[83] The name Balanus was used, almost as at present, by Lister and Hill, before the introduction of the binomial system. Since that period the first two authors, as far as I know, who used this name, were Da Costa, in his ‘Hist. Nat. Test. Brit.,’ in 1778; and Bock, in the ‘Naturforscher,’ for the same year; Bock, however, applied it to a Chelonobia.
CONOPEA (pars generis). Say. Journal Nat. Sc. Philadelphia, vol. ii, part ii, 1822.
MESSULA (do.). Leach. Zoological Journal, vol. ii, 1825.
CHIRONA (do.). J. E. Gray. Philosoph. Transacts., 1835, p. 37.
Compartments six; basis calcareous or membranous; opercular valves sub-triangular.
Distribution.—Mundane: in the warmer seas.
General appearance.—The shape of the shell in the different species varies from depressed conical to cylindrical; the latter form being generally assumed when specimens are crowded together; but some species, as B. balanoides, crenatus, and lævis, seem more subject than others to be thus affected. The colour is either white, generally tinted by the yellowish or brownish epidermis, or any colour intermediate between bright pink and rich blue, purple being the prevailing tint. The persistence of the so-called epidermis is very different in different species, being even sometimes highly variable in the same species. The surface is either smooth or more commonly folded longitudinally, or sharply ribbed. The orifice differs in form from diamond-shape to trigonal; the carinal end, owing to the shape of the carina, being always sharper or narrower than the rostral end. The size of the orifice, in proportion to the shell, varies accordingly as the latter is more or less conical or cylindrical. The orifice is either entire or more or less deeply toothed, in proportion to the degree of obliquity of the summits of the radii and alæ. The radii almost always have smoother surfaces than the parietes. In some few species the radii are not developed, the sutures being marked only by fissure-like lines; in others they are very narrow, and in this case their upper margins are generally rounded and smooth, instead of being straight and jagged. The carino-lateral compartments are usually much narrower than the lateral compartments, occasionally in an extreme degree, as in B. allium. The shell is generally strong, sometimes to a wonderful degree; but the strength and thickness vary in the individuals of some of the species. By the action of hot caustic potash, the compartments in several species, such as B. Hameri and crenatus, separate on a touch; in others, they adhere so strongly as to prove that the sutures must be calcified together. In this genus we have the largest known sessile cirripede, viz., the B. psittacus, and on the other hand many small species; but it is very difficult, except in well-known species, to ascertain the average or even the maximum dimensions.
Scutum.—This valve is almost triangular, with the basi-tergal corner more or less rounded off. The prominent lines of growth are sometimes crossed by longitudinal striæ. Internally, the articular ridge projects to a very different degree in the different species; its lower end is sometimes (as in B. lævis, Pl. 4, fig. 2 c) produced downwards as a small, sharp, free style; there is always an articular furrow receiving the inflected margin of the tergum. There is always an impression left by the attachment of the adductor scutorum muscle; and often its lower side is bounded more or less closely by a sharp adductor ridge, running some way down the valve; this ridge is occasionally almost confluent, in its upper part, with the articular ridge, and in this case sometimes it forms, together with the inflected tergal margin, a large tubular cavity, running up, as in B. psittacus (Pl. 2, fig. 3 c), almost to the apex of the valve. Almost invariably there is a slight pit or depression for the lateral depressor muscle; sometimes within the depression there is a little ridge, as in B. perforatus and nubilus (Pl. 4, fig. 3 a, and Pl. 6, fig. 2 a); and in the case of B. vestitus, flosculus, and imperator (Pl. 8, figs. 3 a, 4 a), there are regular crests for this same purpose. The rostral depressor muscle is usually attached in a small pit formed by the folding over of the lower part of the occludent margin: in B. imperator (Pl. 8, fig. 4 a) there are regular crests for its attachment, and traces of them may be discovered in B. vestitus.
