A Monograph on the Sub-class Cirripedia · Charles Darwin
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_Mouth_: labrum with six teeth: mandibles with the fourth and fifth teeth small and rudimentary: maxillæ, with a small notch under the upper pair of spines; in the lower part there is a single large spine. _Cirri_, dark brownish purple, making a singular contrast with the white operculum and shell; first pair, with one ramus, having twenty-six segments, and about twice as long as the shorter ramus, having twelve or thirteen segments, with their front surfaces protuberant. In the second pair the segments are but little protuberant: third pair about one third longer than the second pair: sixth pair, elongated, having in the same individual forty-six segments; these segments have shield-shaped fronts, bearing five pairs of spines, with some minute intermediate bristles. There is the usual point at the dorsal base of the penis.
_Range: Geological History._--This species is common on the shores of Scotland and Ireland; the most southern point of Europe whence I have happened to see a specimen is Tenby, in South Wales: but I have no doubt it is found further south; and Mr. Jeffreys, who knows this species well, has found it common on the extreme southern shores of England. In the United States, it is found on the shores of Maine and Massachussetts: northward, I have seen specimens from Iceland, from Davis's Straits, and from Lancaster Sound, in lat. 74° 48′ north; these latter I owe to Sir J. Richardson. It is an inhabitant of deep water; in Mr. Thompson's collection there are several specimens from the Bay of Belfast, marked twenty-five fathoms, and one group said to have come from "about fifty fathoms, on the coast of Antrim:" one specimen from Cape St. Anne, Massachussetts, is marked as having come from only five fathoms. This species is commonly associated, on both sides of the Atlantic, with _B. crenatus_, and sometimes with _B. Hameri_ and _Verruca Strömia_: mollusca, such as pectens, modioli, and oysters, offer the most usual surfaces of attachment: I have, however, seen many specimens on crustaceans, on rocks, and even on the roots of the larger sea-weeds. This species is very common in the glacial deposits of Uddevalla, of Skien in Norway, and of Canada, and is associated with the same species as in the living state: I have seen, also, specimens from the same formation in the Island of Bute, Scotland. I have seen numerous specimens from the mammaliferous crag, and a few from the Red Crag of England. I owe to the kindness of Mr. J. de C. Sowerby an inspection of the original specimens of the _B. tesselatus_ of the Mineral Conchology, which is certainly the present species.
_Affinities._--This species is very distinct from every other; it comes nearest, as shown in all the characters derived from its opercular valves, to _B. nubilus_, and in this latter species we have seen the basis plainly tending to lose its pores and thus become solid. _B. porcatus_ is perhaps allied in some degree to _B. trigonus_, and slightly to _B. crenatus_. The rather broad radii, with their summits hardly oblique, give this species a very different aspect from those species of the genus amongst which it must be placed. _Varieties._--A conical specimen, sent to me from the coast of Massachussetts, is remarkable from the radii not having been at all developed, being represented by mere fissures. I have seen a few specimens of _var._ (_a_), (one collected by Sir E. Parry in the arctic seas) which had a remarkably different aspect from the common forms, but which, after a careful examination of the opercular valves and of the animal's body, I feel convinced are not specifically distinct: they are characterised by the walls being smooth and absolutely destitute of the external longitudinal ribs; by the shell being more cylindrical, with broader radii, and with the orifice larger and more rhomboidal; the walls and radii are much thinner, and the internal lamina is less plainly ribbed: the beak of the tergum is not purple. As most of these specimens had grown in a group crowded together, the difference of shape, and perhaps the thinness of the walls, is thus explained. In a specimen from Davis's Straits, in Mr. A. Hancock's collection, most of the above characters are in an intermediate condition; there are only a few external longitudinal ribs on the parietes; and the terga have not purple apices. In Mr. Cuming's collection there are some fine, brilliantly white specimens (_without opercula_) from the coast of China; these have thin walls and radii, and the walls are not longitudinally ribbed, but they are not smooth: the orifice is not large, nor the shape of the whole shell cylindrical. It is just possible that these latter specimens may be a distinct and representative species, but I do not think so.
