A Monograph on the Sub-class Cirripedia · Charles Darwin
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_Affinities._--This species in general appearance closely approaches _B. crenatus_ and _balanoides_, and it is related to them in many essential parts, such as in the opercular valves. It agrees with _B. balanoides_, and differs from _B. crenatus_, in the smallness and imperfection of the parietal pores, and in the radii having rounded summits; it agrees with _B. crenatus_ in the structure of its basis, and in the prominent longitudinal ribs on the internal surface of the parietes, and differs from that species in the spur of the tergum being squarer, and in the scutum having an adductor ridge. _Range._--From the appearance of the Californian specimens, I suspect that they had adhered to tidal shells and to wood. The specimens in the British Museum, adhering to _Pollicipes polymerus_, consist of two lots, one of unknown origin, and the other certainly brought from the southern half of the Pacific Ocean by Sir James Ross: it deserves notice, that the _Pollicipes polymerus_, the supporting object, ranges from California to the southern Pacific Ocean. _Section_ E. _Basis membranous._ 29. BALANUS BALANOIDES. Pl. 7, fig. 2 _a_-2 _d_. LEPAS BALANOIDES. _Linn._ Fauna Succica, 1746, et Syst. Naturæ, 1767. ---- ---- _O. Fabricius._ Fauna Groen., p. 424, 1780. ---- ---- ET CLIVATUS. _Montagu_ (!). Test. Brit., 1803. BALANUS VULGARIS (?) _Da Costa._ Hist. Nat. Testacea, Pl. 17, fig. 7, 1778.
_General Appearance._--The shell, in middle-sized and old specimens, is almost invariably folded longitudinally and irregularly; it is either dirty white or very often pale brown, and punctured from the outer lamina having been corroded, to which action it is extremely subject. In very young specimens, the surface is usually quite white and smooth. The shell is sometimes much depressed; generally conical, but when crowded together, cylindrical or club-shaped, one specimen being even more than five-and-a-half times as long as wide. In Mr. Jeffreys' collection there is a specimen 2.5 of an inch long, .45 in diameter at the summit, only .2 in the middle, and rather more than .2 near the base. Another specimen was 1.8 in length, its greatest diameter being .35 of an inch at the summit. On the other hand, I have seen a very depressed variety, with deeply folded walls, in Mr. Thompson's collection from near Dublin, which was no less than four times as wide as high; so that the difference in proportion of height and greatest width, in the two extreme specimens, was nearly as 10 to 1. Occasionally, from some unknown cause, isolated specimens become cylindrical. The orifice of the shell, in the much elongated specimens, is generally deeply toothed. The radii are always narrow, sometimes extremely narrow, and have their summits smooth and rounded.
English specimens do not usually attain half an inch in basal diameter; I have, however, seen one from near Yarmouth .9 of an inch in diameter. Specimens from Massachussetts seem rather larger than the average size of British specimens, many being .6 of an inch, and one specimen a whole inch in basal diameter. The _opercular_ valves so closely resemble those of _B. crenatus_, that the description is necessarily comparative; in some cases they could hardly be discriminated; generally, owing to the disintegration to which this species is subject, the tips of the scuta are worn off, and hence the articular ridges together form (Pl. 7, fig. 2 _a_) a square projection, indenting the two terga; but I have examined young specimens and others when not disintegrated, in which the opercular valves, viewed externally, presented no difference whatever from those of _B. crenatus_. The _scuta_, however, are, I think, generally rather thicker, with the growth-ridges more prominent, and with the tips certainly less reflexed than is usual with _B. crenatus_. Internally, the articular ridge is rather less prominent: there is no distinct adductor ridge. The _terga_ are often rather narrower in proportion, and this especially holds good in the elongated varieties; in these latter, there is occasionally a moderately deep longitudinal furrow: the spur is often exactly the same shape as in _B. crenatus_, but it is apt to be rather longer (Pl. 7, fig. 2 _c_) and more pointed: in _var._ (_a_) it is pointed (fig. 2 _d_) in a very remarkable manner. Internally, the articular ridge is decidedly more prominent than in _B. crenatus_; the crests for the tergal depressor muscles are either well developed or almost absent. From this description it will be seen, how singularly the opercular valves of the common varieties of these two species resemble each other. I may mention that in some of the much elongated specimens, the muscles going to the opercular valves partially lose their transverse striæ, and become ligamentous.