Tergum.—This valve is more nearly triangular than any other shape, with the spur more or less prominent. The apex generally projects a little above the level of the scutum; in some species it consists of a triangular and solid, in others (Pl. 2, fig. 3 b) of an almost cylindrical, extremely sharp, inwardly curved, and very prominent beak. This beak is generally purple; it is sometimes hollow, and occupied by a thread of corium. Its formation, and the apparent sliding up of the whole tergum, so as to project above the scutum, has been described under the family. From an account given to me by a person who kept B. porcatus alive, the beaks appear to be used, when the operculum is touched, as an organ of defence,—the animal striking with them. The tergal margin is more or less inflected; and the carinal margin is convex in different degrees, and, in some species, is added to by upturned zones of growth. The basal margin either forms a nearly straight line on opposite sides of the spur, or more commonly slopes towards it in various manners. The spur, or basal projection, is rarely placed in the middle of the basal margin, generally near, sometimes close to the basi-scutal angle; it varies much in length and breadth, and is sometimes even half the width of the valve. The surface of the valve is almost always more or less depressed, sometimes so much as to form a deepish furrow, the “longitudinal furrow,” which extends from the apex to the extremity of the spur. When the furrow is deep, its sides, as the specimen grows old, almost always become folded inwards, so as to touch, and then the furrow becomes converted into a closed fissure: in this latter case the folded sides generally form a central crest on the spur. Internally, in the middle of the upper part of the valve, the articular ridge is more or less prominent, forming the carinal margin of the articular furrow, in which the articular ridge of the scutum is lodged; occasionally, however, this articular ridge can hardly be said to exist. In most species the tergal depressor muscle is attached to sharp crests on the basi-carinal corner of the valve, but these are almost obliterated in other species.
Compartments.—The external appearance of the shell has already been described. In the most typical species, the parietes consist of an outer and inner lamina, separated by strong longitudinal septa; these septa are denticulated on both sides at their bases, but only close to the inner lamina; in fact the inner lamina is apparently formed by the union, thickening, and production, of some of the denticuli. As it is not the innermost of the denticuli on the basal edges of the longitudinal septa, which thus become united into a solid layer, the longitudinal septa form slightly projecting, longitudinal ribs on the inner lamina. These internal ribs are longitudinally striated; in old specimens they often become obliterated, especially in the upper part of the shell. The parietal tubes or pores (occupied by threads of corium) are generally square and large; but in B. Ajax they are very small, and in B. glandula often extremely minute. In the upper part of the shell, and sometimes low down, they are generally crossed by thin, transverse, calcareous septa: in some species, as in B. perforatus, and in some varieties of B. amphitrite, the upper ends of the tubes are filled up solidly with shell. In some varieties of B. crenatus and of amphitrite, the longitudinal septa, near the outer lamina, divide, thus giving rise to a very imperfect row of outer short tubes. In B. vinaceus (Pl. 2, fig. 7 d) the inner lamina is cancellated instead of being solid, which is caused by the basal denticuli of the longitudinal septa being simply united together by their ends and crossed by transverse septa, instead of being consolidated into a mass. In several species, as in B. Hameri, the walls consist only of the outer lamina with longitudinal ribs, no inner lamina having been formed; the ribs here evidently answer to the longitudinal septa in the foregoing species. In B. flosculus and imperator the walls are solid, their basal margins being formed of irregular, elongated points, and little ridges (Pl. 8, fig. 4 c), which apparently prefigure the more regular longitudinal ribs or septa. In B. balanoides the walls are generally either nearly smooth and solid, or irregularly cancellated; in B. cariosus (Pl. 7, fig. 3 b) two or three rows of short irregular tubes are formed by unequally branching septa, almost as in the genus Tetraclita.
