A Monograph on the Sub-class Cirripedia · Charles Darwin
LEPAS ET BALANUS AUCTORUM.
_Basis._--The basal membrane is thin, and is divided, but not very plainly, into concentric slips, marking the successive increments of growth. In the middle of it, in two young specimens, I found with great difficulty the pupal prehensile antennæ: they were of small size, measuring from the extreme edge of the main or second segment to the end of the disc, only 27/6000ths of an inch: the disc appeared narrow (as in Pollicipes and Scalpellum), with a single spine at the proximate end: the ultimate segment, placed as usual at about right angles to the disc, bore two groups of shorter and longer spines, but I could not count how many. The antennæ were enveloped in a mass of cement of a yellow colour, resembling in all its characters the cement of other Cirripedes. In only one case, I believe I saw bifurcating cement-ducts, of extreme tenuity, viz. 1/15,000th of an inch in diameter. The sheet of cement on the whole under side of the basal membrane, not rarely shows a very irregular reticulated structure. For convenience sake, it will be best to defer the discussion on the very anomalous, though slight, powers of excavation which this genus possesses, and which I must attribute to the effects of some substance secreted probably by the cement-organs. I will here only mention, that the specimens which have excavated a depression, are less firmly attached than those, which have not acted on their support; and that, in the former case, the basal membrane, for a considerable space in the middle, becomes quite detached.
_Animal's Body._--The body is much flattened and, owing to the little development of one side of the shell, lies parallel to the surface of attachment. The prosoma is but little protuberant. The articulations of the thorax are unusually straight and transverse. The _Mouth_ is also much flattened: it is placed rather distantly from the adductor scutorum muscle, owing to the production of the lower margin of the labrum. The _Labrum_ is not notched, or even hollowed out in the middle, or (excepting in _V. nexa_) bullate; its crest is surmounted by about eight (more numerous in _V. nexa_) little teeth, or by some fine bristles. The _Palpi_ are of moderate size, with their tips nearly meeting; they are slightly curved, and have bristles only on their outer sides and extremities: they are apparently capable of being lifted up and down by a muscle attached to them, just outside the rounded swelling on each side of the labrum to which they are articulated: in _V. nexa_, however, the palpi are very small and narrow, and their tips do not nearly meet. In this genus, therefore, we find the swollen state of the labrum and the size of the palpi--characters generally invariable and of high classificatory importance--variable. The mandibles have three upper main teeth, with two or three minute lower teeth, or, in _V. nexa_, with the lower part pectinated with small spines: in _V. Strömia_, I have seen traces of the second tooth being laterally double--a character of some importance. The _Maxillæ_ have a notch under the upper pair of large spines, with the lower part bearing, as usual, a double row of bristles, and forming a large step-formed projection: these organs are furnished with the usual apodeme and muscles. The _Outer Maxillæ_ are prominent, and deeply lobed on their inner surfaces, the two lobes being clothed with bristles.
_Cirri._--The first pair are attached, as usual, on each side of the mouth, and stand some way apart from the five posterior pairs. The second and third pairs differ considerably in structure from the three posterior pairs, which are much elongated. The _first pair_ (excepting in _V. nexa_) is short, with the two rami slightly unequal in length, and with the segments thickly clothed, as usual, with spines. The _second pair_ is remarkable from the posterior ramus being more than twice as long, and containing thrice as many segments, as the anterior ramus, which is barely as long as the shorter ramus of the first pair: the segments in the anterior ramus of the second pair (only five in number in a full-sized specimen) are broader and more protuberant in front, and more thickly clothed with spines (the terminal spines being doubly pectinated), than are the segments on the posterior ramus; on the latter, the uppermost segments have their bristles arranged in front in simple pairs, with the dorsal spines long, the lower segments being more thickly clothed with bristles, owing to the development of lateral rows. The _third pair_ resembles in every respect the second pair, except in being a little longer, and in the bristles on the posterior ramus being less crowded, more resembling the arrangement of those on the posterior cirri. In _V. nexa_, however, there is not so great an inequality in length or dissimilarity in structure in the two rami of the second cirrus, and only a very slight difference of any kind in the two rami of the third pair. _Fourth_, _fifth_, and _sixth pairs_ have numerous elongated segments, bearing four or three pairs of long slender spines in front, with a single minute bristle between each pair, and with two or three slender spines in the dorsal tuft.
