The writings of Charles Darwin on the web

by John van Wyhe Ph.D.

Darwin, A monograph on the sub-class cirripedia, with figures of all the species. The lepadidae, or, pedunculated cirripedes. 1851.

-Front matter; preface; introduction (pp. -7)
-Family—lepadidæ; capitulum; Genus—lepas; pæcilasma. nov. genus
(pp. 8-115)
-Genus—dichelaspis; oxynaspis. gen. nov.; genus—conchoderma; Genus—alepas; anelasma. gen. nov.; Genus—ibla
(pp. 115- 214)
-Genus—scalpellum; [sub-carinâ nullâ.] 1. scalpellum vulgare; 2. scalpellum ornatum; 3. scalpellum rutilum; [sub-carinâ presente.] 4. scalpellum rostratum; 5. scalpellum peronii; 6. scalpellum villosum
(pp. 215-281)
-Summary on the nature and relations of the males and complemental males in ibla and scalpellum; Genus—pollicipes; Genus—lithotrya; Species mihi non satis notæ, aut dubiæ (pp. 281-375)
-Explanation of the plates
(pp. 377-393)
-Index (pp. 395-400)

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Cirripedia pedunculo flexili, musculis instructo: scutis* musculo adductore solummodô instructis: valvis cæteris, siguæ adsunt, in annulum immobilem haud conjunctis.

Cirripedia having a peduncle, flexible, and provided with mucles. Scuta* furnished only with an adductor muscle: other valves, when present, not united into an immovable ring.

Metamorphoses; larva, first stage, pp. 9—12; larva, second stage, p. 13; larva, last stage, p. 14; its carapace, ib.; acoustic organs, p. 15; antennaæ, ib.; eyes, p. 16; mouth, p. 17; thorax and limbs, p. 18; abdomen, p. 19; viscera, ib.; immature cirripéde, p. 20; homologies of parts, p. 25.

Description of mature Lepadidæ, p. 28; capitulum, ib.; peduncle, p. 31; attachment, p. 33; filamentary appendages, p. 38; shape of body, and muscular system, p. 39; mouth, ib.; cirri, p. 42; caudal appendages, p. 43; alimentary canal, 44; circulatory system, p. 46; nervous system, ib.; eyes, p. 49; olfactory organs, p. 52; acoustic(?) organs, p. 53; male sexual organs, p. 55; female organs, p. 56; ovigerous lamellæ, p. 58; ovigerous fræna, ib; exuviation, p. 61; rate of growth, ib.; size, ib.; affinities of family, p. 64; range and habitats, p. 65; geological history, p. 66.

Metamorphoses.—I will here briefly describe the Metamorphoses, as far as known, common to all Cirripedia, but more especially in relation to the present family. I may premise, that since Vaughan Thompson's capital discovery of the larvæ in the last stage of development in Balanus, much has been done on this subject: this same author subsequently published† in the 'Philosophical Transactions,' an account of the larvæ of Lepas and Conchoderma (Cineras) in the first stage ; and seeing how totally distinct they were from the larva of the latter stage in Balanus, he erroneously attributed the difference to

* The meaning of this and all other terms is given in the Introduction, at pp. 3-7.

† Philosophical Transactions, 1835, p. 355, Pl. VI.

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the difference in the two families, instead of to the stage of development. Burmeister* first showed, and the discovery is an important one, that in Lepas the larvæ pass through two totally different stages. This has subsequently been proved by implication to be the case in Balanus, by Goodsir,† who has given excellent drawings of the larva in the first stage; and quite lately, Mr. C. Spence Bate, of Swansea, has made other detailed observations and drawings of the larvæ of five species in this same early stage, and has most kindly permitted me to quote from his unpublished paper‡. I am enabled to confirm and generalise these observations, in all the Cirripedes in the Order containing the Balanidæ and Lepandidæ.

