A Monograph on the Sub-class Cirripedia · Charles Darwin
LEPAS ET BALANUS AUCTORUM.
Thus it is rendered probable that this fixed valve is a modified scutum; but a surface of attachment for one end of the adductor muscle might, perhaps, have been developed on any other valve, or a scutum might have become fused with a lateral valve of the shell; the shell on this latter view being rendered in idea more symmetrical. But when a very young specimen is carefully examined, it is found that the moveable and fixed scutum, the moveable tergum and its opposed valve or fixed tergum, at the first period of calcification, resemble each other quite closely; but that, as each zone of shell is added, the differences become rapidly greater and greater: hence, it may be considered as directly proved, that the two fixed valves (S′ and T′ in all the figures in Pl. 21), which are opposed to the moveable valves of the operculum (S and T), consist of an extraordinarily modified scutum and tergum.
It has been shown (p. 129), that at the period of the metamorphosis, the two scuta, the two terga, and the carina of the Lepadidæ, commence their growth, under the form of the so-called "primordial valves," and so differ from all the other valves when such occur: now, in two species of Verruca, I have found closely analogous primordial valves on the apices of both the moveable and fixed scutum and tergum (thus affording strong additional evidence that their nature has been rightly interpreted), and on one of the two remaining valves, namely, that at the posterior or carinal end of the shell. Hence, we may safely infer, that this latter valve, which, though very much more developed on one than on the other side, is so far medial as to curl round and cover the line of opening between the moveable and fixed tergum, is really a carina. The sixth valve differs only very slightly in shape from the carina, and is directly opposed to it; therefore, in accordance with all analogy, it must be the rostrum. Consequently, the shell in Verruca consists of a moveable scutum and tergum, a fixed scutum and tergum, a carina and rostrum, and, as we shall immediately see, a membranous basis--the basis being, as in all sessile cirripedes, the homologue of the peduncle in the Lepadidæ.
The moveable scutum and tergum stand at about right angles with the fixed pair; and as these latter form a part of the wall of the shell, which is always steep on this side, the moveable pair, which close the orifice, are nearly horizontal or parallel to the basis and surface of attachment. Hence, the animal's body, which is attached between the two scuta, but nearest to the moveable scutum, also, lies nearly parallel to the surface of attachment; and I was consequently at first led to suspect that the basal membrane was one side of the shell in a modified condition; but the presence of the prehensile antennæ of the pupa in nearly the middle of this membrane, and the sheet of cement-tissue on its under side, demonstrate that this membrane, though lying on one side of the animal, is the true basis. To make all the parts in Verruca hold the same position as in other cirripedes, relatively to the surface of attachment, we must develope the carina and rostrum equally on both sides of the true longitudinal axis of the shell, and insert the newly-developed portion between the basis and the fixed scutum and tergum, reducing the latter in size, and tilting a little up the moveable scutum and tergum; and by this means the animal's body would be turned, so that its dorso-ventral longitudinal plane would stand at right angles to the basal membrane.
Extraordinarily great as is the difference between the right and left sides of the whole shell, yet in all the species it seems to be entirely a matter of chance whether it be the right scutum and tergum with the right side of the rostrum and carina, or the left scutum and tergum with the left side of the rostrum and carina, which become abnormally developed. Nor does there seem to be any relation between the side of the operculum to be attached, whether right or left, and the nature of the surface of attachment; for I have seen many specimens adhering to perfectly level surfaces, and to quite cylindrical branches of Laminariæ; and in these cases, however the larva might attach itself, there could be nothing to favour the development of one side more than the other. Although the attached scuta and terga are larger than the moveable pair, yet, owing to the small development of the carina and rostrum on the attached side, the upper or unattached side must be considered as the most developed. In this respect, and in the circumstance of either right or left side being modified, we are reminded of the structure of _Pœcilasma Kæmpferi_ (described in my former volume on the Lepadidæ), in which the valves on the side of the capitulum, nearest to the crab's body, to which the specimens were attached, were somewhat less developed than those on the opposite side. I may add, that in ordinary Crustaceans, as I am informed by Professor Bell, the unequal development of the thoracic limbs seems quite capriciously to affect either the left or right side of the body.
