A Monograph on the Sub-class Cirripedia · Charles Darwin
TABLE OF CONTENTS.
On parts of the shell where there are no bristles, similar tubuli penetrate the shelly layers, and come to the surface. The tubuli running to the lowest and last-formed row of bristles, just after a period of exuviation, are so delicate as hardly, or not at all, to be distinguished; in the row above, they are plain and longer, and for the next two or three upper rows they are, in some cases, as in _Tetraclita porosa_, longer and longer, having been added to during each successive thickening of the valve. These tubuli consist of chitine, and no doubt first existed as threads of corium; they are so tough that they must serve to strengthen the successive layers of shell, but I imagine their chief function is to keep up the vitality of the newly-formed layers of shell. May we not, also, venture to suppose that by their means, some degree of sensibility is given to the bristles? I need only further remark, that in some species of Balanus and of Chthamalus, the under side of the shell is penetrated by irregular pores, large enough to be visible to the naked eye, into which threads of corium penetrate; but these can hardly be said to appertain to the microscopical structure; and are more nearly related to those pores and furrows, formed by the greater or less development of the longitudinal septa, and in which the threads of corium deposit, or rather become changed into, transverse septa, or solid shelly matter, as previously described.
The above-mentioned muscles are attached at their upper ends to the opercular valves, and at their lower ends to the basis. There are, in fact, three pairs, but the pair attached to the basi-carinal angles of the two terga (Pl. 25, fig. 1, _i_), are almost invariably confluent, forming one great bundle; the second pair is attached to the lateral or basi-tergal corners of the two scuta, and are hidden in the figure; the third pair (_h_) is attached also to the scuta, to their rostral angles. These muscles can only act as depressores; they are often extremely powerful; they belong to the voluntary class, for they are transversely striped. By their action, the opercular valves are capable of varied slight movements, within the limit allowed by the width of the flexible opercular membrane. By the action of the lateral scutal depressores, the orifice leading into the sack is opened, the movement being generally aided by the protrusion of the cirri. By the sudden contraction of the rostral scutal depressores, the blows which are sometimes given by the beaked terga at the opposite end of the operculum, are probably effected. By the contraction of all three pairs of muscles, the opercular valves are held down with quite surprising force. The valves can be raised only by the action of the animal's body against the basis.
In Coronula these muscles are more spread out, and do not extend down to the basis; their lower portions, as is likewise the case in Tubicinella, do not exhibit transverse striæ, and hence tend to pass either into the involuntary class, or into ligament. This condition of the muscles, in the above two genera, accords with the little-developed state of their opercular valves. In Xenobalanus, there is no longer any evidence of the muscles being collected into five or six bundles, for they are thinly and almost uniformly spread out, and show in no part transverse striæ. I may add that in much elongated specimens of _Balanus balanoides_, these muscles become in their lower part ligamentous, and destitute of striæ. _Branchiæ._
In the Balaninæ, a pair of Branchiæ is always present: they lie on each side, in a somewhat curved position, in the angle between the sides of the shell and the basis. In Pl. 25, fig. 1, they are exactly covered, on the further side, by the body of the animal. They are attached near each other at the carinal end of the sack in a vertical line, and likewise on each side in a transverse line, extending from close beneath the spur of the tergum towards the point of attachment of the body to the scutum. In Balanus, as in the figure (Pl. 25, fig. 3) of _B. tintinnabulum_, each branchia consists of a medial fold of skin, a little curved conformably with the sack, and slightly tapering towards its rostral and free extremity; but this fold is almost hidden by the vertical sub-folds or membranous ridges, themselves plicated and sub-plicated, which project on both sides: these vertical folds are free at their tips: at their lower attached ends, they are thickest. On the side nearest the wall of the shell, the whole branchia has a bilobed appearance, owing to a very deep indentation caused by the projection of the scutal lateral depressor muscle; the sub-folds on this side are also more plicated.