The ova, and consequently the larvæ of the Lepadidæ, in the First Stage, whilst within the sack of the parent, vary in length from ·007 to ·009 in Lepas, to ·023 of an inch in Scalpellum: my chief examination of these larvæ has been confined to those of Scalpellum vulgare; but I saw them in all the other genera. The larva is somewhat depressed, but nearly globular; the carapace anteriorly is truncated, with lateral horns; the sternal surface is flat and broad, and formed of thinner membrane than the dorsal. The horns just alluded to are long in Lepas and short in Scalpellum; their ends are either rounded and excessively transparent, or, as in Ibla, furnished with an abrupt, minute, sharp point: within these horns, I distinctly saw a long filiformed organ, bearing excessively fine hairs in lines, so exactly like the long plumose spines on the prehensile antennæ of the larvæ in the last stage; that I have not the least doubt, that these horns are the cases in which antennæ are in process of formation. Pos-

* Beiträge zur Naturgeschichte der Rankenfüsser, 1834. Mr. J. E. Gray, however, briefly described, in 1833, (Proceedings, Zoological Society, October,) the larva in the first stage of Balanus; in this notice the anterior end of the larva is described as the posterior.

† Edinburgh New Philosophical Journal, July 1843, Pls. iii and iv.

‡ This will appear in the October number (1851) of the 'Annals of Natural History.'

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teriorly to them, on the sternal surface, near each other, there are two other minute, doubly curved, pointed horns, about ·004 in length, directed posteriorly; and within these I again saw a most delicate articulated filiformed organ on a thicker pedicel: in an excellent drawing, by Mr. C. S. Bate, of the larva of a Chthamalus (Balanus punctatus of British authors), after having kept alive and moulted once, these organs are distinctly shown as articulated antennæ (without a case), directed forwards: hence, before the first moult in Scalpellum, we have two pair of antennæ in process of formation. Anteriorly to the bases of these smaller antennæ is seated the heart-shaped eye, (as I believe it to be,) ·001 in diameter, with apparently a single lens, surrounded, except at the apex, by dark-reddish pigment-cells. In some cases, as in some species of Lepas, the larvæ when first excluded from the egg, have not an eye, or a very imperfect one.

There are three pairs of limbs, seated close together in a longitudinal line, but some way apart in a transverse direction: the first pair always consists of a single spinose ramus, it is not articulated in Scalpellum, but is multi-articulate in some genera; it is directed forwards. The other two pair have each two rami, supported on a common haunch or pedicel: in both pair, the longer, ramus is multi-articulate, and the shorter ramus is without articulations, or with only traces of them: the longer spines borne on these limbs (at least, in Scalpellum and Chthamalus), are finely plumose. The abdomen terminates, a little beyond the posterior end of the carapace, in a slightly upturned horny point; a short distance anteriorly to this point, a strong, spinose, forked projection depends from the abdominal surface.

Messrs. V. Thompson, Goodsir, and Bate, have kept alive for several days the larvæ of Lepas, Conchoderma, Balanus, Verruca, and Chthamalus, and have described the changes which supervene between the first and third exuviations. The most conspicuous new character is the

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great elongation of the posterior point of the carapace into an almost filiform, spinose point in Lepas, Conchoderma, Chthamalus, and Balanus, but not according to Goodsir, in one of the species of the latter genus. The posterior point, also, of the abdomen becomes developed in Balanus (Goodsir) into two very long, spear-like processes, serrated on their outer sides; in Lepas and Conchoderma, according to Thompson, into a single, tapering spinose projection; and in Chthamalus, as figured by Mr. Bate, the posterior bifid point, as well as the depending ventral fork, increase much in size. Another important change, which has been particularly attended to by Mr. Bate, is the appearance of spinose projections and spines (some of which are thick, curved, and strongly plumose, or, almost pectinated along their inner sides) on the pedicels and lower segments of the shorter rami of the two posterior pairs of limbs.

The mouth in its earliest condition alone remains to be described; in S. vulgare, it is seated on a very slight prominence, in a most remarkable situation, namely, in a central point between the bases of the three pairs of legs. I traced by dissection the oesophagus for some little way, until lost in the cellular and oily matter filling the whole animal, and it was directed anteriorly, which is the direction that might have been expected, from the course followed by the oesophagus in the larva in the last stage, and in mature Cirripedes. Mr. A. Hancock has called my attention to a probosciformed projection on the under side of the larva of Lepas fascicularis, when just escaped from the egg. Mr. Bate has described this same proboscis in Balanus and Chthamalus, and states the important fact, that it is capable of being moved by the animal; and, lastly, I have seen it in an Australian Chthamalus, and in Ibla, of remarkable size. This proboscis, which is always directed posteriorly, (like the mouth in the mature animal,) certainly answers to the mouth as made out by dissection in Scalpellum; and I believe I saw, as has Mr. Bate, a terminal orifice: it certainly does