26. BALANUS PATELLARIS. Pl. 6, fig. 5 _a_-5 _c_. LEPAS PATELLARIS, (_Gmelin_). _Spengler._ Schriften der Berlin. Gesellschaft, &c. b. i (1780), Tab. 5; Chemnitz, Neues Syst. Couch., Tab. 98, fig. 839. _Shell depressed; brown, generally with obscure longitudinal violet stripes: radii (in full-grown specimens) with their summits rounded and surfaces finely ribbed parallel to the basis: basis sometimes permeated by imperfect pores. Scutum internally with an adductor ridge._ _Hab._--Bengal, on wood, Mus. Brit.; on a shell, Mus. Stutchbury; Philippine Archipelago (young specimen), Mus. Cuming. According to Spengler, on the Coromandel and Malabar coasts. _General Appearance._--Shell depressed, sometimes much depressed: orifice elongated, rhomboidal, but little toothed; surface smooth, but in old specimens sometimes with the walls slightly folded longitudinally. The radii are rather narrow, with their summits oblique; in old specimens their summits are rounded, and their whole surface finely ribbed parallel to the basis. Colour, in old specimens dirty brown, tinged with violet, sometimes in longitudinal bands, and with whiter irregular marks in the upper parts owing to disintegration: in young specimens the walls are regularly banded longitudinally, with violet-brown and dirty white; the radii being generally of a paler dirty red or violet. Basal diameter of largest specimen .9 of an inch. _Scuta_, externally rather smooth; internally, articular ridge prominent, reflexed, with the lower edge hollowed out so as to be slightly hook-formed: adductor ridge small; there is a slight pit for the lateral depressor. _Tergum_, with the spur bluntly pointed, placed at about its own width from the basi-scutal angle; there is no longitudinal furrow, only a slight depression; carinal margin arched and protuberant: internally, articular ridge extremely prominent, running down in the direction of the middle of the spur: crests for the tergal depressores well developed.
_Parietes_, with the pores rather large; the internal lamina is very strongly ribbed, the ribs being but slightly denticulated at their bases: the parietal pores do not appear to be crossed by transverse septa: sheath closely attached to the walls. The radii have jagged oblique summits forming an angle of about 45° with the horizon; in old specimens they become more oblique and narrow: and are then very remarkable from their summits being arched and rounded, with a crenated edge, and with their whole surface transversely ribbed in horizontal lines; this is likewise the case with the recipient furrow in the opposed compartments: in young specimens the radii are externally quite smooth: the septa on the sutural edges are bluntly denticulated; the interspaces being filled up solidly. The alæ have their summits oblique, but much less oblique than the summits of the radii; their sutural edges are very finely crenated. _Basis_ thin, either quite solid, that is, not permeated by pores, but only furrowed in lines radiating from the centre, or permeated by pores towards the circumference, the pores being of very small diameter;--so that we here have an important character variable within the limits of the same species. Base flat, and this holds good, as remarked by Spengler, even when the specimens are attached to cylindrical pieces of wood.
_Animal's body_ unknown. _Affinities._--In the basis being sometimes permeated towards the circumference by pores, and by the colouring (the other species in this and the next section being dirty white), _B. patellaris_ has almost as strong a claim to be ranked in the last as in the present section: in the rounded summits of the radii, and in the state of the basis, it, perhaps, shows more affinity to _B. improvisus_ than to any other species; it is, however, almost equally allied to _B. glandula_. 27. BALANUS CRENATUS. Pl. 6, fig. 6 _a_-6 _g_. B. CRENATUS. _Bruguière._ Encyclop. Method. (des Vers) 1789. LEPAS FOLIACEA, _var. a_. _Spengler._ Skrifter af Naturhist. Selskabet, b. i, 1790. ---- BOREALIS. _Donovan._ British Shells, Pl. 160 (1802-1804). B. RUGOSUS. _Pulteney_ (?) Catalogue of Shells of Dorsetshire, 1799. ---- ---- _Montagu_ (?) Test. Brit. 1803. ---- ---- _Gould_ (!). Report on Invertebrata of Massachussetts (1841), fig. 10. B. GLACIALIS (?) _J. E. Gray._ Suppl. Parry's Voyage, 1819. B. ELONGATUS (!), CLAVATUS (!), _Auctorum variorum_. _Shell white: radii with their oblique summits rough and straight. Scutum without an adductor ridge: tergum with the spur rounded._ _Hab._--Great Britain, Scandinavia, Arctic Regions as far as Lancaster Sound, in 74° 48′ N. (Mr Sutherland); Behring's Straits (Captain Kellett); United States; Mediterranean; West Indies, (Mus. Brit.); Cape of Good Hope, (Mus. Krauss). Generally attached to shells and crustacea in deep water; sometimes to ships' bottoms. Very common.