The branchia essentially is an inward plicated fold of the membranes of the sack; for its outer, very thin tunic is continuous with and moulted with that lining the sack; and within it we have two layers of delicate, pulpy, transparent corium, united together (as is best seen in Coronula) by ligamentous fibres, branched at their two ends, all exactly as in the corium surrounding the sack. There are here no distinct vessels, any more than in other parts of the body, but a fluid could easily circulate in the interspaces of the corium. From the large size of this organ, and its simplicity of internal structure, being adapted exclusively to expose a great surface of skin to the water, I do not doubt that it has been correctly considered as a respiratory organ. By the voluntary movements of the opercular valves (_i. e._ part of the carapace) the water is constantly being pumped in and out of the sack; the movement, indeed, may be almost compared to the heaving of a man's chest. Moreover, the branchiæ on each side are attached so closely to the spur of the tergum, that each time the latter is moved, the whole branchia must, I think, be agitated, and the folds opened, as by the action of a lever.
In our two commonest, tidal, sessile Cirripedes, viz. _Balanus balanoides_ and _Chthamalus stellatus_, I have observed that, when left uncovered by water, they kept the orifice of their operculums a little open, with a bubble of air within their sacks, so that the orifice was in fact closed by a thin septum of water, with air beneath; when disturbed, they closed their operculums with force, and expelled the bubble of air with a clicking noise, which has been noticed by Dr. Coldstream,[27] and has been thought to be made by the movement of the operculum itself. _Bal. crenatus_, a deep-water species, when out of water, keeps its operculum closed. [27] 'Encyclopædia of Anatomy and Physiology;' article Cirrhopoda. In Coronula, Platylepas, Tubicinella, and Xenobalanus, each branchia[28] consists of two unequal folds, both plicated on both sides: in the two latter genera, they extend far down the deep and elongated sacks, and hence the area of surface altogether gained is extremely great. In most of the species of Chthamalus, the branchia consist of a small fillet barely plicated: in the allied _Chamæsipho columna_, they are rudimentary, forming a smooth little pouch only 1/100th of an inch in length: in _Chthamalus scabrosus_ they are quite aborted, being perhaps represented by a slight hairy ridge; but in _Chthamalus dentatus_, and therefore within the limits of the same genus, the branchiæ (and this seems to me a singular fact) are large, each being composed of two plicated folds, as in Coronula. Tapering filaments situated near the bases of the cirri, such as those occurring in several species of the Lepadidæ, are not found in any sessile Cirripede; but I have observed nearly similar filaments, projecting upwards and inwards at the base of the sack, in several species of Balanus and in Coronula; those which I examined were simply occupied by delicate corium, and no doubt must aid in exposing a greater surface of corium to the circumambient water.
_Thorax and Body._ _Parts of the body included within the shell or carapace._--These parts (Pl. 25, fig. 1) consist of the prominent mouth, and of the thorax (_c′_), with its largely developed portion, called the prosoma (_c_), and with its appendages. The abdomen is quite rudimentary, being represented merely by a small portion of membrane surrounding the anus, and sometimes inserted like a wedge between the inwardly inflected posterior thoracic segments; in only two genera (Catophragmus and Pachylasma), its nature is rendered somewhat plainer by supporting caudal appendages. The probosciformed penis lies folded under the thorax; and I believe (from what is seen in the anomalous genus Proteolepas), that it normally arises from the ventral surface of the terminal point of the rudimentary abdomen.[30] The thorax is laterally compressed, the ventral surface being very narrow, with the bases of the cirri placed closely together. It consists, in appearance, of two very different portions; one a soft, more or less rounded bag, which I have called the prosoma; and the other, which supports the five posterior pairs of cirri, is narrower, invested with stiffer membrane, and is more or less distinctly composed of five segments. These segments (Pl.
