Part 58

_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.

_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.