Chapter 7

_Exuviation; Rate of Growth; Size._--I have had occasion repeatedly to allude to the exuviation of the Lepadidæ: with the exception of the genus Lithotrya,[20] in which the calcareous scales on the peduncle, together with the membrane connecting them, is cast off, neither the valves nor the membrane uniting them, nor that forming the peduncle with its scales and styles, are moulted; but the surface gradually disintegrates and is removed, perhaps sometimes in flakes, whilst new and larger layers are formed beneath. In Scalpellum, I ascertained that the new membrane, connecting together the newly-formed calcified rims under the valves of the capitulum, was formed as a fold, with the articulated spines which it bears, all adpressed in certain definite directions. This fold of new membrane, when the old membrane splits and yields, of course expands, and thus the size of the capitulum is increased. In the peduncle, lines of splitting can seldom be perceived, except, indeed, in the sub-globular, embedded, downward-growing peduncle of Anelasma, as described under that genus. I do not understand what determines the complicated lines of splitting of the old membrane between the several valves of the capitulum,--without it be simply, that along these lines alone, the old membrane is not strengthened by the new membrane being closely applied under it, the new being formed, as we have just said, in a fold, in order to allow of increase in size. Although, as I believe, there is strictly no exuviation in the outer membranes of mature Lepadidæ, it seems that narrow strips of membrane are cast off from between the valves, for the few first moults, after the final metamorphosis of the larva. I may here remark that, in most sessile Cirripedes, the outside membrane connecting the operculum and shell, is regularly moulted.

[20] The external integuments being moulted in Crustacea, but not in the Cirripedia, may appear, at first, an important difference: but we here see that non-exuviation is not universal amongst the Lepadidæ, and, on the other hand, according to M. Joly, (‘Annales des Sciences Naturelles,’ 2d series, Zoolog.), there is one true crustacean, the _Isaura cycladoides_, which has a persistent bivalve shell.

The delicate tunic lining the sack, (a mere duplicature of that thick one, forming the outside of the capitulum, and generally transformed into valves,) and the integuments of the whole body, are regularly moulted. With these integuments, the membrane lining the oesophagus, the rectum, and the deep olfactory pouches, and the horny apodemes of the maxillæ, are all cast together. I have seen a specimen of Lepas, in which, from some morbid adhesion, the old membrane lining one of the olfactory pouches had not been moulted, but remained projecting from the orifice as a brown shrivelled scroll. The new spines on the cirri (and on the maxillæ) are formed within the old ones; but as they have to be a little longer than the latter, and as they cannot enter these up to their very points, their basal portions are not thus included, but are formed, running obliquely across the segments of the cirri; and what is curious, these same basal portions are turned inside out, like the fingers of a glove when hastily drawn off. After the exuviation of the old spines, the new spines have their inverted basal portions drawn out from within the segments, and turned outside in, so as to assume their proper positions.

All Cirripedia grow rapidly: the yawl of H. M. S. _Beagle_ was lowered into the water, at the Galapagos Archipelago, on the 15th of September, and, after an interval of exactly thirty-three days, was hauled in: I found on her bottom, a specimen of _Conchoderma virgata_ with the capitulum and peduncle, each half an inch in length, and the former 7/20ths in width: this is half the size of the largest specimen I have seen of this species: several other individuals, not half the size of the above, contained numerous ova in their lamellæ, ready to burst forth. Supposing the larva of the largest specimen became attached the first day the boat was put into the water, we have the metamorphosis, an increase of length from about .05, the size of the larva, to an whole inch, and the laying of probably several sets of eggs, all effected in thirty-three days. From this rapid growth, repeated exuviations must be requisite. Mr. W. Thompson, of Belfast, kept twenty specimens of _Balanus balanoides_, a form of much slower growth, alive, and on the twelfth day he found the twenty-first integument, showing that all had moulted once, and one individual twice within this period. I may here add, that the pedunculated Cirripedes never attain so large a bulk as the sessile; _Lepas anatifera_ is sometimes sixteen inches in length, but of this, the far greater portion consists of the peduncle. _Pollicipes mitella_ is the most massive kind; I have seen a specimen with a capitulum 2.3 of an inch in width.