26, fig. 8) on their dorsal and dorso-lateral surfaces, are generally driven like wedges one into the other, with their points directed anteriorly: on the ventral surface the articulations are transverse. The prolongation (_e_) of the thin membrane (_a_) surrounding the anus (_b_), that is, the rudiment of the abdomen, which sometimes carries caudal appendages, almost divides (in appearance, whether really I know not) the hindermost thoracic segment along the medio-dorsal line, into two parts. I have given the above drawing of these segments, but with the dorsal surface much flattened, in _Coronula diadema_; in most species of Balanus, however, the wedges formed by one segment being driven into another, are much sharper; on the other hand, in Xenobalanus they are nearly straight and transverse. The three posterior segments are always the most distinct; the two next segments are also distinct laterally, but along the dorsal surface they become, excepting in Xenobalanus and some few other cases, completely confluent. The greater distinctness of the posterior segments is conformable to what takes place in the higher Crustacea.
The articulations between the segments are folded inwards, and are formed of thin membrane, which in some cases, as in _Coronula diadema_, forms a marked contrast with the much thicker, stiffer, and yellowish membrane of the segments themselves; in _Balanus tintinnabulum_, however, the whole membrane of the five thoracic segments is very thin, excepting small wedge-shaped portions along the medio-dorsal line. The infolded articulations between the segments supporting the three anterior pairs of cirri (at least in the Balaninæ), are much wider than those between the three posterior segments; the former segments, with their cirri, being consequently capable of being moved further apart from each other. Could there have been any doubt as to the distinctness and reality of the five thoracic segments, it would have been set aside by the arrangement of the muscles attached to them, as will presently be described. I need only add, that in many genera there are shield-like swellings at the exterior bases of the pedicels of the posterior cirri, which I for some time thought were the epimeral elements of the thoracic segments; but I now believe them to be parts of the pedicels of the cirri. The basi-exterior margin, moreover, of the pedicel of the third pair of cirri, in many species of the Balaninæ (Pl. 25, fig. 1), is produced as a plate, thickly fringed with fine hairs, half across the dorsal surface of the thorax; serving, apparently, as a brush to clean the sack, or to prevent the ingress of any intruding substance.
_Muscular System._ _Attachment of the Body to the Shell._--The prosoma which carries the posterior thoracic segments, and in appearance the mouth, is the only part of the body which is attached to the general covering (Pl. 25, fig. 1), namely, to the opercular valves. Except through the continuity of the lining membranes of the sack, the body lies free within the walls of the shell. The area of attachment (shown by a sinuous broken line round _a_ and _b_) extends from about the middle of the two scuta down to their basal margins. As these valves lie obliquely across the orifice of the shell, the animal's body comes to be suspended almost in the middle of the sack. The two scuta, as we have seen, have the power of opening and shutting a little; and are brought together by the adductor scutorum muscle (_a_), which is generally very powerful. The body is attached to these valves, round and beneath the adductor, so as to hide it until one of the valves be removed. The attachment is chiefly effected by three pairs of widely expanded, superficial muscles, two pairs of which are spread over the flanks of the prosoma, and the third pair over its rounded (properly dorsal) surface, which lies close to the rostral compartment (A, fig. 1) of the shell.
I should have stated, that my chief examination of the attachment of the body to the scutal valves, has been made on _Coronula balænaris_, and less closely on _Balanus tintinnabulum_. Within these three pairs of superficial muscles, there are (besides the adductor) no less than five other pairs; of these one long pair is attached at one end to the basal margin of the labrum (_e_), and at the other end, to the under side, near to the basal margin of the scuta: two other, shorter, parallel pairs of muscles are attached at one end to the interspace of membrane between the basal edge of the labrum and the adductor scutorum muscle, and at the other end, to the under side of the scuta, above the attachment of the first pair: the fourth and shortest pair curls close under the adductor, and is there attached at both ends beneath it. The action of these four pairs of muscles must be to draw back, from the orifice of the shell, the mouth, and that interspace of body between the basal margin of the labrum and the adductor muscle. This movement I saw in living specimens.