_Fossil_ in glacial deposits of Scandinavia and Canada, Mus. Lyell; in the mammaliferous, and Red, and Coralline Crags, Mus. S. Wood, J. de C. Sowerby, Bowerbank; Miocene formation, Germany, Mus. Krantz. I find, in most collections, this species confounded with _B. balanoides_; I have even seen the two species, placed by Leach, on the same tablet in the British Museum: _B. balanoides_ is, moreover, generally confounded with _Chthamalus stellatus_; nor has any one hitherto separated the present species from _B. improvisus_. On the other hand, trifling varieties, both of _B. balanoides_ and _B. crenatus_, have commonly been considered as specifically distinct. From these facts it will be seen in what confusion our commonest British species of Balanus have been left. After due deliberation, I have little doubt that this is the _B. crenatus_ of Bruguière, and probably the _B. rugosus_ of Montagu, but this latter author omits all reference to the really important diagnostic characters between this species and _B. balanoides_. The _B. crenatus_ is certainly the _B. rugosus_ of Dr. Aug. Gould. In various collections, I find specimens of _B. crenatus_, when coming from the arctic regions, called _B. glacialis_, _arcticus_, and _borealis_; though I have not met with an authentic specimen of the _B. glacialis_ of Gray ('Supp. Parry's Voyage,' 1819, p. ccxlvi), I have little doubt that it would prove to be the present species.
_General Appearance._--White, usually of a dirty tint, from the yellowish or brownish persistent epidermis: conical, generally (fig. 6 _a_) with the parietes rugged and irregularly folded longitudinally; but sometimes much depressed and extremely smooth (6 _b_); often cylindrical and very rugged; occasionally club-shaped (6 _c_), the upper part being much wider than the lower: specimens in this latter condition sometimes have extremely narrow parietes, like mere ribs, and wide radii. The orifice in the cylindrical varieties is often most deeply toothed. The radii are generally narrow, and have jagged oblique summits; but not infrequently they are so narrow as to form mere linear borders to the compartments. The orifice is rhomboidal, passing into oval, either very deeply or very slightly toothed. _Dimensions._--The largest British specimen which I have seen was only .55 of an inch in basal diameter: specimens from Greenland and the northern United States frequently attain a diameter of three-quarters of an inch, and I have seen one single somewhat distorted specimen actually 1.6 of an inch in basal diameter. The specimens from the glacial deposits of Uddevalla and Canada appear, on an average, to attain as large or larger dimensions than those from the United States: on the other hand, the specimens from the mammaliferous and Red Crag are smaller, the largest being only .35 in basal diameter. When individuals have grown crowded together, their length is often twice, and even occasionally thrice, as great as their greatest diameter; thus I have seen a Greenland specimen 1.6 of an inch in length, and only .75 in diameter. In the British Museum there are some arctic specimens, one and a half inch in length, only half an inch in diameter at the summit (fig. 6 _c_), thence tapering downwards to a blunt point.
_Scuta_; the lines of growth are but little prominent: the surface is generally covered by disintegrating membrane. The upper ends are usually a little reflexed, so that the tips project freely as small flattened points. Internally, the articular ridge is highly prominent and somewhat reflexed: there is no adductor ridge, but a very distinct impression for the adductor muscle: the depression for the lateral depressor muscle is small, but variable. The _terga_ are rather small: the spur is short, and placed at rather less than its own width from the basi-scutal angle; the basal margin slopes a little towards the spur, of which the lower end is rounded or bluntly pointed in a variable degree. There is no longitudinal furrow, hardly even a depression. Internally, the articular ridge is very prominent in the upper part; the crests for the tergal depressores are well developed, but variable.