The _Parietes_ are either quite solid, or more commonly are permeated by minute pores, or by small irregular square tubes (Pl. 7, fig. 2 _b_), which only run up each successive zone of growth, for very short distances, giving to the shell a cancellated structure, which from corrosion is often externally visible. In the rather rare variety (_a_) the parietes are permeated by regular tubes, extending up to the apices of the compartments, but crossed by transverse septa. The longitudinal septa, when such can be said to occur, in no case are denticulated at their bases. The internal surface of the parietes is either quite smooth or is traversed (Pl. 7, fig. 2 _b_) by very slight anastomosing ridges, but never, even in _var._ (_a_), by regular longitudinal ribs, as in most other species. The carinal margin of each compartment, on the inside, projects, as in _B. crenatus_, inwards, beyond the general surface of the shell, and running down, rests on the basal membrane. The lower edge of the sheath is rarely hollow beneath. The walls are lined by purplish, or pale brown, or sometimes by almost black corium; numerous tubuli penetrate the under sides of the walls and opercular valves; and it is the intersection of these tubuli that gives the punctured appearance to the often corroded surface of the shell. The _radii_ are narrow, generally very narrow; they have their upper and outer margins, as seen externally, very oblique, rounded and (when well preserved) smooth; their sutural edges are either quite smooth, or sometimes just perceptibly pitted, like the basal margin of the walls, or occasionally furnished with globular or arborescent little ridges.
The _alæ_ are also very oblique, but to a variable degree, sometimes only slightly oblique: their sutural edges are either smooth or obscurely crenated. _Basis_, membranous; in some much elongated specimens, during continued growth, the basal edges of the compartments approach each other so closely as almost to touch, so that the whole shell becomes pointed at the bottom; but on careful inspection I have never failed to find, even in the most pointed specimens, a minute basal membrane; in other much elongated varieties, in which the shell has apparently become too large for the animal's body, the basal membrane, instead of being flat, becomes drawn up deeply inwards, so as to touch the surface of attachment only close round the basal edges of the shell.
_Mouth_: labrum with the teeth on each side of the central notch unusually variable in number; I have seen specimens with only two on each side, with four on each side, with five on one side and four on the other, with five on one side and none on the other, and with six on both sides; hence the total number ranges from four to twelve. Mandibles, with the fourth and fifth teeth small, or quite rudimentary. Maxillæ, with scarcely even a trace of a notch under the upper pair of spines. _Cirri_; first pair, with one ramus one third or one fourth longer than the other; in one specimen the number of segments were nine and sixteen in the two rami: second and third cirri short, very nearly equal in length, having in the just-mentioned specimen respectively ten and eleven segments; the sixth cirrus in this same specimen had twenty-five segments, each segment being about as long as broad, and supporting six pairs of spines. In the singular variety (_a_) the posterior cirri are more elongated, and each segment supports seven or eight, and in one case even ten pairs of spines! the third pair is also in this variety proportionally rather longer. At the base of the third pair there is a tuft of fine spines. The penis has not, as in _B. crenatus_, a point at its dorsal basis. The branchiæ are very little plicated.
_Varieties._ Of the varieties having much elongated, club-shaped, hour-glass shaped, and depressed shells, there is no necessity to say anything in particular. With respect to the remarkable variety (_a_), I at first named and described it as a distinct species: I have received two lots, both from North America, one being sent me by Professor Agassiz from Cape Cod. These agreed in having the parietes permeated by regular tubes; in having the spur of the tergum most sharply pointed; in the third pair of cirri being proportionally longer compared with the second pair; in the sixth pair having more numerous segments, namely, three times as many as in the third pair; in the segments of the posterior cirri being more elongated, and especially in the number of pairs of spines on each segment--amounting in one case even to ten, a number unparalleled in other cirripedes. It may naturally be asked why I have not retained so well marked a form as a distinct species? In the first place, I found the most remarkable character in _var._ (_a_), namely, the number of pairs of spines on the posterior cirri variable, there being in one lot seven or eight pairs, and in the other lot nine or ten pairs on each segment.