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not possess any trophi. In Ibla (in which the larva is large enough for dissection), the base of the proboscis arises posteriorly to the first pair of legs, and the orifice at the other end reaches beyond or posteriorly to the point, where the mouth in Scalpellum opens, namely between the middle pair of legs. The mouth being either so largely probosciformed or seated only on a slight eminence, in two genera so closely allied as Ibla and Scalpellum, and (judging from Mr. Thompson's figures, and from what I have seen myself,) in the species of the same genus Lepas, is a singular difference: in the cases in which, at first, the proboscis is absent, it would probably soon be developed. I cannot but suppose that the inwardly directed spines on the bases of the two posterior legs, which are so rapidly developed, serve some important end, namely, as organs of prehension for the larvæ like the mandibles and maxillæ of mature Cirripedes, for seizing their prey, and conveying it to their moveable mouths, conveniently seated for this purpose.

The first pair of legs answers, as I believe from reasons hereafter to be assigned, to the outer pair of maxillipods in the higher crustacea; and the other four legs to the first two pair of thoracic limbs in these same crustacea; this being the case, the highly remarkable position of the mouth in the larva, either between the bases of the two posterior pair of legs, or at least posteriorly to the first pair, together with the probable functions of the spiny points springing from the basal segments of the two hinder pair of true thoracic limbs, forcibly bring to mind the anomalous structure of the mouth being situated in the middle of the under side of the thorax, in Limulus,—that most ancient of crustaceans, and therefore one likely to exhibit a structure now embryonic in other orders. I will only further remark, that I suspect that the truncation of the anterior end of the carapace, has been effected by the segments having been driven inwards, and consequently, that the larger antennæ within the lateral horns, though standing more in front than the

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little approximate pair, are normally the posterior of the two pair. According to Milne Edwards, the posterior pair are normally seated outside the anterior pair, and this is the case with those within the lateral horns.

Larva in the Second Stage.—Notwithstanding the considerable changes, already briefly given, which the larva undergoes during the first two or three exuviations after leaving the egg, all these forms may be conveniently classed under the first stage. The larva in the Second stage is known only from a single specimen described, figured, and found by Burmeister,* adhering to sea-weed in the midst of other larvæ of Lepas in the last stage. In its general shape and compressed form, it seems to come nearer to the last than to the first stage. It has only three pair of legs, situated much more posteriorly on the body than in the first stage, and all directed posteriorly; they are much shorter than heretofore, and resemble rather closely those of the last stage, with the important exception that the first pair has only one ramus. It is this circumstance which leaves no doubt on my mind, that we here have the three pair of limbs, of the first stage metamorphosed. The body is prolonged some way behind these limbs, and ends in a blunt, rounded point, in which, probably, are developed the three posterior pair of legs and the abdomen of the larva in the last stage. The mouth is now seated some way anteriorly to the limbs, is large and probosciformed, and is, I presume, still destitute of trophi. There are now two closely approximate eyes, but as yet both are simple. The smaller pair of antennæ has disappeared. The whole animal was attached to the sea-weed by a (I presume, pair of,) "fleischigen Fortsatz," which Burmeister considers as the prehensile antennæ to be presently described, in an early state of development. I have little doubt that this is correct, for in an abnormal Cirripede of another order, in which the larva appears in the first stage with prehensile antennæ, the eggs have two great projecting horns including

* Beiträge zur Naturgeschichte der Rankenfüsser, s. 16, Tab. i, figs. 3, 4.

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these organs, and attached by their tips, through some unknown means, to the sack of the parent, apparently in the same manner as Burmeister's larva was attached to the sea-weed. I will only further remark on the larva of this Second stage, that its chief development since the first stage, has been towards its anterior end. The next great development, to be immediately described, is towards the posterior end of the animal.

Larva, Last Stage.—My chief examination has been directed, at this stage of development, to the larvæ of Lepas australis, which are of unusual size, namely, from ·065 to even almost ·1 of an inch in length; I examined, however, the larvæ of several other species of Lepas, of Ibla and of Balanus, with less care, but sufficiently to show that in all essential points of organisation they were identical; this, indeed, might have been inferred from the similarity of the larval prehensile antennæ, preserved in the bases of all mature Cirripedes, and which I have carefully inspected in almost every genus. The larvæ in this final stage, in most of the genera, have increased many times in size since their exclusion from the egg; for instance, in Lepas australis, from ·007 to ·065, or even to ·1 of an inch. They are now much compressed, nearly of the shape of a cypris or mussel-shell, with the anterior end the thickest, the sternal surface nearly or quite straight, and the dorsal arched. Almost the whole of what is externally visible consists of the carapace; for the thorax and limbs are hidden and enclosed by its backward prolongation; and even at the anterior end of the animal, the narrow sternal surface can be drawn up, so as to be likewise enclosed. As in several Stomapod crustaceans, the part of the head bearing the antennæ and organs of sense, in front of the mouth, equals, or even exceeds in length, and more than exceeds in bulk, the posterior part of the body, consisting of the enclosed thorax and abdomen. I will now briefly describe, in the following order, the carapace, the organs of sense, mouth, thorax and limbs, abdomen, and internal viscera.