_Compartments._--The internal carinal margin of each compartment, from the sheath to the basis, generally, but not invariably, projects a little inwards beyond the general internal surface of the shell, in a manner not common with the other species of the genus: the basal edge of this projecting margin rests on the calcareous basis, and is crenated like the basal edges of the longitudinal parietal septa. The whole internal surface of the shell is ribbed, but the ribs are not very prominent. The parietal tubes are large, and are crossed in the upper part, and often low down, by transverse thin septa: the longitudinal parietal septa are only slightly denticulated at their bases; occasionally they divide at the basis close to the outer lamina of the parietes, making some short outer subordinate pores. In the circular furrow beneath the lower edge of the sheath, there are sometimes little ridges, dividing it into small cells: sometimes, however, this furrow is filled up by irregular knobs of calcareous matter. The _radii_ are always rather narrow, and often they form mere linear ribbons of nearly uniform width along the edges of the compartments. Their summits or edges are always more or less irregular and jagged: they form an angle with the horizon of generally above 40°. Their septa are fine, and barely or not at all denticulated. The alæ have oblique summits: their sutural edges are rather thick and distinctly crenated. _Basis_ flat, calcareous, very thin, with the surface slightly marked by radiating furrows, which furrows answer to the radiating pores that occur in the bases of most species. In a club-shaped arctic specimen, one inch and a half in length, the summit being half an inch and the base only one fifth of an inch in diameter, the basis was still calcareous, thick, and not permeated by pores.
_Mouth_: labrum with six teeth: mandibles with the fourth tooth minute or rudimentary, and the fifth generally confluent with the inferior angle. Maxillæ with generally, but not invariably, a small notch under the upper pair of great spines. _Cirri_, first pair with the rami very unequal in length, one ramus being nearly twice the length of the other; in a large specimen having a cylindrical shell the proportional numbers of the segments in the two rami of the first cirrus were ten to twenty-three; in a small conical specimen the numbers were only eight to thirteen. The second cirrus has only two or three more segments than the shorter ramus of the first pair: the third cirrus has one or two more segments than the second; but it is nevertheless decidedly longer than the second. On the dorsal surfaces of both segments of the pedicel of the third cirrus, there is a tuft of fine spines. The segments of these three pairs of cirri are not much protuberant in front. The segments of the posterior cirri have, each, four, or five, or six pairs of spines. _Penis_, with a straight, sharp, short point on the dorsal basis.
_Range, habits, &c._--I have received specimens from all parts of the coast of Great Britain and Ireland, generally attached to crustacea and mollusca, and never hitherto from rocks uncovered by the tide. This species is also attached to floating timber, sticks, fuci, and occasionally to pebbles at the bottom of the sea. Mr. Thompson has sent me specimens from twenty-five fathoms depth in Belfast Bay: others on a Pinna from _about_ fifty fathoms on the coast of Antrim; others from between three and six fathoms attached to Laminaria digitata: there is a specimen in Mr. Jeffreys' collection marked forty-five fathoms. It is often associated, both on the coasts of America and Britain, with _B. porcatus_, and though these species are so distinct, yet when both have their surfaces similarly affected by being attached, as is often the case, to large Pectens, it is not at first easy, by external characters, to distinguish them, except by close inspection of the terga, which in _B. porcatus_ are beaked and purple. The _B. crenatus_ is sometimes associated in deep water with _B. Hameri_. At Ramsgate, in Kent, I saw a rudder of a ship, in which the two or three upper feet were thickly coated with _B. balanoides_, and the two or three lower feet with _B. crenatus_ and _improvisus_ mingled, together with a few of _B. balanoides_: occasionally vessels are thickly encrusted with this species, but I have never seen an instance of its concurrence with _B. tintinnabulum_ and _amphitrite_--the commonest species on ships coming from the south. I have seen specimens from Greenland, Baffin's Bay, the coast of Labrador, and other specimens marked simply, "Arctic regions," and, again, others from the shores of Maine and Massachussetts. The arctic specimens, and those from the northern United States, are larger than the British. I have seen one single minute specimen on a crab, marked as having come from the Mediterranean.
In the British Museum, amongst some specimens of _B. eburneus_, ticketed as having been sent from Jamaica, there was a small group of specimens, differing in no one essential respect from the common varieties of _B. crenatus_: at first I concluded that this was an erroneous habitat, and that the specimens had really come from the United States, where _B. eburneus_, is found as well as in the West Indies: for it appeared to me exceedingly improbable that an animal which can exist in lat. 75° N. should inhabit the hot shores of Jamaica: but subsequently I have received a specimen from Prof. Krauss, collected by himself in Algoa Bay, which is perfectly characterised, and even has the little cells in the furrow under the sheath: so that I am compelled to admit this enormous range and capability of resisting the most extreme climates. That this species should live in the tropical seas is the more surprising, as the large size of the specimens in the northern seas and in the glacial deposits, might fairly have been supposed to have indicated special adaptation for a cold climate. The great geographical range of this species accords with its range in time from the present day to the Coralline Crag period.
