5/3000ths of an inch in width, but is produced upwards, so that a considerable length projects freely, and the rounded, properly external, margin can be traced down for a length of about 20/3000ths of an inch. In a lateral view of the mouth, the extreme tip of the mandibular organ could sometimes be seen just projecting out of the hood. The mandibular organ, when separated and carefully examined, presents the appearance, represented from a camera drawing, in (Pl. 24, fig. 2): we here see three groups of teeth; of these the lower set (_c_) consists of blunter teeth, placed more transversely, and easily separated from the others, and altogether clearly appears like a distinct organ. I do not feel nearly so sure regarding the other two sets; my first impression was strongly that they were distinct organs, closely united laterally together,--one (_a_) probably representing the mandibles, and formed into a single large tooth; the other (_b_) formed of three teeth, and probably representing the outer maxillæ; the first-mentioned set of teeth, which seemed to me to arise from between the other two sets, being the inner maxillæ.
If this view (and it must be remembered how excessively minute the parts are) be not, as I now suspect, correct, we must suppose that the outer maxillæ are aborted, and we have seen some tendency towards this in other cirripedes; the compounded organ being formed only of the mandibles (having on this view four teeth) and the inner maxillæ. As far as the mandibles are concerned, their existence, I may remark, is plainly shown by the presence of the palpi, which in all cases belong to and form part of the mandibles. The ventral surface of the mouth, immediately beneath the free portion of the compounded mandibular organ, consists of a triangular projection, but I could see no appearances to make me suppose that this part represented the outer maxillæ. The compound organ--in general shape, and in the oblique manner in which the front part is cut off and terminates in ligamentous apodemes, to which muscles are attached,--presents an unmistakable likeness to a mandible. It is hollow within, and muscles appear to extend some way up, perhaps to the transversely toothed portion, which represents, as I believe, the inner maxillæ: these two groups of teeth, anyhow, seemed to have some power of sliding over each other, and altered their positions during the course of dissection. On each side of the mouth, there is a muscle attached by its lower end to the basal edge of the labrum, and two others, one above the other, attached by their lower ends to about the middle of the labrum; these muscles, which are distinctly striated or voluntary, I infer, from analogy, run up to the ligamentous apodemes of the compound mandibles. There appeared to be other more delicate muscles attached to the basal articulation of the mouth on the ventral face, and these, I presume, would run to the supposed inner maxillæ.
The mouth in forming a prominence separated by a distinct articulation from the body, and in the union of the palpi and labrum (though here carried to excess), is constructed so far on the cirripedial type; but how are we to account for the extraordinary reversed position of the united mandibles and maxillæ, with their backs almost touching each other, and their toothed edges twisted round so as to face outwards in a manner unexampled, I believe, in any other articulate animal? It might, perhaps, be at first suspected, that the compounded mandible had not really been twisted round, but that the teeth had been abnormally developed on the outer convex margin: this view, however, certainly cannot be admitted, for the properly outer convex margin can be traced running far down the mouth, in a manner utterly inexplicable, if this were really the inner side; and equally inexplicable on this view would be the position of the ligamentous apodemes.
Hence I cannot doubt that this compounded mandibular organ has really rotated on its axis; and if the course of development could be followed, I suspect that the twisting would be seen to be effected as follows: we know in all cirripedes that the outer and inner maxillæ, and to a certain extent the mandibles, instead of facing each other, are directed towards the labrum; they therefore have already been twisted round a quarter of a circle, as may be seen in the diagram (Pl. 24, fig. 4), copied from the mouth of Ibla. Now let us drive inwards the front of the mouth, along a narrow medial line; these organs would then (fig. 5) be compelled to turn round a quarter of a circle more, and so face directly outwards. In this process, the integument between the lower and outer part of the mandible and the base of the palpus, which normally are in close contact, would have to be greatly stretched. By a movement of this order, the mandibles would come to stand posteriorly or exteriorly to the other gnathites; and as far as I could make out (previously to my having any theory) the large single toothed portion of the compound organ which most resembles a mandible, did really stand outside the other toothed portion.
With respect to the action of this singularly constructed mouth; if its ventral and oblique surface were applied to any yielding object, as the adjoining soft prosoma of the Alepas, the compound mandibles would be worked within an absolutely closed chamber. The action of these mandibles would be to make a transverse slit, and subsequently to serve as a grapnel to keep the mouth closely adpressed to its prey: the other teeth might act in keeping the wound open. When the mouth was thus closely adpressed over a wound, the great power of shortening the whole body which the animal possesses (the œsophagus being closed), would, by the subsequent action of the elasticity of the outer membrane, almost certainly create suction, and thus cause the nutritious juices of the Alepas to flow into the body of the parasite. Hence I have called the mouth suctorial.