_Affinities._--Considering the close affinity between the several genera, there are, I conceive, no grounds for dividing the Lepadidæ into sub-families, as has been proposed by some authors, who have trusted exclusively to external characters. In establishing the eleven genera in the Lepadidæ, no one part or set of organs affords sufficient diagnostic characters: the number of the valves is the most obvious, and one of the most useful characters, but it fails when the valves are nearly rudimentary, and when they are numerous: the direction of their lines of growth is more important, and fails to be characteristic only in Scalpellum: with the same exception, the presence or abscence of calcified or horny scales on the peduncle is a good generic character. For this same end, the shape of the scuta and carina, but not of the other valves, comes into play. In three genera, the presence of filamentary appendages on the animal’s body is generic; in Pollicipes, however, they are found only on three out of the six species. The number of teeth in the mandibles, and the shape of the maxillæ, often prove serviceable for this end; as does more generally the presence of caudal appendages, and whether they be naked or spinose, uniarticulate or multiarticulate; in Pollicipes alone this part is variable, being uni-and multi-articulate; and in one species of Scalpellum they are absent, though present in all the others. The shape of the body, the absence or presence of teeth on the labrum, the inner edge of the outer maxillæ being notched or straight, the prominence of the olfactory orifices, the arrangement of the spines on the cirri, and the number and form of their segments, are only of specific value.

Comparing the pedunculated and sessile Cirripedes, it is, I think, impossible to assign them a higher rank than that of Families. The chief difference between them consists, in the Lepadidæ, in the presence of three layers of striæ-less muscles, longitudinal, transverse and oblique, continuously surrounding the peduncle, but not specially attached to the scuta and terga; and on the other hand, in the Balanidæ, of five longitudinal bundles of voluntary muscles, with transverse striæ, fixed to the scuta and terga, and giving them powers of independent movement. In the Lepadidæ, the lower valves, or when such are absent, the membranous walls of the capitulum, move with the scuta and terga when opened or shut; and the lower part of the capitulum is separated by a moveable peduncle from the surface of attachment; in the sessile Cirripedes, the lower valves are firmly united together into an immovable ring, fixed immovably on the surface of attachment. I will not compare the softer parts, such as the cirri and trophi, of the Lepadidæ with those of the Balanidæ, as my examination of this latter family is not fully completed: I will only remark, that there is a very close general resemblance, more especially with the sub-family Chthamalinæ.

_Geographical Range; Habitats._--The Pedunculated Cirripedes extend over the whole world; and most of the individual species have large ranges, more especially, as might have been expected, those attached to floating objects; excepting these latter, the greater number inhabit the warmer temperate, and tropical seas. Of those attached to fixed objects, or to littoral animals, it is rare to find more than three or four species in the same locality. On the shores of Europe I know of only three, viz., a Scalpellum, Pollicipes, and Alepas. At Madeira (owing to the admirable researches of the Rev. R. T. Lowe), two Pæcilasmas, a Dichelaspis, and an Oxynaspis are known. In New Zealand, there are two Pollicipes and an Alepas, and, perhaps, a fourth form. From the Philippine Archipelago, in the great collection made by Mr. Cuming, there are a Pæcilasma, an Ibla, a Scalpellum, Pollicipes, and Lithotrya. Of all the Lepadidæ, nearly half are attached to floating objects, or to animals which are able to change their positions; the other half are generally attached to fixed organic or inorganic bodies, and more frequently to the former than to the latter. Most of the species of Scalpellum are inhabitants of deep water; on the other hand, most of Pollicipes,[21] of Ibla, and Lithotrya are littoral forms. The species of Lithotrya have the power of excavating burrows in calcareous rocks, shells, and corals; and the singular manner in which this is effected, is described under that genus. Anelasma has its sub-globular peduncle deeply embedded in the flesh of Northern Sharks; and I have seen instances of the basal end of the peduncle of _Conchoderma aurita_, being sunk into the skin of Cetacea; in the same way the point of the peduncle in the male of Ibla, is generally deeply embedded in the sack of the female. I believe in all these cases, the cementing substance affects and injures the corium or true skin of the animal on which the creature is parasitic, whilst the surrounding parts, being not injured, continue to grow upwards, thus causing the partial embedment of the Cirripede. In the case of Anelasma, we have growth at the end of the peduncle, and consequently downward pressure, and this may possibly cause absorption to take place in the skin of the shark at the spot pressed on.