The Radii, in all the species, are constructed essentially on the same plan as the parietes; thus, in the typical forms, there is an outer and inner lamina, with septa, which, near the inner lamina, are furnished with denticuli on both sides; hence the radii are permeated by pores or tubes, like the parietes; but this holds good only in the first section of the genus, for, in the other species, the tubes are filled up quite solidly. The denticuli on the septa often occur only on one side, or disappear altogether; and, lastly, the septa themselves often appear merely like little teeth, or disappear altogether as in B. Hameri, or occur only near the bases of the radii, as in B. amaryllis. A slight furrow in the compartment, against which each radius abuts, is generally marked by the septa and their denticuli. In regard to the alæ, their lateral or sutural edges are either thin and smooth, or, more commonly, finely crenated or ribbed. The little transverse crenations are homologous with the septa in the radii and parietes. The edges of the alæ are usually received in a furrow. The diametric growth of the shell is effected by the growth of the radii and alæ, and chiefly by that of the former. The sutural and lateral edges of both radii and alæ are added to, either quite up to their summits, or only low down, and during the continued growth of the shell, lower and lower down; in accordance with this difference in growth, the summits of the radii and alæ become either very oblique, or they extend parallel to the basis, that is, from tip to tip of the adjoining compartments. When the radii and alæ are added to, as is most usual, above the level of the opercular membrane, and therefore above the sack, ribbons of corium run up the sutures from the sack, higher or lower, according to the height to which, in the different species, the edges of the radii and alæ continue to be added to. The obliquity of the summits of the radii and alæ varies, in some cases, in the same species. It often happens that when the summits of the radii are very oblique, the summits of the alæ are but little so; and the converse; both, however, are often either equally oblique, or both have square summits. The sheath extends either one third or more than half down the shell; its basal margin often (Pl. 25, fig. 1, K′) freely depends or overhangs the inner lamina of the walls.
Basis.—In typical species the basis is calcareous, and consists of an upper and lower lamina, separated by radiating septa, forming pores. In the same manner as the septa of the parietes sometimes, though rarely, become irregularly divided near the outer lamina, forming outer pores, so it is, but in a much more marked degree, with the basis. The basis in such cases becomes extremely thick, and consists of an upper, thin lamina, with the regular radiating septa and pores, and of an underlying, thick, cancellated mass, which seems wholly to result from the dividing and sub-dividing of the septa. The basal radiating pores, like the parietal pores, are closed at intervals by calcareous transverse septa. The basal points of the parietal septa enter the orifices of the basal pores, and the threads of corium pass into the latter, between the denticuli of the parietal septa. In some species, as in B. crenatus and Hameri, the basis is perfectly solid, the upper lamina being absent, just as in some species, the internal lamina of the parietes is absent. In B. flosculus the basis is calcareous, but consists of so excessively thin a film as hardly to be distinguished: it presents, moreover, as also is the case with B. imperator, a beaded structure. Again, in some few species, as in B. balanoides, the basis is simply membranous. When the basis is thin, it is always flat, and is closely moulded to the irregular surface of attachment; and in this case, when specimens are crowded together, their elongation is effected exclusively by the growth of the walls; but, when the basis is thick, it sometimes becomes, in crowded groups, deeply, but irregularly, cup-formed, or cylindrical, as in B. psittacus and perforatus. In B. allium, however, which inhabits massive corals, the basis is as regularly concave or cup-formed as in the genus Pyrgoma: in B. calceolus and its allies, and in some varieties of the fossil B. inclusus, the basis is boat-formed, with its lower surface deeply grooved longitudinally from clasping the stem of the Gorgonia or other zoophyte, to which it was attached. In certain varieties of B. lævis it is very remarkable that the deeply cup-formed basis becomes, owing apparently to the whole shell having grown too deep for the animal, half-filled up with irregular, calcareous, transverse plates (Pl. 4, fig. 2 a), resting one upon the other by irregular points or pillars. The cementing apparatus has been sufficiently described under the Family.