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The form of the Carapace has been sufficiently described; it consists of thick chitine membrane, marked with lines, and sometimes with stars and other patterns; it is obscurely divided into two halves by a line or suture along part of the dorsal margin; these halves or two valves are drawn together by an adductor muscle, in the same relative position as in the mature Cirripede. The part overhanging and enclosing the thorax is lined by an excessively delicate membrane, obviously homologous with the lining of the sack in the mature animal, and is nothing but a duplicature of the carapace, rendered very thin from being on the under or protected side: a layer of true skin or corium, probably double, separates these two folds.

Acoustic Organs.—On the borders of the carapace, at the anterior end, on the sternal surface, there are two minute orifices, in L. australis ·002 in diameter, sometimes having a distinct border round them; the membrane of the carapace on the inside is prolonged upwards and inwards in two short funnel-shaped tubes, lodged in closed sacks of the corium: within these sacks on each side a delicate bag is suspended, and hangs in the mouth of the above funnel; at the upper end a large nerve could be distinctly seen to enter the bag: I cannot doubt that this is a sense-organ; from its position and from the animal not feeding (as we shall presently see), I conclude that it is an acoustic organ.

Antennæ.—These are large and conspicuous; they are attached very obliquely on the sternal surface, a little way from the anterior end of the carapace, beyond which, when exserted, they extend;* they can (at least in Ibla)

* Mr. J. D. Dana, who has examined these organs in the larvæ of Lepas, informs me in a letter, that in his opinion they correspond with the inferior antennæ the superior being wanting, as in most Daphnidæ." He continues—"I know of no case in which the inferior are obsolete when the superior are developed; but the reverse is often true." In position these antennæ certainly correspond to the inferior and central pair of the larva in the first stage, which belong, as it would appear, to the first segment of the body; but judging from the drawing by Burmeister of the larva in the second stage, I am, in some respects, more inclined to consider that they correspond to the larger pair seen within the lateral horns of the carapace in the first stage.

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be retracted within the carapace. They consist of three segments: the first or basal one is much larger than the others, and apparently always has a single spine on the outer distal margin. The second segment consists either of a large, thin, circular, sucking disc, or is hoof-like (Tab. V, figs. 5, 10, 11, 12); in all cases it is furnished with one or more spines, (seven very long ones in Lepas,) on the exterior-hinder margin. The third and ultimate segment is small; it is articulated on the upper surface of the disc, and is directed rectangularly outwards; it is sometimes notched, and even shows traces of being bifid; it bears about seven spines at the end; some of these spines are hooked, others simple, and in Lepas and Conchoderma, two or three are very long, highly flexible, and plumose, a double row of excessively fine hairs being articulated on them. I can hardly doubt that these latter spines, (within which the purple corium could be seen to enter a little way,) floating laterally outwards, serve as feelers. The antennaæ, at first, are well furnished with muscles. They serve, in Lepas, according to Mr. King, and in Balanus, according to Mr. Bate, and as I saw myself in another unnamed order, for the purpose of walking, one limb being stretched out before the other; but their main function is to attach the larva for its final metamorphosis into a Cirripede. The disc can adhere even to so smooth a surface as a glass tumbler.* The attachment is at first manifestly voluntary, but soon becomes involuntary and permanent, being effected by special and most remarkable means, which will be most conveniently described in a later part of this Introduction. I will here only state that I traced with ease the two cement-ducts running from two large glandular bodies, to within the antennæ up to the discs.

Eyes.—Close behind the basal articulations of the antennæ, the sternal surface consists of two approximate, elongated, narrow, flat pieces, or segments. These

* Rev. R. L. King. Annual Report of R. Institution of Cornwall, 1848, p. 55.