There is a considerable amount of variation in the proportional length, and in the number of the segments, of the several cirri in _V. Strömia_; in some specimens the two rami of the fourth pair were unequal in length; in some, nearly all the cirri on the lower or attached side were shorter than those on the upper side. _Caudal Appendages._--These are of most unusual length, sometimes even exceeding those of _Ibla quadrivalvis_, which surpasses, in this respect, all other cirripedes. They arise on each side and over the anus. They consist of numerous (sometimes as many as twenty-four), unequal, cylindrical, thin segments, bearing, at their upper ends, a circle of long and very slender spines. They sometimes equal two thirds or even four fifths of the length of the sixth cirrus; but their length, and the number of their segments, (sometimes imperfectly divided), varies much in different specimens of the same species, and sometimes even on opposite sides of the same individual. In some very young shells, as big as a pin's head, the caudal appendages were proportionally extremely short, and consisted of only two or three segments. No muscles enter these organs; and when the animal is taken out of its sack, they project straight out behind, instead of being curled in, like the cirri.
_Anatomical Structure._--The animal's body is attached to the two scuta by the adductor scutorum, and by the other usual muscles running towards the mouth, and surrounding the prosoma. The whole external covering or shell has no other muscles; Verruca thus differing from the Balanidæ and Lepadidæ; but the shell is attached all round, near its circumference, to the basal membrane, by a band of very short fibres, appearing like muscles, but really ligamentous, as determined for me by Professor Quekett. _Branchiæ_ are entirely absent. The _alimentary canal_ presents all the usual characters, but in the prosoma is rather abruptly bent back on itself. The orifices of the two _olfactory pouches_ are not at all prominent; they are placed directly under the outer maxillæ, (homologically in their middle segment), just above a small, medial, tongue-like apodeme. The orifices of the _acoustic sacks_ appeared to be in their usual position beneath the basal articulations of the first pair of cirri. The _vesiculæ seminales_ occupy their usual position in the prosoma; they are not much convoluted; they unite before entering the penis. The probosciformed penis is imperfectly ringed; it is thick and short, and tapers much more abruptly than is usual; it supports a few very thin hairs.
The _ovarian cæca_ are spread over the basal membrane, at the bottom of the sack; hence they in fact lie almost on one side of the animal: they consist of two main trunks, proceeding out of the animal's body at the rostral end of the sack, which then branch and inosculate. In specimens of _V. Strömia_ collected by Mr. Peach for me, in Cornwall, during the first week of April, there were included two ovigerous lamellæ, placed transversely across the rostral and the carinal end of the sack: the lamellæ were .11 of an inch in length; they appeared loose and not attached, as in the Lepadidæ, to any ovigerous fræna. The ova, in their earliest age, have one end much pointed, and are 8/1000ths of an inch in length; they become blunter and increase a little in size before the larvæ burst forth. The larvæ, both during their earliest stage and after the first moult, have been excellently figured and described[137] by Mr. C. Spence Bate: they present no particular characters distinct from the larvæ of other Cirripedes. I will only further add, that the structure of the prehensile antennæ still adherent to the basal membrane, indicates that the larva in its last stage,--that is the locomotive pupa,--has a normal character. [137] 'Annals and Mag. of Nat. Hist.,' 1851, Pl. 7, fig. 8-10.