The last and fifth pair of muscles is small, but of considerable length; it is a diverging pair, attached at the converging end, above and exteriorly to the adductor muscle; and at the diverging end, low down on the under side of the scuta; I am very doubtful regarding the function of this pair. Altogether we have seen that round and within the fleshy pedicel, by which the body is attached to the scuta, there are no less than eight pairs of muscles. The central space between these muscles is hollow, and here many lacunal channels seem to converge. These muscles receive nerves from the supra-œsophageal ganglions. The interspace above alluded to, between the basal edge of the labrum and the adductor scutorum muscle, occupies a very different position according as the animal's body is protruded as far as it can be, or is retracted. It is homologically part of the third cephalic segment; and consequently the mouth ought to have stood posteriorly (_i. e._ above, in the position figured in Pl. 25, fig. 1) to this interspace; yet, in fact, when the animal is retracted within its shell, the mouth usually lies almost directly beneath this interspace and the adductor scutorum muscle.
Besides these muscles of attachment, the prosoma is furnished with several other muscles. There are superficial muscles running up towards the basal margin of the sides of the mouth; and other deeper muscles, to which, I presume, the movements of the mouth, as a whole, are due. The muscles moving the gnathites do not, as far as I could make out, extend beneath the basal edge of the mouth. There are, also, powerful muscles giving movement to the basal segments of the pedicel of the first pair of cirri. Again, there are superficial muscles running to the next succeeding thoracic segment; the anterior ends of which are separated by a clear interspace from the lower ends of the above-described superficial muscles, by which the prosoma is attached to the scuta. On each flank, moreover, but more deeply imbedded, are the long flexor and extensor muscles, presently to be described, running to the five posterior thoracic segments. The last muscle which I need here mention, is a deep-seated diverging pair, attached near the upper end of the stomach, on its ventral surface, and diverging from this point to the sides of the prosoma high up beneath the mouth. The probable action of this pair, as well as of the three superficial pairs of muscles by which the body is attached to the scuta, is to draw up the whole prosoma towards or from the orifice; and likewise to contract it firmly, so as to serve as a fulcrum for the movements of the five posterior thoracic segments, together with the cirri, which they carry.
The muscles of these five thoracic segments are numerous and powerful; they are also complicated, chiefly owing to the segments on their dorsal and dorso-lateral surfaces being driven, like wedges, one into the other. As far as I could make out, there are on each side three, superficial, dorso-lateral and lateral muscles (generally, if not always, destitute of striæ), which do not cross the articulations, but extend merely from articulation to articulation; and of which the function can be only to contract each separate segment, and consequently to open out the intermediate infolded articulations; the effect of this would be to separate slightly the cirri from each other,--more especially those borne on the two or three anterior segments, between which the infolded articulations are deeper or broader. There are other more deeply imbedded, powerful, long, dorso-lateral extensor, and ventri-lateral flexor muscles, attached at one end within the flanks of the prosoma, and at the other end to the successive segments of the thorax. The action of the former is to straighten and stretch out the thorax; of the latter, or ventri-lateral muscles, to retract it.
In tracing these muscles, a fascia could be seen to become attached to a segment, and then this same fascia would run on to the next succeeding segment: the effect of this must be, that each segment can be retracted and protracted either from the prosoma as a fulcrum, or from the antecedent segment as a fulcrum: we have, also, seen that each segment can, by the agency of the superficial, non-striated muscles, contract itself. Hence these thoracic segments are capable of diverse movements, as was very evident when the shell of a living specimen was opened. By one movement in common, the whole five posterior segments could be drawn back, so as to become even partly imbedded in the prosoma: lateral, twisting or wriggling movements were also quite distinct: the three posterior segments seemed to be capable of less independent movements than the anterior segments; and I observed that the more powerful flexor and extensor muscles did not run into these three posterior segments. The cirri, of course, partake of the movements of the thorax; and in watching, in an uninjured specimen, the alternate, protruding, gracefully sweeping and retracting movements of the posterior pairs of cirri, it was evident that the thorax was the chief agent in their movement. Besides the muscles now mentioned, there are some immediately to be noticed, which extend from within the thoracic segments to within the pedicels of the cirri.