Mouth.—The labrum is always notched; sometimes it has no teeth, but generally there are three on each side; in B. balanoides there are five or six on each side; and in B. improvisus and eburneus there is a whole row of teeth (Pl. 26, fig. 2, e′), graduated in size, on each side of the notch. The palpi are large, with their apices nearly touching, and furnished with long spines. The mandibles have, as it appears, normally, five teeth, but the two lower teeth are always small and often rudimentary, and almost confluent with the inferior, sometimes spinose angle. The maxillæ have either a simple edge, or a notch under the pair of large upper spines, or the lower part forms (Pl. 26, fig. 7) a step-formed projection: there are generally two lower spines, placed singly or not in pairs, larger than the others, with the exception of the uppermost pair. The outer maxillæ are, on their inner faces, obscurely divided into two lobes.
Cirri.—The rami of the first pair are unequal, the shorter one sometimes not being more than half the length of the other ramus: the segments of the shorter ramus are broad, and are, together with the lower segments of the longer ramus, thickly clothed with spines; in some species, as in B. perforatus, the anterior surfaces of the segments, more especially of the shorter ramus, and of both rami of the second pair are produced (Pl. 29, fig. 4), so as sometimes to form very remarkable projections. The segments of the second and third pairs are always thickly clothed with spines, as also are their pedicels. The third pair is rather longer than the second; but in B. vestitus and imperator it is much longer, and is otherwise somewhat different. The dorsal and basal margin of the pedicel of the third pair, in some of the species, as in B. tintinnabulum, is produced backwards on the thorax, and forms a membranous plate fringed with fine spines. The three posterior and longer pairs of cirri have from three to rarely eight or ten pairs of long spines on each segment, with generally one or two minute spines in the middle between each pair: their pedicels have a regular double row of spines.
The penis is long and hairy: in most of the species there is, at its dorsal basis, a small, sharp, flattened, imperforate projection; first observed by Poli: but this is sometimes absent, as in B. crenatus, though present in the closely allied B. balanoides; and its presence is variable in B. tintinnabulum. All the species have large plicated branchiæ. The base of the sack in several species is furnished with inwardly projecting filamentary appendages. In B. perforatus, crenatus, and improvisus, and I believe in other species, the upper part of the stomach is furnished with a circle of branching cæca.
On the variation of the species; their arrangement and affinities; value of the characters used; changes during growth.—Owing to the great variation in external characters, to which almost all the species are subject, and likewise to the genus being a very natural one, that is, to the species following each other in close and natural order, it is not easy to exaggerate the difficulty of identifying the species, except by a deliberate examination of the internal and external structure of each individual specimen. Every one who has collected sessile cirripedes must have perceived to what an extent their shape depends on their position and grouping. The surface of attachment has a great effect on that of the shell; for as the walls are added to at their bases, every portion has at one time been in close contact with the supporting surface; thus I have seen a strongly-ribbed species (B. porcatus) and a nearly smooth species (B. crenatus) closely resembling each other, and both having a peculiar appearance, owing to their having been attached to a pecten. Dr. Gray has pointed out to me specimens of B. patellaris, curiously pitted like the wood to which they had adhered; and numberless other instances might be added. Quite independently of the effect produced by the surface of attachment, the degree to which the longitudinal folds and ribs are developed on the parietes, is variable in most of the species, as in B. tintinnabulum, vestitus, and even in B. porcatus; the presence or entire absence of these ribs often surprisingly alters the whole aspect of the shell. The persistence of the so-called epidermis is in some degree variable; and in B. lævis we have groups of specimens absolutely naked, and others uniformly clothed with a brown membrane. Again, some species in certain localities are all subject to the disintegration of the entire outer lamina of the walls; and in such cases (as with B. perforatus) there is not the smallest resemblance between the corroded and perfect specimens. The size of the orifice, and consequently of the operculum, compared with the shell itself, varies accordingly as the shell is more or less conical or cylindrical; in the latter case, the summits of the radii are generally more oblique and the aperture consequently more deeply toothed than in the more conical varieties. Size is a serviceable character in some cases, but very many specimens are required to ascertain the average or maximum size of each species, for there is no method of distinguishing a half-grown from a full-grown specimen; and I believe, as long as the individual lives, so long does it go on moulting and growing. Colour is of very considerable service; though the precise tint varies greatly in almost every species; and what is a far more serious evil, the majority of the species have their white or nearly white varieties, the latter sometimes as numerous as the coloured ones: in B. perforatus, lævis, flosculus, amphitrite, and in several other species, the common white varieties are eminently deceptive.