_Affinities._--These have been sufficiently discussed under the family; I need here only remark that all the species, with the exception of _V. nexa_, are intimately allied together. _Range--Habits--Geological History._--The genus Verruca ranges, being represented by four species, from Iceland to Cape Horn. The species that is found in Tierra del Fuego extends up the west coast to Peru. Our northern form, _V. Strömia_, (if I may trust a specimen in the British Museum, apparently ticketed in an authentic manner), occurs also in the Red Sea; and this is the only locality in the eastern hemisphere whence I have seen this genus. The species seem generally to live in rather deep water: I procured _V. lævigata_ from nineteen fathoms, on the east coast of Patagonia: _V. Strömia_ is found, according to information given me by Professor Forbes, on the British shores, between five and fifty fathoms, and on the steep shore off Mull, in ninety fathoms; but Mr. Thompson assures me that he once saw it adhering to tidal rocks and likewise to some floating bark. Generally the species are attached to living organic bodies, especially shells of Mollusca and of Cirripedes, to Gorgoniæ, and Laminariæ; less frequently to rocks. We shall immediately see that it has slight powers of excavation. This genus is geologically older than any true sessile cirripede or member of the Balanidæ: _V. Strömia_ is found in the Glacial Deposits and in the Red and Coralline Crag of England: another species (in a state not to be identified) occurs in the ancient Tertiary formations of Patagonia; and another in the Chalk of England and Belgium. The fact of this Family ascending to a Secondary epoch accords, in an interesting manner, with its affinities; inasmuch as though in appearance a sessile cirripede, it is almost equally related to the Lepadidæ and Balanidæ, and is more nearly related to the Lepadidæ than to the Balaninæ, or typical members of the Balanidæ: of the latter, none have hitherto been found in any Secondary deposit, whereas the Lepadidæ culminated during the Cretacean period.
_Powers of Excavation._ My attention was called to this subject by Mr. Hancock, whose excellent researches on the boring of Mollusca are well known. _Verruca Strömia_, when attached to shells destitute of an epidermis, excavates, as he informed me, a slight depression, deepest in the middle; but when the epidermis is present no effect whatever is produced. We shall presently see that the central depression is in some degree distinct from that of the circumference. I have since found Mr. Hancock's observations strictly applicable to _V. lævigata_, _V. Spengleri_, and to an ancient tertiary species from Patagonia. From having found that the following cirripedes, viz., Lithotrya, Alcippe, and Cryptophialus, all form their deep excavations by mechanical means, and from having read the above-mentioned memoirs by Mr. Hancock on the boring of mollusca, I was strongly impressed with the idea that the action in Verruca would likewise prove mechanical: but from the following facts I have come to the conclusion that the excavation must be due to a solvent, probably poured out from the cement-ducts, which debouch on the under side of the basal membrane.
In the first place, an epidermis, as just stated, perfectly preserves the shells of the various species of mollusca and certain cirripedes, to which I have seen Verruca attached: this is well shown by comparing the effect produced on the same shell in parts covered by the epidermis and in parts whence it has been abraded; or where the shell of the Verruca had fixed itself, whilst very young, within a crack in the epidermis, and had subsequently, by its growth, turned up the edges, and had then acted on the underlying shell; whereas the specimens attached to the sound epidermis had not produced the smallest effect. Again, I have seen an epidermis-covered mussel-shell encrusted by a hard nullipora, on which _V. lævigata_ was attached; and here the calcareous nullipora, under the middle of the basal membrane, was entirely corroded away, whilst the underlying epidermis and the shell beneath it, were not in the least affected. The protection afforded by the epidermis is still more strikingly shown by contrasting shells with very sharp prominent ridges, when thus invested and when naked, to which Verrucæ have been attached: I have given a figure (Pl. 21, fig. 6) of a piece of an invested Venus, from the surface of which a _V.
Spengleri_ had been just removed; on the other hand, I have seen a Peruvian Discina in which even sharper ridges, covered by epidermis, were left absolutely untouched, although projecting deeply into the shell of an attached _V. lævigata_. I have seen several specimens of this latter Verruca (which has the power of corroding naked shell as deeply as its congeners), attached to the membrane-covered variety of _Balanus lævis_, the shell of which was thus perfectly preserved: now this membrane is little more than the 1/2000th of an inch in thickness; it is not hard, and so brittle that it generally separates with the Verruca, leaving the underlying shell of the _B. lævis_ with its lines of growth glossy and perfect: it appears to me impossible that a membrane so thin and brittle could resist an action, if mechanical, which has worn away from twenty to forty times as great a thickness of hard shell; but the thinnest film of any matter on which acid does not act, as of grease in certain forms of printing, will perfectly preserve the underlying substance, and as I have ascertained by putting on a drop of acid, is the case with this membrane. I have removed several scores of shells of _V.