_General Appearance of the Shell._--The shell is in most cases much depressed and irregularly circular; the side formed by the fixed scutum and tergum is always steeper than the other side: the colour is white or pale brownish, and in _V. nexa_ pale red. The surface is naked. The size is small, rarely exceeding a quarter of an inch in diameter, and the whole shell often appears like a mere scale on the surface of attachment. The most remarkable feature in the external aspect is due to the suture between the rostrum and carina, which is formed by oblique, interlocking plates or folds; as all these plates continue to be added to at their extremities during growth, the upper plates become longer than the lower ones; and the plates on both sides of the suture together form a triangular area, with the broad end uppermost, somewhat like the radius of a sessile cirripede: they act, also, like a radius, for their growth serves to separate these two valves, and so adds to the diameter of the shell. The suture between the rostrum and fixed scutum and that between the carina and fixed tergum are nearly of the same nature, but the former is more conspicuous than the latter; neither are so conspicuous as that between the carina and rostrum: accordingly as the right or left scutum and tergum are moveable, so the suture, second in plainness, (see Pl. 21, fig. 1 _a_, and 1 _d_,) is placed to the left or right hand. The fourth suture, between the fixed scutum and tergum, as viewed externally, is straight, and so very obscure that it has been overlooked by some authors, and the shell described as consisting of only three nearly equal pieces, for the fixed scutum and tergum together are about equal in size to the carina or rostrum. The orifice approaches more nearly to an unequal-sided triangle, with the apex broadly truncated, than to any other figure. The operculum fits with remarkable closeness, and is surrounded by a slight rim, formed by the edges of the four other valves.
_Moveable Scutum and Tergum._--The scutum (S in 1 _b_ and 5) is narrow and very small, barely equalling half the size of the tergum, and therefore proportionally much smaller than in any other cirripede; in a very young shell, however, (of _V. Strömia_) less than a pin's head in size, the scutum equalled the tergum in size. The valve is remarkably thick; it is generally depressed down the middle; but in _V. nexa_ this part is longitudinally ribbed. The occludent margin is curved. On the tergal margin there are two articular ridges (with a deepish furrow between them), of which the upper one (′ in S, in fig. 1 _b_, and 5) extends from the apex about half-way down the valve; and the other, or lower articular ridge (′′ in S), generally runs down nearly to the basal margin: an angle, running from the apex to the basi-tergal corner of the valve, appears like a third articular ridge, but cannot properly be considered such. The above two articular ridges interfold with analogous ones on the scutal margin of the tergum, and so lock the valves together. On the under side (fig. 1 _f_), the surface is bounded along the occludent margin by a slight rim: there is generally a very slight depression for the adductor muscle; but in _V. Spengleri_ there is a straight, short, sharp (Pl. 21, fig. 2), prominent adductor ridge.
The _moveable tergum_ is broad and rhomboidal. Externally a prominent axial ridge (′′′ in T, in fig. 1 _b_, &c.), which widens downwards, runs from the apex of the valve to the basal point, and there projecting slightly, causes the scutum to be indented; this indentation on the scutum appears like a third articular ridge, lying beneath (′′), S, in fig. 1 _b_, &c. Above the lower and axial ridge, on the scutal margin of the tergum, there is a middle articular ridge, which locks in, between the lower (′′) and upper articular ridges (′) of the scutum (S). Again above the middle ridge there is an upper and third articular ridge (′), which is either quite distinct, as in fig. 5, T, or more commonly is formed by the occludent margin of the valve, as in T, fig. 1 _b_. The broad extremity of this upper articular ridge is often produced into a slight projection, or shoulder, and this always underlies the scutum, of which the under and upper surface is indented or furrowed (see fig. 1 _f_), in order to receive this shoulder. The upper articular ridge of the scutum (′, S, 1 _b_) locks in between the upper articular ridge or occludent margin (′, T), and the middle ridge (′′, T) of the tergum. Hence, altogether, there are three articular ridges on the scutal margin of the tergum, the occludent margin being generally counted as one; whereas, on the tergal margin of the scutum, there are only two ridges, though, as before noticed, an outer indentation, which is developed as a ridge in _V. nexa_ (fig. 5, S), might almost be counted as a third articular ridge.
I may here just remark, that the furrow between the two ridges on the tergal margin of the scutum, resembles the articular furrow in the scutum of the Balanidæ; but it may be doubted whether the resemblance be more than superficial, as this furrow, in the case of Balanidæ, receives the edge itself of the tergum, whereas here it receives only a ridge, proceeding from the apex of the tergum, to a nearly middle point on its scutal margin. Finally, I may add, that the tergum in this genus, in general shape, in growth (presently to be referred to), in the manner in which the upper scutal shoulder is overlapped by the scutum, and in the presence of the axial ridge, presents a very striking resemblance to certain old fossil species of Pollicipes, and to a limited extent to the living species of Lithotrya. The scutum and tergum being interlocked, move together; they can be firmly shut by the contraction of the long adductor scutorum muscle. Their opening appears partly due to the elasticity of the membranous hinge (representing the opercular membrane), by which they are attached transversely, just beneath the summit of the carina and rostrum. No doubt the protrusion of the cirri effectively aids the act of opening. These valves are not capable (nor, of course, the other valves) of any other movement; for there are no muscles for such movements.
