A Monograph on the Sub-class Cirripedia · Charles Darwin
TABLE OF CONTENTS.
_Tubicinella._--The cementing apparatus is here less symmetrical; but this, I believe, is chiefly owing to the basal membrane being formed of successively larger discs (not slips) of membrane, thrown down not quite concentrically one over the other; each new disc of membrane seems to cover the last-formed cement-glands and ducts; and there are as many ducts and glands as there are discs of membrane, all adhering together. In one specimen, it appeared that normally there were four sets of cement-ducts, as in the allied genera of Coronula and Platylepas; but in other specimens, the ducts were distributed very irregularly. In one case the two cement-trunks extended parallel and close together, one of them terminating long before the other. I have given a figure (Pl. 28, fig. 3) of three of the cement-glands, removed from the basal membrane, together with their ducts. The cement-trunk (_f f_) seemed to be a little enlarged, and to be crossed by septa, as it entered the glands (_h_), but I could not make out this structure clearly enough to be represented. Whilst young, the cement-glands stand some little way apart from each other; and in the figure given of some of the last-formed glands, they are hardly separate enough.
Each gland gives out obliquely, on one side, a cement-duct (_c_) which I traced in several cases to the margin of one of the discs of basal membrane, where cement issued from it; and on the opposite side, a tapering spur (_b_), varying in length, which may be called, and I believe really is, a rudimentary duct. Of these spurs we shall meet many instances in other genera. The duct (_c_) and the spur (_b_), close to where they entered the gland, in some specimens gave off, at about right angles, short blunt points, or rudimentary branches. This duct and spur correspond, I believe, with the two ducts in Coronula; but besides these, a duct (_a_) is given off from one end of the gland, from the surface opposite to that at which the cement-trunk enters. This duct (_a_) is very singular, from always forming a loop (_a′_), with two spurs projecting from it: these two spurs occasionally spring from a common point: I have seen nothing like this structure in any other Cirripede. This duct (_a_) runs, like the duct (_c_), to the margin of its own disc of basal membrane, where it debouches.
Besides these ducts, in the best specimen which I examined, there were two other sets of ducts, which were slightly zig-zag, and at each angular bend, a mere knob or point, or at most a short branch, was given off; but this branch seemed never to run to the margin of the basal membrane or to give out cement; whereas the main branch did give out cement. I was not able to trace these ducts to their glands. In these zig-zag ducts, and in the rudimentary points sometimes observed at the base of the duct (_c_), and likewise at the base of the spur (_b_), we see the first indication of that tendency to bifurcation, so strongly characteristic of the cement-ducts in all the genera, excepting those already described, which are allied to Coronula.
_Chelonobia patula._--The cementing apparatus is here chiefly remarkable for the thinness and straightness of the main trunk, (_f f_, Pl. 28, fig. 2), and from the great distance at which the glands stand apart; had another gland been drawn, it would, on the scale here used, have stood exactly under the two upper, (_c′ c′_) in fig. 1 _c_. We here see that the trunk (_f_), before entering the gland (_h_), has an enlarged portion (_g_); this, I suspect, is a very general structure. Each gland gives out, on opposite sides, two ducts (_a a_, _b b_), larger even than the main trunk; and these ducts bifurcate repeatedly, and inosculate. By this inosculation it is not improbable that all four ducts, proceeding from the two glands of the same age, may be connected together; certainly the bifurcating branches from the same duct thus become repeatedly connected. For the first two or three bifurcations the ducts decrease very little or not at all in diameter; but nearer the circumference they become smaller. The ducts, also, proceeding from the younger and smaller glands, are, of course, proportionably smaller. In one case I was able to count four bifurcations in the duct between the gland and the edge of the basal membrane.
It follows from this structure, that the basal membrane, at each period of growth, is cemented down by cement issuing from several orifices; but we shall presently find that in other genera the cement proceeds from many more orifices. In fig. 2 there is represented, by the aid of the camera, a small portion (from the outer (_a_) to the outer (_b_) being 12/100ths of an inch in length) of the basal membrane, with all the several cement-ducts adhering to it, which I could distinguish, and drawn of their proper relative sizes; this figure also shows some of the bifurcations, but no inosculation happened to be included in the space here given; the basal membrane itself has not been represented. In taking a view of a considerable portion of the basal membrane, especially towards the circumference, some parallelism in the branches could be perceived; one set of branches tending to run in the direction of the ray of the circle, and the other set in the line of the circumference.