_Movements and Muscles of the Cirri._ Although the cirri have not been described, it will be most convenient here to treat shortly of their muscles. Each cirrus consists of a pedicel, having a long basal and a short upper segment, supporting two multiarticulate rami. The lower segment of the pedicel can be drawn forward by an adductor muscle, attached low down within this segment, and crossing at right angles (at least in the case of the anterior cirri) the corresponding muscle of the opposed cirrus, on the central, ventral surface of the thorax. This segment can also be drawn back by a muscle springing from the dorso-lateral surface of the thorax, and running only a little way within the segment: I am far from sure that the lower segment does not possess other muscles. The short upper segment of the pedicel can be moved backwards and forwards, as I saw in living specimens, independently of the lower segment; this movement being best seen in the anterior cirri, which are much more often moved independently of each other than are the posterior cirri. The rami are capable, I believe, of being moved backwards and forwards _as a whole_, by the movement of the few lower segments, which are generally more or less confluent. They can, also, be curled up and uncurled by the combined movement of each separate segment. The uncurling seems to separate the two rami a little laterally. Each ramus, at least in the two or three anterior pairs, can be moved to a certain extent, independently of the other ramus of the same cirrus; and the few terminal segments, either of both rami or of one ramus, are often a little moved and curled (and this is especially the case with the long anterior ramus of the first pair), without the lower segments or the pedicel being moved.
The flexor and extensor muscles, which, as I believe, move the upper segment of the pedicel (_a_ and _b_, Pl. 29, fig. 1), are attached at their upper ends to its basal margin, and are thus enabled to draw it a little way down within the lower segment, and so move it. The short flexor muscle (_c_), which is attached at its lower end within the upper segment of the pedicel, and the longer extensor (_d_), also, attached within this same lower segment, serve, I believe, to move the lower, partially confluent segments of each ramus as a whole. In the case of these muscles, and of those last mentioned, I am surprised that the extensors (_b_) and (_d_) are not attached nearer to the exterior and dorsal surface. Other muscles (_e_, _f_) attached at their lower ends within the upper segment of the pedicel, run up each of the two rami to their tips, with some of the fasciæ terminating within each segment: of these muscles, the outer one (_f_, _f_) appears to be the extensor, and the inner one (_e_, _e_) the flexor. But besides these, there are other short flexor muscles (_g_, _g_) which run on the anterior face,[31] from segment to segment, serving to pull the front edge of one segment within the edge of the next lower segment. These muscles differ much in plainness in the several genera: they are very distinct in Coronula. In some specimens of this genus, a few of the articulations between the basal segments of the rami having been obliterated, the short muscles (_g_, _g_) running from articulation to articulation were absent, and their presence and nature in the upper segments thus rendered the plainer. The muscular system in the several pairs of cirri seems to be the same, with the exception of the first pair, in which the muscle answering, as I suppose, to (_a_), namely, the flexor of the upper segment of the pedicel, is much spread out at its lower end, and is there attached to the exterior surface of the lower segment.
When a Cirripede is alive, the action of the cirri is really beautiful: from the position of the thoracic segments, the posterior cirri (three pairs in the Balaninæ and four pairs in the Chthamalinæ) form a sort of semicircle facing the mouth: the anterior cirri stand further apart, and are opposed in pairs to each other, with the first pair pointing beyond the mouth. Together the cirri form a hollow cone, not circular but elongated, with the mouth situated at the lower anterior end. The posterior cirri are protruded, by the movement of the whole thorax, curled up, close along the carinal end of the orifice; as they are protruded, they diverge, both by the movement of their pedicels, and, as I believe, by the separation of the thoracic segments. As the two rami of each separate cirrus are uncurled, they also diverge a little; as do the double rows of spines on the segments in each ramus, by their elasticity. By the movement of the thorax, the cirri are then swept towards the rostrum; and, lastly, they are brought perpendicularly down towards the mouth with a rapid movement, which would be beautifully adapted to catch any object floating or swimming in the water; hence I have called the action of the cirri, captorial.
Ideas discussed in this chapter
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