_Fixed Scutum and Tergum._--The fixed scutum is larger than the fixed tergum, and therefore has the same proportions as the homologous valves in ordinary cirripedia, but reversed proportions compared with the moveable scutum and tergum. The shape of neither valve can hardly be described. The _fixed Scutum_ (S′ in all the figs.), externally, seems at first to consist of two portions, namely, a curved occludent rim (_a_), closely resembling the opposed occludent margin (_a_) of the moveable scutum (S), having in fact undergone very little modification, (as may be best seen in Pl. 21, fig. 1 _b_); and secondly, of a much modified portion (marked ′′), which resembles in outline and state of surface the rest of the walls of the shell, and may be called the parietal portion. The rostrum (A) curls round the end of the occludent portion, under an edge (_b_), evidently answering to the basal margin (_b_) of the moveable scutum, and is simply united to this portion by membrane, but beyond this part, it is articulated to the parietal portion (′′) of the fixed scutum, by oblique interlocking ridges, like those forming the suture between the rostrum and carina. In _V. nexa_, however, (fig. 5) the rostrum (A) does not curl round any part of the interlocking fixed scutum (S′), but articulates with it by a straight suture. Internally, the fixed scutum (S′ in figs. 1 _e_, 1 _c_, of reversed shells) has a surprisingly large, thin adductor plate (_m_ in fig. 1 _c_, 1 _b_), with a rounded outline, projecting nearly parallel to the basis or surface of attachment; the adductor scutorum muscle is attached to its upper surface, and consequently the animal's body lies between this plate and the moveable scutum. In the fixed scutum of _V. nexa_, however, there is a deep pit, instead of a plate, for this muscle.
The _fixed Tergum_, likewise, consists of two portions--a middle and lower, or parietal portion, and a rim or upper portion; the rim consists of two unequal arms, answering to the two upper margins (not merely the edges) of the rhomboidal moveable tergum; the longer rim (_x_ in T′, see fig. 5) answers to the carinal margin (_x_) of the moveable tergum, and may be called the carinal rim; and the shorter rim (_o_ and ′ in fig. 5) answers to the occludent margin (_o_ and ′) of the moveable valve, and may be called the occludent rim. The carina curls round the end of the carinal rim, under an edge, _z_ (much foreshortened in T′ in fig. 1 _b_, and best seen in fig. 5), answering to at least a large part of the basal margin (_z_) of the moveable tergum, and interlocks, by a serrated suture, with the edge of the parietal portion of the valve. Internally (fig. 1 _e_, less plain in 1 _c_) there is a transverse ledge, notched in the middle, and sometimes deeply hollow beneath, running across the valve in about the line of the adductor plate of the fixed scutum: this ledge, in fact, marks and is partly caused by, the line of separation between the central or parietal, much modified, and the scarcely modified, upper or rim portion of the valve. The use of this ledge is apparently to give attachment, as does the under side of the adductor plate of the fixed scutum, to ligamentous fibres, presently to be mentioned, by which the shell is attached to the basal membrane: the carina and rostrum being so much more gently inclined, do not stand in need of a ledge for their attachment.
By comparing the moveable scutum and tergum with the corresponding fixed valves, in all the species, the modification of the latter may be clearly made out to have been effected as follows; and the case appears to me a striking and interesting one. The moveable scutum and tergum lie in the same plane, and are articulated by the means of three ridges on the tergum (including the occludent margin), and by two on the scutum. The fixed scutum and tergum have to be curved, and to be greatly increased in size; and this is brought about, as we shall see, by the large development of a certain small portion of each valve. Comparing first the moveable tergum (T) with the fixed tergum (T′), the umbo of growth matches the umbo of the four margins of the moveable valve, the carinal (_x_), basal (_z_), and occludent (_o_ in fig. 5), margins can be identified with certainty in the fixed valve, from their close similarity in shape, their absolute apposition, or correspondence in position.
There remains only the scutal or articular margin, with its three articular ridges; of these, the uppermost (′), inasmuch as in most of the species it is hardly distinct from the occludent margin, can, as we have just seen, be clearly identified, and is overlapped, as it normally should be, by the upper tergal corner of the fixed scutum: the second or middle articular ridge, though not so distinct as in the moveable valves, can be plainly recognised (′′), T′, in fig. 1 _b_, and 5; and it serves its normal function of articulating the two valves together. But when we look in the fixed valve for the third or axial ridge (′′′), we find in its exact place, namely, extending from the umbo to the extreme opposite end of the valve, between the second articular ridge (′′) and the basal margin (_z_, see fig. 5), only that portion of the valve which I have called the parietal portion; consequently, I do not doubt that this really is the axial ridge largely expanded.