Besides the slight variation in the obliquity of the summits of the radii and alæ, dependent on the more or less cylindrical form of the shell, in some species, as in B. tintinnabulum, amphitrite, improvisus, trigonus, and porcatus, their obliquity also varies occasionally from unknown causes, and thus greatly affects the general appearance of the shell. In some few species, as in B. perforatus, the radii are often either not at all developed, or are of very variable width; in others, when the shell has become cylindrical, or when very old, the radii cease to grow, and from the disintegration of the whole upper part of the shell, with the continued growth of the lower part, the radii at last come to exist as mere fissures: I have seen instances of this in B. psittacus, nigrescens, and porcatus. Nevertheless, the obliquity of the upper margin, and the breadth of the radii are useful characters; and still more useful is the fact whether the upper margins are smooth and arched, or straight and jagged. The fact of the terga being more or less beaked is useful: as is, likewise, the presence of striæ, or furrows, or rows of pits, radiating from the apices of the scuta; but to ascertain the presence of these marks, it is almost invariably necessary to take out the scuta, clean, and examine them with a lens; these ridges and furrows, moreover, in some species, as is strikingly the case with B. tintinnabulum, and in less degree with B. trigonus, lævis, and concavus, appear and disappear, and vary without any apparent cause.
Now if we reflect that form, size, state and nature of the surface, presence of epidermis, relative size of the orifice, presence of longitudinal ribs, tint, and often the existence of any colour, are all highly variable in most of the species; and that the obliquity of the summits of the radii, and the presence of longitudinal striæ on the scuta, are variable in several species; we shall perceive how difficult it must ever be to distinguish the species from external characters. As some evidence of this, I may mention that, after having described nearly 40 species, and when my eye was naturally able to appreciate small differences, I began carefully to examine varieties of B. tintinnabulum, amphitrite, improvisus, porcatus, vestitus, &c., without even a suspicion that they belonged to these species, at that time thoroughly well known to me; yet in the cases here referred to, there could be no doubt, when a perfect series was examined, that the specimens were only varieties. From this cause the labour of naming a collection is great. Let no one attempt to identify the species of this genus, without being prepared to disarticulate, clean, and carefully examine with a microscope the basis and parietes, and both the under and upper surfaces of the opercular valves; for I feel convinced, that he will otherwise throw away much labour. Moreover, in many cases, it is almost necessary, on account of the variability of the characters, to possess several specimens. From these facts, I have not hesitated to form my sections on characters which require close examination, though I would gladly have seized on external characters, could I have found such even moderately constant.