_Elminius Kingii._--The cement-glands here resemble those of Chelonobia, but the trunk does not seem to be enlarged before entering the gland. The glands are situated rather far apart; and the chief peculiarity is, that the trunk connecting the glands is as tortuous as the track of a worm. Each gland gives out two ducts, which bifurcate repeatedly, and often inosculate, making, in parts, an hexagonal mesh-work: some of the branches do not debouch on the basal membrane, but terminate in blunt points. So numerous are the ducts, that the basal membrane may be compared to pieces of paper with the fine fibrous branching roots of some plant dried and heaped on it. Some of these ducts are very regularly jointed, and resemble a conferva,--an appearance which I believe is owing to divisions in the contained cement; other ducts are partially marked by little wrinkles, as presently to be described under Balanus. The cement, instead of, as heretofore, invariably forming a slip round and beneath the circumference of the basal membrane, here often forms little, independent, circular, and irregularly-shaped discs, each of which has issued from a single orifice. I may here add that in two species of _Tetraclita_ I saw the cement-ducts repeatedly bifurcating, with some of the branches inosculating, as in Elminius and Chelonobia.
In _Balanus balanoides_, which, like all the Cirripedes hitherto mentioned, has a membranous basis, I could only make out an amazing number of bifurcating and inosculating cement-ducts, of very various diameters. The cement-tissue, on the under side of the basal membrane, generally consisted of little circular discs, on an average from 2 to 5/1000ths of an inch in diameter; but there were also globules and short tortuous threads of cement. In some very much elongated and crowded specimens--in which, during the downward growth of the walls, the basal membrane had ceased to reach the surface of attachment, and being thus suspended had become, as viewed from the outside, deeply concave--the cement had apparently continued to try to reach the rock, and now hung down in the form of two thickish roots, some tenths of an inch in length. These roots were round, and tapered either to a fine or blunt point; one was doubled on itself, and so had become united; in the other, I could perceive five layers or sheaths of the cement-tissue, one within the other; the innermost of these layers, which once, no doubt, formed the outside surface of the root, was only a quarter of its present length.
In _Balanus tintinnabulum_, the basis is calcareous: when its upper surface is cleaned, dried, and examined under a good light, the numerous larger cement-ducts can be seen, even by the naked eye, or under a weak lens, and present an elegant appearance. These larger ducts run in parallel lines from the two chains of glands towards the circumference. They are all encrusted with calcareous matter, and in the more central parts are hidden under it; at each period of growth, when the basis is added to round the circumference, it would appear that a layer of excessive tenuity of shell is thrown down over the whole surface, just in the same way as in Tubicinella, at each period, a new and larger disc of membrane was thrown down over the pre-existing membranes with their cementing apparatus. The cement-glands, in the middle of the basal plate, seem often to give rise to small abnormal depositions of calcareous matter. When the basis (it is best to take a young specimen) is slowly dissolved in acid, all the cementing apparatus is left uninjured, adhering to the delicate tissue which before existed in a calcified condition. Near the middle I saw the two antennæ of the pupa; and from them the two cement-trunks extended about half-way towards the circumference. These two chains of glands are often placed very irregularly, but they tend to form, as in Coronula, a large angle, open towards the rostral end of the shell. The glands, close to the old antennæ, commence abruptly, of rather large size: the later-formed glands, with their ducts, are in regular order larger than the younger ones, and stand much closer together. After immersion in acid all the glands and ducts appeared empty, instead of the older ones being, as in Coronula, filled with cement. In one case I counted on each trunk twenty-five glands, besides some smaller obscure ones close to the centre.