Secondly, all the characters by which this variety differs from the common _B. balanoides_, are those which are variable in the latter; this is especially the case with the structure of the parietes, and in a lesser degree with the spur of the tergum. Thirdly, I found a specimen in Mr. Cuming's collection, from Sweden (so that this _var._ (_a_) is not confined to North America), in which the cirri quite resembled those of the American specimens, but the spur of the tergum was in an intermediate condition as compared to that of ordinary varieties; and the parietal tubes were of unequal sizes, and scarcely more regular than sometimes in the true _B. balanoides_. And lastly, I have seen specimens from Ayrshire, with the parietes permeated by regular tubes, but with the tergum in an intermediate condition, and with the segments of the posterior cirri not more numerous or more elongated than in _B. balanoides_, supporting only six or seven pairs of spines, that is only one more than is common with _B. balanoides_; so that it was impossible to decide whether to rank the Ayrshire specimen under _var._ (_a_) or under the common form, so that I was compelled to give up _var._ (_a_) as a species.
_Monstrous individuals, with the male organs aborted: Parasite._--Amongst some specimens, chiefly elongated ones, sent to me from Tenby, in South Wales, I found no less than seven individuals with some of the posterior cirri distorted, unequal on the opposite sides, and in an almost rudimentary condition, and in each case with the penis truncated, without any muscle entering the stump, which was _absolutely imperforate_: the vesiculæ seminales were much shrunk; in one case without any zoosperms; in another case with headless zoosperms cohering in an unusual manner; hence it is certain that these individuals were functionally only female, and could not impregnate their own ova; yet in two instances the ova had been impregnated, no doubt by neighbouring perfect individuals, for they contained well-developed larvæ. Several of these monstrous individuals were infested by one, two, or three curious crustaceans, which have been described by Mr. Goodsir,[97] as the male of the Balanus; but these supposed males are females, and were distended with ova containing almost mature larvæ; I believe that they are the females of the unnamed genus, belonging to the family of Ioniens, described by Mr. Goodsir, which live parasitic within the sack (as I likewise found) of the same individual Balani. [97] 'Edinburgh New Philosophical Journal,' July, 1843.
_Diagnosis._--I have seen several specimens of this species and of _B. crenatus_, absolutely undistinguishable in external appearance. I may specify one of _B. balanoides_, imbedded in an alcyonidium, and one of _B. crenatus_, imbedded in a sponge, and therefore neither at all abraded. Generally, the tips of the scuta in _B. crenatus_ are a little reflexed, whereas in _B. balanoides_, when the shell has been at all disintegrated, the tips form a square projection locked into the terga. _Bal. crenatus_ never assumes the punctured appearance so common in _B. balanoides_. Very young specimens of the latter can be distinguished by their dead white colour and smoothness. The edges of the radii are almost always smoother than in _B. crenatus_, and they are never so wide as is sometimes the case with _B. crenatus_. When a specimen is disarticulated, our present species can at once be distinguished from _B. crenatus_ (and from _B. improvisus_), by its membranous basis, and by the solid or cancellated walls, which are rarely permeated by regular tubes or pores; and the walls when porose are not internally ribbed. I have already pointed out the few very trifling points, in which the opercula of the two species differ. The mouth and cirri offer likewise very few differences: in _B. balanoides_ there are often more teeth on the labrum than in _B. crenatus_; the rami of the first cirri are perhaps here rather less unequal; the second and third pairs of cirri are certainly in most cases more equal in length; and lastly, the segments of the sixth cirri, even in the common varieties, bear, in equal-sized specimens, more pairs of spines than in _B. crenatus_. We shall see that in habits, with regard to depth, the two species differ, _B. balanoides_ inhabiting much shallower water than _B. crenatus_.