The least varying, and therefore most important characters, must be taken from the internal structure of the parietes, radii, and basis: not that these characters are absolutely invariable; thus the porosity of the parietes is slightly variable in B. glandula, and highly variable in the fossil B. unguiformis; it is also highly variable in B. balanoides, but under a systematic point of view this is unimportant, as the section including this latter species is well defined by the membranous basis. The porosity of the basis is in some degree variable in B. nubilus, improvisus, and patellaris; and in B. flosculus we see signs of a passage from a calcareous to a membranous basis. Characters derived from the general shape, and from the ridges and depressions on the under side of the scuta and terga, especially of the scuta, are highly serviceable. The terga, indeed, in many species, as in B. amphitrite, vary considerably, and are affected by the general shape of the shell. Unfortunately the differences are not very great between the scuta of the different species. The cause of the opercular valves offering more useful characters, as far as outline is concerned, than do the walls of the shell, is no doubt due to their being almost independent of any influence from the nature of the surface of attachment. Even the ridges and depressions on the under side of the scuta, which are in direct connection with the muscles and soft parts of the animal, vary to a certain extent: thus the length and prominence of the adductor ridge is decidedly variable in B. concavus and tintinnabulum, and in a less degree in B. lævis; the size and form of the little cavity for the lateral depressor muscle varies in many species; so does the exact shape and degree of prominence of the articular ridge. There is one character in the terga, which at first would be thought very useful, namely, whether an open longitudinal furrow, or a closed fissure runs down the valve from the apex to the spur; but it is found that the furrow almost always gradually closes up during growth; and as a consequence of this, the width of the spur compared to that of the whole valve, as well as its distance from the basi-scutal angle, and the form of its basal extremity, all vary in some degree. The length of the spur sometimes varies considerably, as in B. concavus and amphitrite. The parts of the mouth are only occasionally serviceable; for the teeth on the labrum, and the state of the lower teeth on the mandibles, and the presence of a step-formed projection at the lower angle of the maxillæ, are all often variable. The relative lengths of the two rami of the first pair of cirri, the degree of protuberance of the segments, and the number of pairs of spines on the segments of the posterior pairs of cirri, are sometimes useful; but the relative lengths of the cirri, and more especially the numbers of pairs of spines on the posterior cirri, are apt to vary. Finally, I must express my deliberate opinion, that every part and organ, internal and external, in Cirripedes, is liable to some amount of variation in some of the species.
I must now point out the principal changes which supervene during growth, and which cannot properly be called variations. In the first place, I think, it is scarcely possible to recognise a species when under the 1/10th of an inch in diameter. In some cases, as in var. d’Orbignii of B. tintinnabulum, the shell is invariably coloured when old, but quite white when very young. Generally the tints become very much darker with age. Some species, which usually or invariably have, when mature, longitudinally folded walls, as B. flosculus and balanoides, are perfectly smooth in youth. The walls in B. eburneus, when young, have white, hyaline, longitudinal lines, and are naked; whereas, with advancing age, these lines disappear, and the subsequently formed shell becomes covered with membrane. The summits of the radii are apt to be oblique in the young of B. Capensis, psittacus, and tintinnabulum, whereas they are generally quite square in old specimens. In B. eburneus, cariosus, and in a lesser degree in B. psittacus, the scuta become longitudinally striated only with age. On the other hand, in very young specimens of B. tintinnabulum, the scuta sometimes are deeply impressed by little pits placed in rows. I have already alluded to the longitudinal furrow on the tergum so entirely changing its character, owing to the edges becoming, during growth, folded inwards; this likewise causes a decrease in the proportional breadth of the spur. In old specimens of B. flosculus, var. sordidus, the whole tergum is much more elongated than in young specimens. The basal margin of the sheath is hollow beneath in the young of B. cariosus and of some other species, but in the old it is continuous with the inner surface of the walls. The inner lamina of the parietes generally loses, to a certain extent, its longitudinally ribbed character in old age. The basis is solid, instead of being porose, in very young specimens of B. improvisus. In all the species, the carino-lateral compartments, in early age, are very narrow in proportion to the width of the lateral compartments; and in all, at this early period, the operculum is large in proportion to the whole shell. The number of spines on the edge of the maxillæ, the number of segments in all the cirri, and the number of spines on each segment, are few in early youth, and go on increasing with each successive exuviation: the pedicels of the cirri are long in proportion to the rami at this same early age.[84]
[84] In some specimens of Balanus perforatus I made the following enumeration of the number of segments in the cirri:—
| Basal diameter of shell 1/20th of an inch. | Basal diameter of shell 1/5th of an inch. | Medium sized specimen about 3/4ths of an inch in basal diameter. | |
|---|---|---|---|
| First cirrus shorter ramus | ? | 11 | 17 |
| Second cirrus | 4 or 5 | 9 | 13 |
| Sixth cirrus | 9 or 10 | 19 | 31, in another 36 |
In the specimen 1/5th of an inch in basal diameter, each segment of the posterior cirri carried five pairs of spines; whereas, in full-grown specimens, there are six or seven pairs. In the 1/20th of an inch specimen, on the inner maxillæ, there were no spines between the upper large and the lower large pair of spines; whereas, in the 1/5th of an inch specimen, there were five intermediate spines, and in larger specimens nine or ten spines.