So again in comparing the moveable scutum (S) with the fixed scutum (S′); two of the three margins of the former, namely, the occludent (_a_, see fig. 1 _b_) and basal (_b_), can be identified without a doubt in the fixed valve: the third and tergal margin remains; this should have two articular ridges; of these the upper one, still serving its normal function, can be detected in all the species (′ in fig. 1 _b_), and can be seen pretty plainly (′ fig. 5) in _V. nexa_: but of the lower and other articular ridge there is no sign,--excepting indeed the whole parietal portion of the valve, which, from holding an exactly homologous position with the lower articular ridge of the moveable valve, I cannot doubt in this ridge expanded and curiously metamorphosed. Hence, in both fixed scutum and tergum, it is the outermost or lowest of the articular ridges which has been modified and expanded, so as to rest on and be fixed to the surface of attachment. It would appear as if it had resulted from one ridge in each of these valves having been thus used up by expansion (so to express myself), that the suture between the fixed scutum and tergum is more simple than any other suture in the whole shell; and it is owing probably to this straightness, and consequent _tendency_ to weakness, that the valves do not grow along this line, and so do not become separated from each other during growth, as on the three other lines of suture. As it actually is, owing to this suture never being separated, it is even stronger than the others; its edges on the inside (fig. 1 _c_), I may add, are a little inflected or prominent.
_Rostrum_ and _Carina_: these valves differ from each other, only in the former (A) being rather the largest, and in being more plainly articulated with the fixed scutum, than is the carina (B) with the fixed tergum. Their umbones stand in their normal places, at the two ends of the orifice leading into the sack, that is, facing the dorso-ventral longitudinal plane of the animal; but they are very unequally developed on the two sides, and hence they rise very obliquely from the surface of attachment. Their summits are nearly square, which is caused by the continued growth on both sides of the oblique plates or ridges, by which they are articulated with the adjoining valves. These plates strikingly resemble, as already stated, the radii in certain species of Chthamalus. Without these articulating plates, the outline of the rostrum and carina would have been triangular, with the apex upwards. In _V. nexa_, in which the walls of the shell are almost perpendicular, the rostrum (A, fig. 5) is very peculiar and patelliformed, with the umbo sub-central: this results from the development of a border at the upper end of the valve. In this same species, the basal edges of the rostrum, carina, fixed scutum and tergum, are rectangularly inflected, so as to form a ledge round the basis, as in the case of some few species of Chthamalus,--the ledge appearing like part of the real basis. During the growth of the shell, the upper internal ends of the carina and rostrum are either rendered solid, or a ledge is formed on the inside across their summits, hollow beneath, like the sheath of the Balanidæ, to which solid or hollow ledge the basal margins of the moveable scutum and tergum are attached by a rim of membrane, forming a hinge.
_Direction of Growth: Minute Structure of Valves._--The shell grows downwards all round its basal margin. As far as the diametric growth of its upper part is concerned, there may be said to be only three valves, for the fixed scutum and tergum never become, as already stated, separated; on the three other lines of suture, the valves are added to on both sides; and thus the whole upper part of the shell, and the orifice, increases in diameter. The moveable scutum and tergum grow along their basal margins, and along the margins by which they are articulated together; but the scutum in this latter respect, less than the tergum. The summits of the moveable scutum and tergum, during continued growth, become either worn away, or they project freely; in this latter case, an internal ledge is added round the upper end of the fixed scutum and tergum, so as to keep the orifice accurately closed. In _V. nexa_ the rostrum, with its sub-central umbo, is anomalous, as already stated, owing to a broad upper internal border growing in a direction almost directly opposed to the basal growth of the moveable opercular valves. In young specimens, on the apices of both scuta and both terga, and on the carina, but not on the rostrum, _primordial valves_ may be distinguished, resembling the valves, so called, which first appear (p. 129, Introduction) after the metamorphosis in, the Lepadidæ. In the Verrucidæ, however, they are calcareous; and the minute transverse cylinders, of which they appear to be composed, stand further apart, causing the surface of the primordial valve to be marked with little separate circles, instead of by hexagons.
The shelly matter of which the valves are composed is translucent: it is remarkably destitute of any investing membrane. The under surface is marked with rows of minute approximate pores, parallel to the lines of growth, into which the corium enters: after a portion of shell has been dissolved in acid, these threads of corium are seen to change, a short distance within, into cylinders of yellow chitine, running obliquely through the substance of the valve. These cylinders are about 1/2000th of an inch in diameter, but in parts they are spindle-shaped and twice as thick: they vary in length, about 1/100th of an inch being the average length: these cylinders at their upper ends suddenly contract into a point, more or less long, or are produced into a very fine tortuous tubulus of chitine, imbedded in the shell: I have seen in no other Cirripedes tubuli of this structure. There are other ordinary tubuli, such as occur in the valves of most Cirripedes, about 1/6000th of an inch in diameter, and which sometimes alternate with the above-described thicker cylinders. There are no external spines. From the number and length of the tubuli of both kinds, the tissue left after the action of acid is singularly complicated.
Ideas discussed in this chapter
Text from Project Gutenberg, public domain in the United States.