[21] I am informed by Mr. L. Reeve that _Pollicipes mitella_ is eaten on the coast of China; and Ellis states (‘Phil. Trans.,’ 1758) that this is the case with _P. cornucopia_ on the shores of Brittany. It is well known that the gigantic _Balanus psittacus_ on the Chilian coast, is sought after as a delicacy; and I am assured, by Mr. Cuming, that it deserves its reputation.

_Geological History._--Having treated this subject at length, in the volume of the Palæontographical Society for 1851, I will not here enter on it: I will only remark, that the Lepadidæ or Pedunculated Cirripedes are much more ancient, according to our present state of knowledge, than the Balanidæ. The former seem to have been at their culminant point during the Cretaceous Period, when many species of Scalpellum and Pollicipes, and a singular new genus, Loricula, existed; Pollicipes is the oldest genus, having been found in the Lower Oolite, and, perhaps, even in the Lias. The fossil species do not appear to have differed widely from existing forms.

_Genus_--LEPAS. Plate I.

LEPAS. _Linnæus._[22] Systema Naturæ, 1767.

ANATIFA. _Brugière._[23] Encyclop. Method. (des Vers), 1789.

ANATIFERA. (_Lister_) et plerumque Auctorum Anglicorum.

PENTALASMIS. (_Hill._) _Leach._ Journal de Physique, July, 1817.

PENTALEPAS. _De Blainville._ Dict. des Sci. Nat., 1824.

DOSIMA. _J. E. Gray._ Annals of Philosophy, vol. x, 1825.

[22] Linnæus, as is well known, included under this genus both the pedunculated and sessile Cirripedes. According to the rules of the British Association, the name Lepas must be retained for part of the genus; and as the sessile division was named Balanus, by Lister and Hill, even before the invention of the binomial system, and subsequently, in 1778, by Da Costa, and again, in 1789, by Brugière, there can be no question that Lepas must be applied to the pedunculated section of the genus. In this instance it is particularly desirable to recur to the Linnean name, as no other name has been _generally_ adopted. Had not Lister and Sir J. Hill published before the binomial system, their names of Anatifera and Pentalasmis would have had prior claims to Lepas.

[23] The date of this publication is almost universally given as 1792, apparently caused by an error in the title-page of the First Part, which has consequently been cancelled. The First Part contains Anatifa and Balanus, and was published in 1789. The Second Part was published in 1792, and has a corrected title-page for the whole _volume_.

_Valvæ 5, approximatæ: carina sursùm inter terga extensa, deorsùm aut furcâ infossâ aut disco externo terminata: scuta subtriangula, umbonibus ad angulum rostralem positis._

Valves 5, approximate: carina extending up between the terga, terminating downwards in an embedded fork, or in an external disc: scuta sub-triangular, with their umbones at the rostral angle.

Filaments seated beneath the basal articulation of the first cirri; mandibles with five teeth; maxillæ step-formed; caudal appendages uniarticulate, smooth.

_Distribution._--Mundane; attached to floating objects.

_Description._--Capitulum flattened, sub-triangular, composed of five approximate valves. The valves are either moderately thick and translucent, or very thin and transparent; and hence, though themselves colourless, they are often coloured by the underlying corium. Their surfaces are either smooth and polished, or striated, or furrowed, and sometimes pectinated. They are not subject to disintegration; they are generally naked, except on the borders, where they are coated, and held together by membrane; in _L. fascicularis_, however, the valves are covered with thin membrane, bearing very minute spines. The manner of growth of the valves will be best described under each. All the valves, even in the same species, are subject to considerable variation in shape, more especially the terga.