Notwithstanding the difficulties now enumerated, I hope that, owing to having examined a vast number of specimens of the most varying species, I have not fallen into very many errors. I have endeavoured to err on the side of making too few instead of too many species. In those cases, however, in which I have seen only a few specimens, I have been sometimes compelled to decide without sufficient evidence.
I would gladly have divided this genus, already including 45 species, into smaller genera; but so far from being enabled to do so, I have been compelled to form my Sections (immediately to be given) on characters not absolutely invariable, and far from obvious. I was particularly anxious to separate the elongated species with a boat-formed basis, which are attached to Gorgoniæ, and which form the genus Conopea of Say, but I was unable to effect their separation even as a sub-genus; for B. navicula and cymbiformis graduate in the most insensible manner through B. galeatus (the type of Say’s genus) and B. calceolus into B. stultus, and this into B. Ajax; yet this latter species has even been described as a mere variety of the typical B. tintinnabulum! Indeed, so insensible is this graduation, that the first and second sections of the genus are hardly distinct. I fully admit, that if B. stultus and Ajax had never existed, B. calceolus and its three allies might have formed as natural a little group, though difficult to be characterised, as does the sub-genus Acasta; or perhaps this group and Acasta might have been combined together. These same species, viz., B. calceolus and its allies, are intimately allied to B. terebratus and inclusus, which are contained in the last section (F) of the genus; and this shows that Dr. Gray’s proposed genus Chirona, including the species with non-porose parietes in sections (E) and (F), could hardly have been instituted, even if the porosity of the parietes had not been variable in B. unguiformis, balanoides, and glandula. My fourth section (D), founded on the basis not being porose, is perhaps the weakest of the divisions, as may be seen in the list of exceptions appended to the sectional headings.
The arrangement of the species is, I think, as natural as a linear one could be made: every one knows how irregularly and in how many directions the lines of affinity in every natural genus branch out. Some few species stand rather isolated, as B. declivis; and B. allium, cepa, and quadrivittatus in a little group by themselves. I have already shown how the species in the first and second sections (A and B) blend into each other; and that the blending species are likewise allied to some in the last section (F); furthermore, I shall have occasion to show that these same species can hardly be separated naturally from the sub-genus Acasta. The first section, moreover, passes into the third (C) by B. tulipiformis; and the third into the fourth (D) by B. improvisus, nubilus, corrugatus, and patellaris: the fifth and sixth (E and F) sections are closely connected together by B. cariosus and flosculus; and these two sections blend into the fourth (D) through B. unguiformis and balanoides, and lastly, into the third (C) section by B. dolosus and improvisus.
The genus, as we have just seen, is hardly separated from the sub-genus Acasta; by B. allium it tends to pass through the sub-genus Creusia into Pyrgoma; and by B. imperator and flosculus it graduates into Elminius and Tetraclita.