Strömia_ from the stems of Laminariæ, and when the latter were washed and slightly dried, generally not the least effect could be seen, except that the spots where the shell had adhered were glossy from the still adherent basal membrane: yet the stems of Laminariæ are far from hard. In some cases, however, the attachment of the Verruca seemed to have produced a very slight depression on the Laminaria, but this, I think, may be safely attributed to the growth of the surrounding surface; for I have seen exactly the same effect produced by the attachment of the discs of the antennæ of a Lepas, and these discs, with their long spines, could not possibly produce any excavation; nor is Lepas or its pupa in any case a burrowing animal. Again, I have seen a few specimens of Verruca attached to Gorgoniæ, and they had not acted in the least on the horny axis. I have examined numerous specimens of _V. Strömia_ attached to three pieces of slate-rock, and to one piece of red sandstone, all from different localities, and no effect whatever had been produced; yet the slate-rock, especially in one instance, was soft.
Mr. Bate, to whom I am indebted for some of these specimens, also informs me that he could discover no impressions on the slate-rocks, whence specimens of the Verruca had been removed. On the other hand, I have had two specimens of limestone, with attached Verrucæ, one coarse and very impure, and the other hard and marble-like; and in both cases there was a distinct central slight cavity, including loose gritty matter. The loose particles evidently resulted from the unequal action either of a solvent or of some mechanical power on the rock, for it is improbable in the highest degree that the shells should have fixed themselves exactly over small collections of loose particles, even if such could possibly have remained on projecting surfaces of sea-washed rocks.
The above facts seem to indicate pretty plainly that the excavation of the support does not depend on its hardness, but on its containing calcareous matter, liable to be acted on by some solvent: but as this view, considering what we know of Lithotrya and of the two other burrowing genera of cirripedes to be hereafter described, appears improbable, I will add a few additional observations. I most carefully examined the shell and basal membrane of Verruca, and likewise the tissues left after the dissolution of the shell in acid, and could detect no structure at all fitted for boring; and what appears more important, there was no apparent difference in the state of the specimens which had and had not excavated a hollow; and this, I think, would certainly have been the case (as in Lithotrya) if the action had been mechanical. It is not easy to ascertain, owing to the small effect at any time produced, at how early an age Verruca begins to act on its support; but I found two sets of specimens only 1/20th of an inch in basal diameter, which had certainly commenced. The ribbed shell, (Pl.
21, fig. 6), especially the middle rib, shows, in a somewhat exaggerated degree, the _typical_ form of the excavation; it may be here seen that the excavation is of the same depth for some little distance from the circumference towards the centre, but that in the middle it suddenly becomes deeper. I have seen several specimens with a central hollow, without any, or with scarcely any, marginal depression, and likewise the reversed case. These several facts show that the central excavation cannot be due to an equable action, prolonged during the whole growth of the shell, having thus affected the middle more than the circumferential parts, for in this case the excavations would have sloped into each other. In specimens which have not at all acted on their support, the whole basal membrane is firmly attached, as in all ordinary cirripedes, to the supporting surface; but in those which have acted, the middle portion of the basal membrane is quite unattached, and the circumferential portion is, I think, less firmly attached than is usual; but between these two portions, there is a circular zone strongly cemented to the supporting surface, and which alone keeps the shell in its place.
Now, on the mechanical theory, to account for the circumferential hollow, the basal edges of the shell together with the circumference of the basal membrane must be subjected to movement, but the shell is united to the basal membrane by corium and by transparent structureless chitine (both of which may be left out of question) and by a circle of short fibres, which adhere at their lower ends to the firmly cemented circular zone, and by their upper ends to the shell; and these fibres have been very carefully examined by Professor Quekett, and pronounced to be not muscular, but exclusively ligamentous, and therefore incapable of moving the edge of the shell. The basal membrane over the central hollow is, as stated, quite loose: its lower surface, formed by a reticulated layer of horny cement-tissue, shows no signs of abrasion, and the membrane is so brittle and tender, that in specimens which have been once dried and then well soaked, it almost invariably cracks when the shell is removed, owing to its mere adhesion to the delicate inner tunics of the sack; yet on the mechanical theory, the wearing of the central hollow must have been caused by the action of this middle portion of the basal membrane,[138] which, it may be repeated, is destitute of muscles.