_Scuta._--These valves are sub-triangular in outline, with the basal margin straight and rather short; and with occludent and tergo-carinal margins more or less protuberant; in _L. fascicularis_, however, the basal (Pl. I, fig. 6), and occludent margins are slightly reflexed and prominent. A ridge, generally runs from the umbo to the upper point. Internally, there is no conspicuous pit for the adductor muscle; under the umbones, there is generally either on both valves, or only on the right-hand side (Pl. I, fig. 1 _c_), a small calcareous projection or tooth, of variable size and shape, even in the same species; it is generally largest on the right-hand valve; these teeth at first sight appear to form a hinge, uniting the opposite scuta at their umbones, but this is not really the case, and their use appears to be only to give attachment to the membrane uniting the valves together, and to the peduncle. The basal margin is internally strengthened by a calcified rim, more or less developed. The umbones (and primordial valves when distinguishable,) are seated at the rostral angles; during growth the basal margin is not added to, and the occludent margin only to small extent; hence the main growth of the valve is at the upper end, and along the carina-tergal margin. In _L. fascicularis_, however, the basal reflexed margin is slightly added to beneath the umbo.

_Terga_,--flat, small compared with the scuta, usually of an irregular quadrilateral figure, with the two upper or occludent margins very short, in proportion to the two (carinal and scutal) lower margins; all the margins are nearly straight. The two occludent margins, generally meet each other at about right angles, forming a small triangular projection; in _L. fascicularis_, however, the occludent margin is formed by a single, slightly curved line. The umbones (and primordial valves when distinguishable) are not seated at the uppermost point, but at the angle where the carinal margin unites to the upper of the two occludent margins: during growth the terga are added to, both on the occludent and on the scutal margins, and slightly along the carinal margin; hence their growth is unequally _quaqua-versal_, except at one angle of the irregular quadrilateral figure.

_Carina._--This is always very narrow and curved, concave within, often carinated and barbed exteriorly; it extends upwards between the terga for one half or two thirds of their length: at the lower extremity it ends (with the exception of _L. fascicularis_), in a small fork (Pl. I, fig. 1, _a_, _b_) rectangularly inflected and embedded in the membrane, beneath the basal margin of the scuta. From comparing this lower part of the carina in _L. australis_ (fig. 5 _a_), with the same part in some of the species of the allied genus Pæcilasma, it would appear that the fork is formed by an oblong disc, more and more notched at the end, and with the rim between the two points more or less folded backwards: conformably with this view, in very young specimens of _L. australis_, instead of a large and sharp fork, there is a small disc. The only use of the fork appears to be to give firm attachment to the membrane uniting the valves and peduncle. In _L. fascicularis_, instead of a fork, there is a broad, oblong disc (figs. 6, 6 _a_), rectangularly inflected; it is much longer than the fork, in proportion to the upper part of the carina; the disc is not more deeply embedded than the upper part. The umbo (and primordial valve when distinguishable,) of the carina is seated just above the embedded fork (or disc in _L. fascicularis_), at the point where the inflection takes place; hence the main growth of the carina is upwards,--the fork, however, being of course, likewise added to at its point: in _L. fascicularis_, the growth is both upwards and downwards.

_Peduncle and Attachment._--The peduncle is generally quite smooth: though with a high power its surface may be seen to be studded with minute beads, or larger discs, of yellowish and hard chitine; in the young of _L. australis_, and I suspect of some other species, it is covered with very minute spines. The peduncle in this genus attains its greatest development. The cement-tissue debouches, I believe, only through the functionless larval antennæ, except in one species, _L. fascicularis_, in which a ball of this substance is formed in a most peculiar manner round the peduncle (Pl. I, fig. 6), apparently for the purpose of serving as a float, as will be presently described.