Geographical Distribution.—This, which is much the largest genus of sessile cirripedes, has its species scattered over the whole world, from the arctic regions, in lat. 74° 48′, where we have B. crenatus and porcatus, throughout the tropical seas, to Cape Horn, where B. flosculus adheres to the coast-rocks. Many of the species have individually very wide ranges; thus B. tintinnabulum and amphitrite are found throughout the warmer seas; but the wide distribution of these species may be partly due to their frequent adhesion to ships’ bottoms: B. crenatus ranges from the frozen seas, in lat. 74° 48′ north, to the West Indies and Cape of Good Hope—a wonderful endurance of the most opposite climates. Balanus improvisus, again, extends from Europe to Nova Scotia, thence southward to Patagonia, and up the western coast of S. America, someway north of the Equator. Most of the species have considerable ranges; thus of the six species found on the eastern shores of northern America, five of them occur in Great Britain. Of the thirty-six species of which the habitats are known, exactly one third, or twelve, inhabit both the torrid and temperate zones, these being divided by the isocryme of 68°; nine are found exclusively in the torrid, and fifteen exclusively in the temperate zones. Within the warmer latitudes, and especially in the southern hemisphere, Tetraclita and Elminius to a certain extent supplant Balanus. In depth, the species range from the upper limits of the tidal zone to even fifty fathoms. Balanus improvisus and eburneus are able to survive in brackish water. The different species are attached to various surfaces—rocks, shells, timber, floating objects, sea-weed, lamelliform corals, Milleporæ, Gorgoniæ, and even to sponges. Mr. G. B. Sowerby has remarked[85] that in the species from the southern hemisphere it is the basis, and in the species from the northern hemisphere it is the parietes, which are elongated, when the individuals, from being crowded together, become cylindrical; but this is erroneous; B. perforatus, in the northern hemisphere, sometimes has an elongated basis; but no doubt the basis of our commonest species, as B. balanoides, crenatus, and porcatus, from being either membranous or thin, does not become cup-shaped; whereas this structure is conspicuous in B. psittacus and lævis, the two commonest species in southern South America.
[85] Darwin’s ‘Geology of S. America,’ p, 264.
Fossil Species.—Having already given, under the Family, some account of the geological history of sessile cirripedes, short as it is, I here only allude to the subject in order to state my conviction that species cannot be satisfactorily distinguished in a fossil state, and rarely in a recent state, without an examination of the opercular valves. Nothing, indeed, could have been easier than to have affixed names to many groups of specimens, having different aspects, but to feel sure that these were really distinct species requires better evidence than can be afforded by the shell, without the operculum. No doubt, in some of the smaller sections of the genus—for instance, in that characterised by a membranous basis—it would have been possible to have distinguished some or several fossil species; but such have not as yet been found. When the specimens are much fossilised, it is, indeed, difficult to make out the primary points of structure—namely, whether the parietes, radii, and basis are porose: to do this it is sometimes necessary to rub down, polish, and carefully examine, a transverse section of a piece of the shell.
Parietes, and basis, and radii permeated by pores.
Parietes and basis sometimes permeated by pores, sometimes not; radii not permeated by pores; shell elongated in its rostro-carinal axis; basis boat-shaped, attached to Gorgoniæ and Milleporæ.
Parietes and basis permeated by pores; radii not permeated by pores.
Parietes permeated by pores. Basis and radii not permeated by pores.
Basis membranous.
Parietes and radii not permeated by pores. Basis sometimes permeated by pores, sometimes not, sometimes excessively thin and hardly distinguishable.[86]
[86] The following species might, owing to variation, or to the obscurity of their structure, without care, be classed in the wrong sections:—in A, Balanus Ajax, from living attached to Milleporæ, is sometimes elongated in its rostro-carinal axis, and from having its radii only finely porose, it might be classed in B: on the other hand, Bal. stultus is sometimes but little elongated, and the basis hardly boat-formed, and hence might be classed in A. The distinction between all the species in (B) and the sub-genus Acasta is artificial.
In sections C and D, I have seen one specimen of B. spongicola with a solid basis, and very young specimens of B. improvisus are thus characterised, and therefore these species are liable to be placed in the wrong section, D: Bal. nubilus, also, has part of its base non-porose, and therefore likewise might be placed in D: on the other hand, the circumference of the basis in B. patellaris is often porose, and hence this species, which belongs to D, might be placed in C.
In Bal. glandula, in D, the parietes of the compartments, without several were examined, might be thought to be solid, and therefore this species might be wrongly placed in F; on the other hand, the fossil B. unguiformis, in F, often has porose parietes, and such specimens would naturally be placed in D. Lastly, without care, B. flosculus might be thought to have a membranous basis, and so get placed in E.
The genus Pachylasma, without an examination of the animal’s body, might easily get misplaced in the wrong genus, amongst the species in the last section (F) of Balanus, yet there can be no doubt that Pachylasma belongs to the Chthamalinæ.