In Pl. 28, fig. 4 _b_, I have given a drawing of two of the cement glands: the cement-trunk (_f f_) is smooth and apparently cylindrical: it becomes enlarged (at _g_) before entering the gland: it seems even to be prolonged across the gland under the form of a narrow bar (not represented), which apparently serves to keep the two ends of the trunk, on the two sides of the gland, in their proper relative places and distances. The gland itself is an elongated bag (_h_), which properly lies exactly over the enlarged portion (_g_) of the trunk, but in the drawing has been purposely displaced: it gives rise, in the later-formed glands, to a sort of neck (see the upper gland), which is either so long as to deserve rather to be called a duct and which soon bifurcates, or is quite short (see the lower gland) and gives rise to two separate ducts. On the opposite side of these glands, there is a spur (_m_), of greater or shorter length, which is evidently a rudimentary duct, for in the younger glands it existed as a perfect duct. Moreover, the first-mentioned duct often gives off branches (_t′_), having an exactly similar appearance with the spur (_m_). The membrane of which the cement-trunk (_f_), with the enlargements (_g_), is composed, is smooth, but that of the glands and of all the ducts, presents a very peculiar appearance, which at first would be called scaled, but more properly perhaps notched,--each notch being apparently formed by a line of thickened membrane, extending obliquely round only a short portion of the tube, and indenting it. The ducts, which I measured, were between 1/3000th and 4/3000ths of an inch in diameter.
In fig. 4 _a_, I have given a drawing of the two chains of glands, but with only those ducts figured which proceeded from the last-formed pair of glands. The specimen here drawn was old; and it is rare to find the structure of the ducts so simple. From both glands[72] a neck or thick duct arises, which soon bifurcates; one branch runs direct into the circumferential duct, and the other (_t_) bifurcates again; of the latter, one branch unites with its fellow from the opposite gland, and then forming a single duct (_t′_) enters, as do the two other branches, the circumferential duct. Thus, into the latter, five main ducts enter: the position of their points of entrance, with respect to the shell, varies considerably; but I think the five points tend to face the middle of the rostrum, and middle of the two lateral compartments on each side. In some other specimens, in which the ducts were nearly as simple, I observed that the neck or main duct at once divided into three branches, instead of into two, with one soon bifurcating; and on one side a rudimentary branch or spur was given off (above _t_), indicating a tendency to an additional bifurcation. In the later-formed glands, the ducts proceed only from the outer sides and form the ends of the glands furthest from the centre; but in the earlier-formed and smaller glands of the same individual, other ducts proceed from the inner sides, where in the older glands the spurs (_m_) are situated: moreover, in the younger glands, all the ducts bifurcate much oftener (how often I was not able to ascertain), before entering the circumferential duct; many of the branches, however, terminating in spur-like points. Now if we imagine twenty or thirty repetitions of the ducts given in fig. 4 _a_, (independently of the greater complication of the ducts of the younger glands), each a very little smaller than the other, and placed, with the main branches parallel, one over and within the other, we shall gain some insight into the wonderfully complicated structure of the cementing apparatus in this and many other species of Balanus.
In the basis, beneath the tubes formed by the just-mentioned radiating septa, there is a cancellated shelly mass (which, in fig. 4 _a_, was of unusual thickness), and along the crests of the branching ridges forming this cancellated mass, the sub-branches of the above branch-ducts (_i′_) run; these soon become so minute as not to be distinguished by the highest powers, and thus form a sheet of cement, which attaches the last-formed zone of the shelly basis to the supporting surface. At what point the membrane forming any one duct ceases, the cement-tissue being alone left, I was not able to ascertain; but the lower parts of the reticulated slip (_z_, _z_, fig. 4 _a_) closely resembled the cement-tissue which surrounds the disc-segment of the pupal antennæ in _Lepas australis_. The circumferential duct, here and there, forms little loops, as may be seen in fig. 4 _a_: and often two branches, running along the crests of two adjoining basal septa, proceed from a common point of the circumferential duct. The cement itself, under different parts of the basis, appears as little separate discs, as threads, globules, and as a fine network, but most commonly as simple layers. As each thick zone of shelly matter is added round the basis, the exterior branches of the ducts, between the circumferential duct and the new layer of cement beneath, are fairly imbedded in shell, and are for ever hidden, without, indeed, acid be used for the dissolution of the calcareous matter: so, also, the pre-existing ducts and glands, and the main trunk, would all have been hidden, if the layer of calcareous matter, which, I believe, is thrown down at each period of growth over the entire surface, had not been of excessive tenuity.