_Range, Habitats, &c._--This species is extraordinarily abundant within the tidal limits round the shores of Great Britain, and apparently of the northern United States. Besides numerous specimens sent to me from very many English localities, the late Mr. W. Thompson, of Belfast, kindly placed in my hands his very large collection; from these materials it appears that _B. balanoides_ is the only tidal species in the northern parts of our island; but in the south and south-west, it is associated with the _Chthamalus stellatus_ and _Balanus perforatus_. I doubt whether this species ever lives below the lowest tides; the case of a few specimens being mingled with _B. improvisus_ and _crenatus_, (mentioned under the latter species,) at the bottom of a rudder of a small vessel, about six feet deep, is hardly an exception, for the water would there be troubled and aërated almost as in a breaker; and on this very rudder the upper two or three feet were coated exclusively by the _B. balanoides_. This species lives on rocks at both the uppermost and lowest limit of the tides; I am informed by Mr. Thompson, that he has seen specimens attached to a spot not covered by water during neap-tides. As a proof of its tenacity of life, Mr. Thompson informs me that he accidentally kept some specimens in a box, in a warm sitting-room, and found them alive seven days afterwards. This same most accurate observer finds, however, that _B. balanoides_ is very susceptible to brackish water; he says, "that having kept some specimens alive for a week in excellent health, the water being changed once in thirty-six hours, they were one day killed instantly by some water, though brought from the same part of the estuary as usual, having been rendered brackish by much rain having lately fallen." I may recall the fact, that _B. improvisus_ lives daily for hours in absolutely fresh running water.
The _B. balanoides_ lives attached, often continuously coating many square feet of the surface, to rocks, pebbles, wooden-piers, littoral shells and ulvæ. The most northern point whence I have received specimens, is in lat. 66° 34′ in North America, collected by Mr. Sutherland; and the most southern point is Delaware Bay, in the United States, in lat. 39°; I do not believe that this species extends into the Mediterranean, for Ranzani (Mem. di Storia Nat.), who particularly attended to the nature of the basis, was not acquainted with any _Balanus_ having a membranous basis; and Poli (Test. Ut. Siciliæ,) describes only two species thus characterised, and these are manifestly _Chthamali_. With respect to the rate of growth of this species, I am indebted to Mr. W. Thompson for the following note:-- "Sept. 29, 1848.--I examined a great number of Balani, in reference to the growth made by them during the present season, and found it to average three lines in diameter, and at most four lines. I saw a few minute specimens, only one line in diameter, showing that the species continued to breed until lately: these latter were probably not more than four weeks old. The young of the present year are plainly distinguished from the older ones, by their pure white colour and fresh appearance. Judging from the size of this year's specimens, and of the older ones on the same stones, I am of opinion that the term of life of the species is two years. Of the older shells, which I examined and found living in the spring, nine tenths are now dead, the walls only remaining, the opercular valves having been washed away."
Mr. Thompson goes on to say, that the individuals which had, on July 3, a basal diameter of from two and a half to three lines, had attained, by the 30th of September, a diameter of four and a half lines, this being here the maximum size of the species. 30. BALANUS CARIOSUS. Pl. 7, fig. 3 _a_-3 _e_. LEPAS CARIOSA. _Pallas._ Nova Acta Acad. Scient. Petrop. tom. ii (1788), p. 240, Tab. 6, fig. 24. _Parietes thick, formed by several rows of unequal-sized pores. Tergum narrow, with the apex beaked, and spur sharply pointed._ _Hab._--Columbia River, west coast of North America, Mus. Brit. and Cuming; Behring Straits (Capt. Kellett); the Kurile Islands, according to Pallas. Attached to shells, and to each other in groups. _General Appearance._--Shell steeply conical, with a rather small oval orifice; or cylindrical, with a large rhomboidal and little toothed orifice. Colour dirty white. Surface either simply rugged, or more commonly covered by numerous, narrow, extremely prominent, longitudinal plaits; from the manner in which these overlap each other, the shell almost appears as if thatched with straw. The upper corroded part of the shell usually exhibits a cancellated and finely punctured surface. The radii are generally very narrow, forming towards the base of the shell a mere narrow ribbon to each compartment, and often hardly distinguishable; but in one specimen they were of considerable width: in the former case, the alæ are often widely exposed. The largest specimen which I have seen was 1.5 of an inch in basal diameter, but Pallas gives 2.2 as the measurement of a specimen from the Kurile Islands.