The specimens from the glacial deposits which I have examined, chiefly in Sir C. Lyell's collection, are very fine and large; they are often associated, like the now living individuals, with _B. porcatus_ and _Hameri_: they come from the well-known formation of Uddevalla and from Canada. There are well-characterised specimens in the mammaliferous Crag, at Bramerton and near Norwich, in Sir C. Lyell's collection, and from Sutton and other places in the Red Crag of the eastern shores of England: these specimens are decidedly not only smaller than the glacial, but than the recent English specimens; for the largest Crag specimens which I have seen had a basal diameter of only .35 of an inch. The specimens which I have seen from the Coralline Crag, and some others sent me by Krantz from the miocene formation of Flonheim bei Abzei, in Germany, had not their opercular valves, yet I cannot doubt, considering how few species there are in the present section of the genus, that I have rightly identified them. _Diagnosis._--Under the head of _B. balanoides_ I shall make a few remarks on the diagnosis between that and the present species; as _B. improvisus_ is found on the British shores, sometimes mingled with _B. crenatus_, I may observe that, externally, the only difference consists in the edges of the radii in _B. improvisus_ being much smoother and rounded, and in the whole shell being less rugged. Internally, in _B. improvisus_ the porose basis, the presence of an adductor ridge on the under side of the scutum, the graduated teeth on each side of the central notch in the labrum, and the little inequality in length of the rami of the first pair of cirri, are clearly and amply diagnostic.
28. BALANUS GLANDULA. Pl. 7, fig. 1 _a_, 1 _b_. _Shell white; parietes with the internal lamina generally strongly ribbed longitudinally, with the pores imperfect and small, sometimes in part absent; radii narrow, with their summits rounded. Scutum with an adductor ridge; tergum with the spur truncated and rounded._ _Habitat._--California, Mus. Cuming, Aug. Gould; attached to shells and wood, together with _B. nubilus_. Southern Pacific ocean, attached to _Pollicipes polymerus_; Mus. Brit. _General Appearance._--Shell steeply conical, or cylindrical and elongated; dirty white; walls rugged, longitudinally folded; radii narrow, with their summits very oblique and rounded; orifice toothed. Basal diameter of largest specimen half an inch. _Scutum_, resembling externally that of _B. crenatus_; rather broad, surface smooth; articular ridge very prominent, and articular furrow very wide; hence, when the summits of the opercular valves are worn down, the two scuta together form a square projection indenting the two terga, as in _B. balanoides_. Internally, there is a small adductor ridge, on the lower side of which there is a pit, as if for a muscle. The depression for the lateral depressor muscle is small, but variable. _Tergum_ without any longitudinal furrow, and hardly a depression: spur broad, with its lower end truncated and rounded; internally, articular ridge very prominent; crests for the depressores well developed.
_Compartments_:--The internal surface of the parietes is smooth in the upper part beneath the sheath, but generally very strongly ribbed in the lower part, the ribs being plainly denticulated at their bases; in other specimens, the ribs are very small, and even in parts quite obsolete. The parietal pores are short and imperfect, sometimes reduced to an extremely minute size, to be detected only when the walls are broken across near the basal edge, and most carefully examined; occasionally not even a trace of a pore exists. Hence in this respect, this species offers a singular case of variation. The _radii_ are narrow, and of nearly the same width from top to bottom; their very oblique summits, when well preserved, are smooth and rounded; their sutural edges are ribbed or crenated with extremely fine, smooth septa; the recipient furrow is plainly marked by these septa. The sutural edges of the alæ are crenated; their summits are less oblique than those of the radii. _Basis_, thin, finely furrowed in lines radiating from the centre; margin sometimes deeply sinuous. _Mouth_: labrum with the central notch rather widely open, with four teeth on each side of it: palpi with very short spines along their inner margins: mandibles with the fourth and fifth teeth forming mere knobs: maxillæ small, with a mere trace of a notch under the two great upper spines. _Cirri_; first pair with the rami unequal by three or four segments, the longer ramus being only one quarter of its own length longer than the other ramus. Second pair short, with the segments (and those of the shorter ramus of first pair) somewhat protuberant. Third pair with the rami one third longer than those of the second pair. Sixth pair with the upper segments elongated, and bearing six or seven pairs of spines.
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