_Body._--This, as already stated, consists of eleven segments, of which the three posterior (abdominal) smaller segments can hardly be distinguished, without dissection, as separate from each other. The body is mainly occupied by a vast ovarian sack (_e_, _e_, fig. 7), filled by innumerable ova: and the three posterior segments by small testes and their vesiculæ seminales (_i_): but I shall return to the internal anatomy. The outer membrane, lined by delicate corium, is thin, transparent, elastic, and covered by groups of excessively minute blunt little points. The segments can be plainly distinguished by their outlines, especially on the ventral surface; but they are rendered unmistakably distinct by the attachment of the muscles; they can also be perceived when the external membrane is perfectly cleaned, by yellowish lines. The muscular system is highly symmetrical and simple: along all eleven segments, there is a narrow, medial, ventral and dorsal clear space; on both sides of which space there is a band of longitudinal muscles, which, though encroaching on the two sides, and rather largely on the dorso-lateral sides, may be called the ventral and dorsal muscles. These muscles are striæ-less, which is the case with the homologous posterior thoracic muscles in some other cirripedes: on the dorsal surface (lower surface in fig. 7) they are more spread out, and consist, on each side of the medial line, of four ribbons: this seems to be the case on the ventral side, but the ribbons are here much more confluent: in the seventh and eighth segments, the ribbons become broader; but in the ninth, tenth, and eleventh, or three posterior segments, they become much narrower, and some of the fasciæ disappear, so that these muscles can hardly be seen from the outside. Each separate ribbon expands a little at its two ends, which are attached to the articulations separating the successive segments: I carefully observed that they did not pass over at either end to the adjoining segments: hence their action must be either simply to shorten and arch each segment separately; or when acting together, to shorten the whole body, or perhaps the ventral or dorsal surface by itself.
In the first segment, and in the three posterior segments, these longitudinal muscles alone occur; but on the seven segments, from the second to the eighth inclusive, there are other oblique latero-ventral muscles. These muscles lie within the longitudinal muscles, and adhere pretty firmly to the coat (_e_, _e_, fig. 7) of the great ovarian sack. At their ventral extremities they are attached, near the anterior margin of each segment, beneath the point of attachment of the longitudinal fasciæ, and thence they run posteriorly in an oblique line to the anterior margin of the next succeeding segment, where they are attached: so that these muscles run obliquely from segment to segment. The first of these oblique muscles, lying chiefly within the second segment of the body, is thinner and longer than the others: those within the third and fourth segments are short: those within the fifth and succeeding segments extend, at their dorsal (or lower in fig. 7) extremities, as far as the outer dorsal longitudinal fasciæ: those within the seventh segment are broad and short, and cross the longitudinal muscles at only a small angle.
In the eighth segment, there is an oblique lateral muscle, like that in the seventh segment, running from the ventral surface towards the dorsal surface; but there is in addition a second oblique lateral muscle, rising from the dorsal surface, and running towards the ventral surface. This muscle does not occur in the other segments, but in the fourth segment, at the dorsal end of the oblique latero-ventral muscle, there may be seen a small branch of fibres, at right angles, which seems to represent a muscle homologous with that just mentioned in the eighth segment: obscure traces, moreover, of similar fibres, can be detected in some of the other segments: had these oblique latero-dorsal muscles been as fully developed in the seven anterior segments of the body, as on the eighth segment, the whole muscular system would have been perfectly symmetrical. The oblique latero-ventral muscle in the sixth segment is distinctly striated transversely; but this is not the case with most of the other muscles, if with any of them; I cannot account for this difference. The muscles of the gnathites are the only other voluntary muscles in the animal's body.
_Homologies of the Body._--It will hereafter be, I think, clearly shown, that when the shell and integuments of the pupa of Proteolepas are shed, no carapace or general covering for the body is formed; the three anterior segments of the head, the backward prolongation of which (as has been elsewhere explained) certainly forms the carapace of ordinary cirripedes, being here almost absolutely aborted. In every cirripede the mouth is formed of three pairs of gnathites, which, no one will doubt, rise from the fourth, fifth, and sixth segments of the head: here in Proteolepas, the mouth, even on the view of the mandibular organ on each side being compounded of only two gnathites, sufficiently resembles the ordinary cirripedial type to make it very probable, that if examined in the earliest stage of its development, three pairs of gnathites would be discovered. In accordance with this conclusion, the segment succeeding the mouth (_i. e._, the first segment of the body in fig. 7) homologically is the seventh, or last cephalic segment. The succeeding seven segments, of course, are the seven thoracic segments, and the three posterior segments are abdominal; the latter are not developed in ordinary cirripedes when mature, but are present during their pupal condition.