I cursorily examined the cementing apparatus in _Balanus galeatus_, _improvisus_ and _crenatus_, which have all calcareous bases, and belong to different sections of the genus; and the structure seemed to be essentially the same. In _Bal. galeatus_, I found the cement-ducts varying in diameter from 1/4000th to 1/10,000th of an inch in diameter. In _B. improvisus_, the cement-glands do not differ much from those of _B. tintinnabulum_; but the cement-ducts bifurcate often before entering the circumferential duct; and the little branches, which proceed from the latter, are very short, and almost immediately, owing to the thinness of the basis, blend into a slip of cement. * * * * * I hope to be excused for describing at such length, the apparatus by which sessile cirripedes are permanently attached to a supporting surface; for this is the great leading character of the sub-class, not hitherto observed in any other Crustacean.[73] It is not easy to overstate the singularity and complexity of the appearance of the basal membrane of a Balanus or Coronula: and when we consider the homological nature of the apparatus, the subject becomes still more curious: I feel an entire conviction, from what I have repeatedly seen in several genera of the Lepadidæ, both in their mature and pupal condition, and from what I have seen in Proteolepas, that the cement-glands and ducts are continuous with and actually a part of an ovarian tube, in a modified condition; and that the cellular matter which, in one part, goes to the formation of ova or new beings, in the other and modified part, goes to the formation of the cementing tissue. To conclude with an hypothesis,--those naturalists who believe that all gaps in the chain of nature would be filled up, if the structure of every extinct and existing creature were known, will readily admit, that Cirripedes were once separated by scarcely sensible intervals from some other, now unknown, Crustaceans. Should these intervening forms ever be discovered, I imagine they would prove to be Crustaceans, of not very low rank, with their oviducts opening at or near their second pair of antennæ, and that their ova escaped, at a period of exuviation, invested with an adhesive substance or tissue, which served to cement them, together, probably, with the exuviæ of the parent, to a supporting surface. In Cirripedes, we may suppose the cementing apparatus to have been retained; the parent herself, instead of the exuviæ, being cemented down, whereas the ova have come to escape by a new and anomalous course.
The only objection which I can see to the separation of the sixteen genera into the above two sub-families, may be drawn from the degree to which they blend together; thus, as far as the shell is concerned, Chelonobia, in one important internal point of structure, tends to assume the character of the Chthamalinæ; and on the other hand, Pachylasma, a member of the Chthamalinæ, has a shell, which if not examined during its earliest growth, would be placed without doubt amongst the Balaninæ. But it fortunately so happens, that in no one character of the body does Pachylasma approach the Balaninæ more nearly, than do the other members of its sub-family; or Chelonobia approach, in the same respects, the Chthamalinæ. It is only in Chthamalus, of which the shell clearly places it in the sub-family bearing its name, that in some of the species, the less bullate labrum,--the larger palpi,--the lower teeth of the mandibles being laterally double,--and the lower segments of the third pair of cirri being thickly clothed, like the lower segments of the second pair, with bristles--all show that these species make an approach in structure to the Balaninæ.
It will be seen that I have divided the Balaninæ into two little groups, according as whether the branchiæ consist of one or of two plicated folds of membrane, and as whether or not the scutum and tergum are articulated together. I have been greatly tempted to follow Drs. Leach and Gray, who have separated the second of these groups, containing the genera Coronula, Tubicinella, Xenobalanus, and Platylepas, into the sub-family of the Coronulinæ. Certainly these genera have a peculiar aspect in common, and agree in being parasitic and imbedded in the skin of Cetaceans, as is the case with the first three genera, or in that of turtles, manatee, and sea-snakes, as in Platylepas. Though these genera possess several peculiar characters, yet I can find none common to all four, excepting their imbedment in the skin of Vertebrata, their double branchiæ, and their non-articulated opercular valves; and these I do not think of sufficient importance to serve for the separation of a sub-family; for in Chthamalus, one species has double branchiæ, one species has no branchiæ at all, and the other species have single branchiæ; so again in Chelonobia, the scutum having only a horny articular ridge, makes an approach to Coronula and its allies. I may further specify that the folded walls, a singular character common to Coronula, Platylepas and Xenobalanus, fails in Tubicinella; the open tubes and the imperfect outer lamina of the parietes towards their bases, are characters which fail in one species of Platylepas; the muscles running to the opercular valves being thinly spread out, and partially without transverse striæ, is also a character which fails in Platylepas; the simplicity of the cement-ducts partially fails in Tubicinella; and lastly, the existence of small intermediate teeth on the mandibles, fails in Xenobalanus: hence, I repeat, I have not thought it prudent to admit the sub-family of the Coronulinæ though in many respects a very natural group.