The opercular valves are united to each other and to the shell by unusually strong membrane; and the upper parts of both valves, in all the specimens seen by me, have been much disintegrated. The _Scutum_, in old specimens, is faintly striated longitudinally, but in some there is hardly a trace of this: the occludent margin is furnished with a few large knobs, not corresponding with every alternate line of growth (as is usual with other species), but with every fourth or fifth line. Internally, the articular ridge is moderately prominent (in young specimens more prominent) and reflexed. The adductor ridge is sharp and prominent; in the upper part it is confluent with the articular ridge, but in young specimens can be seen to be distinct; in the lower part it borders a large deep cavity for the lateral depressor muscle, in the middle of which there is a very slight longitudinal ridge; this cavity sometimes is almost closed or arched over in its upper part. In one specimen, the basal margin of the scutum was deeply hollowed out in the middle. The _Tergum_ is remarkably narrow, with its apex produced into a triangular beak, hollow within, and sometimes faintly tinged purple. A deep, closed, longitudinal furrow runs down the valve. The spur is long, remarkably narrow, and pointed. Internally, the spur is produced up the valve as a ridge: the inflected scutal margin, and the prominent articular ridge, are both nearly straight, and parallel to the spur. The crests for the depressores are sharp and very prominent.
The _Parietes_ are very thick and strong: unlike every other species of the genus, they consist of several very irregular rows, of unequally sized, round or angular tubes (3 _b_). These tubes or pores are generally short, and are at frequent intervals crossed by transverse septa; they often rather deserve to be called cells than tubes. New tubes are formed along the inner as well as along the outer lamina. They are lined by dusky purple corium. The internal surface of the parietes is smooth in the upper part, and in the lower, it is reticulated by slight, irregularly branching ridges. The carinal internal margin of each compartment projects a little, as in the case of _B. crenatus_ and _balanoides_. The lower edge of the sheath is either hollow beneath, or is united to the walls. The _radii_ in one specimen were broad, with slightly oblique, jagged summits; generally they are extremely narrow, forming more ribbons along the lower edges of the compartments, barely extending up as high as the sheath. They can sometimes hardly, or not at all, be seen, until the shell is disarticulated: in rather young specimens the sutural edge is sometimes quite smooth; in old specimens the lower part of the edge has coarse arborescent septa, with the interspaces filled up solidly, whilst the upper part is smooth. The _alæ_ are conspicuous from the outside, owing to the little development of the radii; but owing to the diametric growth not having been great, the part added during such growth is narrow; the summits of the alæ are only slightly oblique: the sutural edge is coarsely crenated, with the teeth denticulated or slightly arborescent.
_Basis_ membranous. _Mouth_: labrum with only four very minute teeth: mandibles with four teeth; the third tooth broader than the first; the fourth small. Maxillæ, with the two upper spines placed on a slight prominence, beneath which there is a small notch. _Cirri_ of a very dark colour (much injured): the segments of the first, second, and third pairs very broad and short, protuberant in front, and most thickly clothed with spines; the third pair is very little longer than the second pair: the sixth pair (in a large specimen above one inch in basal diameter) had the segments broader than long, each furnished with seven pairs of spines. _Affinities._--This species, though very distinct, evidently comes near to _B. balanoides_, especially to _var._ (_a_). By merely doubling or trebling the irregular rows of short tubes in the walls of _B. balanoides_, with their reticulated inner lamina and longitudinally folded outer lamina, we should have the structure exhibited in _B. cariosus_. We have seen, also, that in _var._ (_a_) of _B. balanoides_, the spur of the tergum is remarkably sharp, as in _B. cariosus_. This species, also, in a very marked manner approaches in many characters, especially in the opercular valves, in the cirri, and to a certain extent in the shell and basis, to _B. flosculus_, and even in external appearance to _var. sordidus_ of the latter--an inhabitant of the opposite extremity of the continent, namely, of Tierra del Fuego. Again, the tergum to a certain extent, and the scutum in a singular manner, resemble these valves in _B. nubilus_, showing an unequivocal affinity to that species. With respect to the most remarkable character of the species, namely, the several irregular rows of tubes or pores in the walls, it deserves notice that in _B. crenatus_, which is certainly closely allied to _B. balanoides_, the longitudinal septa sometimes divide near the outer lamina, thus giving rise to a few additional tubes. Of the above several species, to which our present species is allied, _B. flosculus_ stands in the next section, and _B. nubilus_ and _crenatus_ in the last: hence we see that _B. cariosus_ has singularly divergent affinities. The peculiar structure of the parietes, together with the general appearance of the shell, made me at the first moment suppose I was examining a Tetraclita (or Conia of Leach); hence, also, it has arisen, that _Lepas cariosa_ of Pallas has often been quite erroneously given as a synonym of _Tetraclita porosa_.
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