_Size and Colour._--The species of this genus are the largest of the Pedunculata, with the exception of some Pollicipes: even in the smallest species (_L. pectinata_), the capitulum sometimes attains a length of about half an inch. The peduncle varies much in length in the same species: in _L. anatifera_, it is occasionally above a foot long. The colours of _L. anatifera_, _L. Hillii_, and _L. anserifera_, are very bright and striking; the membrane bordering the valves and that round the top of peduncle in two of the species, is of the brightest scarlet-orange; the valves, owing to the underlying corium, are pale blueish-grey, and the interspaces between them dark leaden-purple. The cirri and trophi are generally dark purple or lead-colour.

_Filamentary Appendages._--These are attached to beneath the basal articulation of first pair of cirri; they vary in the several species, from one to five or six on each side, the lowest being always the longest. Several of them are occupied by testes. In _L. pectinata_, generally, not even one is developed. They are subject to great variation in their proportional lengths, and in number, in the same species. These organs have generally been considered to serve as branchiæ; I see no reason to believe that they are more especially designed for this end, than is the general surface of the body.

_Mouth._--The labrum is moderately bullate, the longitudinal diameter of this part equalling about one third, or half of that of the rest of the mouth. The palpi are moderately developed. The mandibles (Pl. X, fig. 5) have five teeth with the inferior point either broad, or very narrow and tooth-like. The maxillæ are step-formed (Pl. X, fig. 9); the first step is sometimes indistinct and curved; and in _L. pectinata_, all the steps vary much, and are more or less blended together. The outer maxillæ (like those at Pl. X, fig. 16), are internally clothed continuously with spines. The olfactory orifices are not at all prominent.

_Cirri._--The first pair is placed near the second pair, and is of considerable length; the second has the anterior ramus thicker than the posterior ramus, and the segments brush-like; the segments (Pl. X, fig. 26) of the four posterior cirri bear from four to six pair of long spines, with a row of small intermediate spines: in the posterior cirri of _L. australis_ the lateral rim spines are much developed; and in those of _L. fascicularis_, the usual pairs of large spines are lost in a broad triangular brush, formed by the increase of the lateral marginal, and intermediate spines.

_Caudal Appendages_ (Pl. X, fig. 18 _b_), very small, either blunt or pointed, and quite destitute of spines.

The prosoma is well developed. The stomach is surrounded in the upper part by a circle of large branching cæca. The generative system is highly developed; the testes coating the whole of the stomach, entering the filamentary appendages and the pedicels of the cirri; the two ovigerous lamellæ contain a vast number of ova; they are united to rather large fræna, of which the sinuous margin supports either a continuous row or separate tufts of glands.

_Distribution._--The species abound over the arctic, temperate and tropical parts of the Atlantic, Indian and Pacific Oceans, and are always, or nearly always, attached to floating objects, dead or alive. The same species have enormous ranges; in proof of which I may mention that of the six known species, five are found nearly all over the world, including the British coast; and the one not found on our shores, the _L. australis_, apparently inhabits the whole circumference of the southern ocean.

_General Remarks and Affinities._--The first five species form a most natural genus; they are often sufficiently difficult to be distinguished, owing to their great variability. The sixth species (_L. fascicularis_) differs to a slight extent in many respects from the other species, and has considerable claims to be generically separated, as has been proposed by Mr. Gray, under the name of Dosima; but as it is identical in structure in all the more essential parts, I have not thought fit to separate it. As far as external characters go, some of the species of Pæcilasma have not stronger claims, than has _L. fascicularis_, to be generically separated; and I at first retained them altogether, but in drawing up this generic description, I found scarcely a single observation applicable to both halves of the genus; hence I was led to separate Lepas and Pæcilasma. If I had retained these two genera together, I should have had, also, to include the species of Dichelaspis and Oxynaspis; and even Scalpellum would have been separable only by the number of its valves; this would obviously have been highly inconvenient. Although some of the species of Pæcilasma so closely resemble externally the species of Lepas, yet if we consider their entire structure, we shall find that they are sufficiently distinct; as indirect evidence of this, I may remark that Conchoderma (as defined in this volume), includes two genera of most authors, and yet certainly comes, if judged by its whole organisation, nearer to Lepas than does Pæcilasma.

1. LEPAS ANATIFERA. Tab. I. fig. 1. (_var._)