The genera in the Balaninæ and Chthamalinæ are founded chiefly on the number of the compartments (the number being apparently due, as previously explained, to the fusion or abortion of certain of the eight typical compartments); and secondarily, on the nature and even form of the basis, and on the porosity of the walls. In Coronula and its allies, the non-articulated opercular valves and deeply folded walls come into play. As a justification for using these characters in distinguishing the genera, and even to a certain extent in separating the two sub-families, I must call to mind that the shell, with the basis, is not merely a dermal envelope, as amongst Molluscs, but actually consists of the first three segments of the head. The parts of the mouth and the cirri are of very little service in distinguishing the genera,--a singular fact, considering that most of the genera amongst the Lepadidæ could be distinguished by these organs,--though trifling details in their structure sometimes come in useful as specific characters. Balanus, with the sub-genus Acasta; Pyrgoma, with the sub-genus Creusia; Tetraclita, and Elminius, are genera of about equal value; though perhaps the two latter are rather more nearly related together than to the others. Chelonobia is more distinct; it shows some little affinity to the Chthamalinæ, and likewise to the four following genera. Coronula, Platylepas, Tubicinella, and Xenobalanus, are genera quite distinct from the foregoing, and from each other; yet, as we have just seen, palpably allied together. Amongst the Chthamalinæ, Pachylasma, Octomeris, and Catophragmus, are more closely related to each other than to the other two genera of the sub-family; yet Pachylasma, as far as the shell is concerned, leads into the Balaninæ, and Catophragmus into the Lepadidæ; Octomeris leads towards Chthamalus, and Chthamalus towards Chamæsipho.
_Variation._--The discrimination of the species in most of the genera, offers very great difficulties. I cannot too strongly impress on any one intending to study this class, not to trust to external characters: he must separate and clean, and carefully examine the internal structure and form of the compartments, and more especially of the opercular valves. After considerable experience, when numerous varieties of a species have been carefully examined, the eye acquires a sort of instinctive knowledge, by which it can recognise the species, though the character cannot be defined by language; but I have found that no amount of experience with some of the commonest species, will save frequent and grave errors, as long as external characters alone are trusted to. Not only does every external character vary greatly in most of the species, but the internal parts very often vary to a surprising degree; and to add to the difficulty, groups of specimens not rarely vary in the same manner.
After having given up several years to the study of this class, I must express my deliberate conviction that it is hopeless to find in any species, _which has a wide range, and of which numerous specimens from different districts_ are presented for examination, any one part or organ,--which from differing in the different species is fitted for offering specific characters,--absolutely invariable in form or structure. I may in one respect even go further, and affirm, that, if in a species, any part or organ differs remarkably from the same part in its congeners, then if many specimens are examined, especially when collected from different districts, such part or organ will be found _eminently_ variable. I may instance the antenniformed third pair of cirri in _Chthamalus antennatus_, the teeth on the posterior cirri in _Acasta sulcata_, the terga in _Pyrgoma dentatum_, the adductor ridge of the scuta in _Pyrgoma cancellatum_ and in Creusia, and other such cases: hence it will not do to found a species on a slight, or sometimes even on a considerable difference, in _any single point or organ_. On the other hand, I am far from asserting, that if only half-a-dozen specimens of some rare species of Cirripede be brought from some one quarter of the world, characters beautifully defined may not be readily discovered.
Ideas discussed in this chapter
Text from Project Gutenberg, public domain in the United States.