Now this conclusion, which is, in fact, deduced from what we know of the front part of the head in other cirripedes, both larval and mature, appears to me most satisfactorily confirmed by the differences in the muscular system of the segments in Proteolepas. In no other way, I believe, can it be explained, why the last cephalic segment and the three abdominal segments should differ from the seven thoracic segments, in the entire absence of the oblique lateral muscles. The abdominal segments, moreover, differ a little in shape, in the indistinctness of their articulation, in the thinness of the longitudinal muscles, and even in their contents. With respect to the two threads enclosing the cement-ducts, which spring from the second segment of the body (or first of the thorax), and which terminate within the prehensile antennæ of the pupa, we shall hereafter see that their apparently most anomalous position, and even the flattened shape of the dorsal surface of the two anterior segments of the body, all accord perfectly with the homologies just given.
_Alimentary Canal._--The œsophagus is thin, and for a cirripede short, for it extends only half-way down the first segment (_i. e._ last cephalic) of the body; the lower end, which is slightly dilated, nearly touches the anterior end of the great ovarian sack. At its upper end, it is surrounded by delicate, striæ-less constrictor muscles; and there are others radiating outwards, evidently serving to open it: the lower part of the œsophagus, differently from other cirripedes, is destitute of muscles, and is only coated by a thin layer of corium, which would serve to produce a new œsophagus at each exuviation. Strange as the fact may be, I am prepared to assert that there is no stomach, rectum, or anus. As I was able to trace so distinctly the œsophagus, and likewise the generally far smaller orifice and ducts of the male generative organs, I consider it quite impossible that I could have missed the stomach. The rectum and anus are absent in Alcippe: and the absence of a stomach is here in some degree the less surprising, as the structure of the mouth shows that Proteolepas must live on the already elaborated fluids of the Alepas, to which, being a cirripede, it is allied. It is of some importance to observe, that the œsophagus is fitted with muscles simply for shutting and opening it, the wave-like swallowing action of which other cirripedes are capable, being, apparently, here impossible; but the contraction of the body and its subsequent expansion, the œsophagus being opened, would allow the blood of its prey to flow inwards.
The nervous system must be much atrophied, for I could not detect it, and the small size of the animal is not sufficient to account for this: I wish I could have seen this system, for then I should almost certainly have beheld an articulate animal without a trace of a supra-œsophageal ganglion. There is no eye, but such could hardly be expected, as the anterior cephalic segments are aborted. There are no branchiæ. I may state that within the abdomen, along the dorsal surface, there was either a lacuna or a delicate vessel, apparently of a circulatory nature, of considerable diameter, which, near the extreme posterior end of the body, gave out branches. _Female Reproductive Organs._--The eight anterior segments of the body, with the exception of a small space at the two ends, are occupied by an immense (_e_, _e_), opaque, ovarian sack. The tissue forming it is delicate, and presents a peculiar cellular aspect: it is slightly attached to the corium on the ventral surface of the body, and to the oblique latero-ventral muscles. Internally, at the anterior end, it is thickly coated by cellular matter, the cells varying from 4/6000ths to less than 1/6000th of an inch in diameter, becoming in parts confluent, and the whole forming a dark orange-coloured mass. In the more central parts of the sack this cellular matter became aggregated into little pellets, which, in proceeding towards the posterior end of the sack, gradually increased in size, from about (4 to 6)/1000ths of an inch in diameter, and at last appeared as almost mature and perfect ova of a broadly oval figure. Their size, as we see, is small, and their number almost infinite. I carefully examined all round this ovarian sack, and could detect no oviducts; nor from analogy could they be expected: I have no doubt that the ova burst forth by the rupture, probably, of the posterior end of the sack and of the overlying corium; and that they accumulate beneath the external membrane of the body, until this is moulted, the rupture beneath being in the meantime healed, when they are freed, or perhaps temporarily protected in the old moulted envelope of the body.
Text from Project Gutenberg, public domain in the United States.