From the presence of the prehensile pupal antennæ, enveloped in cement, nearly in the centre of the basal membrane, it is certain that this spot was originally attached to the supporting surface, and has since been detached from it; as, moreover, the central hollow goes on increasing in diameter with the growth of the shell, it is certain that the inner edge of the firmly attached circular zone of basal membrane must likewise continually go on becoming detached: it may, then, be asked by what force can the basal membrane, seeing that it is united to its own shell above only by fibres of ligament near the circumference, be continually torn away from the underlying support, to which it is strongly cemented? On the other hand, on the theory of a solvent slowly poured out from the cement-ducts, its separation from its support is simply explained.
It might be supposed that the calcareous matter, when dissolved, would not be able to escape from the central hollow, owing to the basal membrane being so firmly cemented all round it; but the attachment is by a reticulated layer of cement; and I infer that it must be permeated by open passages, from the fact of the hollow being often filled, in dried specimens, by a bubble of air, instead of the basal membrane being pressed closely down into the hollow, as would have been the case had the hollow been hermetically sealed up. I have seen a few instances in which the bottom of the central hollow was occupied, (as was remarked to me by Mr. Hancock), by a little chalky and gritty matter; and in the case of one of the specimens of calcareous rock, before alluded to, by coarse grains and oxide of iron; this seems quite compatible with a solvent acting more readily on certain parts of the rock or shell than on other and less soluble parts or particles. [138] Mr. Hancock suggests to me that the basal membrane, on the mechanical theory, need not itself move; the motion of epithelial scales, were they transferred into cutting agents, might be supposed to be sufficient. But of such scales, though I used very high powers, I could see no trace; and their presence on the under side of the layer of cement seems hardly possible. Moreover, according to Von Siebold ('Anatomie Comparée,' tom. 1, p. 412), ciliary action has not been observed in any Crustacean, or indeed any Articulate animal. This same statement is likewise made in Annals and Mag. of Nat. Hist. 1854, p. 136, by Dr. T. Williams.
The greatest depth of the central hollow, in any specimen seen by me, even measuring from the top of a rib in the case of a ribbed shell, to the deepest point, was only 1/50th of an inch; but considering how much depressed the shell of Verruca is, I have no doubt that this small gain of space is of service to the animal: we must suppose the loose middle portion of the basal membrane is stretched slightly, or splits and is repaired, so as to fit the hollow. With respect to the even much slighter circumferential excavation, it barely equals in depth the thickness of the extreme edges of the walls; it must, I presume, give strength to the shell when _laterally_ pushed; but it certainly appeared to me that the individuals which had excavated a depression for themselves, could be pried vertically up much more easily than those which had not acted on their support. Finally, we must suppose that the hypothetical solvent is poured out of the cement-ducts at the extreme circumference of the basal membrane, which is almost loose and destitute of cement, so as to slightly corrode outwards and downwards the calcareous support; the action here then stops, and this rim of basal membrane becomes, after a new rim has been formed and as the shell grows outwards, firmly cemented down to the now slightly excavated surface of attachment; but during all the time the solvent goes on acting in the middle, and continues, during the whole growth of the shell, to encroach on and dissolve the supporting surface from under the inner edges of the previously cemented down, circular zone of basal membrane. I have discussed this subject at considerable length, as it appears to me an interesting one. In this case we have the action of ciliæ[139] and of respiratory currents, to which in the case of Mollusca so much has been attributed, entirely eliminated. It is, also, an interesting fact, that within the same Order we should have some Cirripedes boring by simply mechanical means, and others by a chemical solvent.[140]
Ideas discussed in this chapter
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