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Burmeister considers as the basal segments of the antennæ: as they are not cylindrical, I do not see the grounds for this conclusion: their posterior ends are rounded, and the membrane forming them is reflected inwards, in the form of two, forked, horny apodemes, together resembling two letters, UU, close together; these project up, inside the animal, for at least one third of its thickness from the sternal to the dorsal surface. The two great, almost spherical eyes in L. australis, each 1/150th of an inch in diameter, are attached to the outer arms, thus, ·UU· , in the position of the two full stops. Hence, the eyes are included within the carapace. Each eye consists of eight or ten lenses, varying in diameter in the same individual from 1/2000 to 3/2000th of an inch, enclosed in a common membranous bag or cornea, and thus attached to the outer apodemes. The lenses are surrounded half way up by a layer of dark pigment-cells. The nerve does not enter the bluntly-pointed basal end of the common eye, but on one side of the apodeme. The structure here described is exactly that found, according to Milne Edwards, in certain crustacea. In specimens just attached, in which no absorption has taken place, two long muscles with transverse striæ may be found attached to the knobbed tips of the two middle arms of the two ·UU·, and running up to the antero-dorsal surface of the carapace, where they are attached; other muscles (without transverse striæ) are attached round the bases, on both sides of both forks. The action of these muscles would inevitably move the eyes, but I suspect that their function may be to draw up the narrow, deeply folded, sternal surface, and thus cause the retraction of the great prehensile antennæ within the carapace.

Mouth.—This is seated in exactly the same position as in the mature Cirripede, on a slight prominence, fronting the thoracic limbs, and so far within the carapace, that it was obviously quite unfitted for the seizure of prey; and it was equally obvious, that the limbs were natatory, and incapable of carrying food to the mouth.


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This enigma was at once explained by an examination of the mouth, which was found to be in a rudimentary condition and absolutely closed, so that there would be no use in prey being seized. Underneath this slightly prominent and closed mouth, I found all the masticatory organs of a Cirripede, in an immature condition. The state of the mouth will be at once understood, if we suppose very fluid matter to be poured over the protuberant mouth of a Cirripede, so as to run a little way down, in the shape of internal crests, between the different parts, and in the shape of a short, shrivelled, certainly closed tube, a little way (·008 of an inch in L. australis) down the oesophagus: Hence, the larva in in this, its last stage, cannot eat; it may be called a locomotive Pupa;* its whole organisation is apparently adapted for the one great end of finding a proper site for its attachment and final metamorphosis.

Thorax and Limbs.—The thorax is much compressed, and consists of six segments, corresponding with the six pair of natatory legs; the anterior segments are much plainer (even the first being distinctly separated a fold from the mouth), than the posterior segments, which is exactly the reverse of what takes place in the mature Cirripede; in the latter, the first segment is confounded with the part bearing the mouth. The epimeral elements of the thorax are distinguishable; the sternal surface is very narrow, and is covered with complicated folds and ridges. The six pair of legs are all close, one behind the other, and all are alike in having a haunch or pedicel of two segments, directed forwards, bearing two arms or rami, each composed of two segments, the outer ramus

* M. Dujardin has lately ('Comptes Rendus,' Feb. 5, 1850, as cited in 'Annals of Nat. History,' vol. v, p. 318,) discovered that the "Hypopi are Acari with eight feet, without either mouth or intestine, and which, being deprived of all means of alimentation, fix themselves at will, so as to undergo a final metamorphosis, and they become Gamasi or Uropodi." Here, then, we have an almost exactly analogous case. M. Dujardin asks—"Ought, therefore, the Hypopi to be called larvæ, when, under that denomination, have hitherto been comprised animals capable of nourishing themselves?"

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being a little longer than the inner one. On the lower segments in both rami of all the limbs, there is a single spine. In all the limbs, the obliquely truncated summit of the terminal segment of the inner ramus bears three very long, beautifully plumose spines: in the first pair, the summit of the outer ramus bears four, and in the five succeeding pair, six similar spines. This difference, small as it is, is interesting, as recalling the much greater difference between the first and succeeding pairs, in the first and second stage of development. The terminal segments of all the rami, bearing the long plumose spines, are directed backwards. The limbs and thorax are well furnished with striated muscles. The animal, according to Mr. King, swims with great rapidity, back downwards. The limbs can be withdrawn within the carapace.

Abdomen and Caudal Appendages.—The abdomen is small, and its structure might easily be overlooked without careful dissection of the different parts: it consists of three segments; the first can be seen to be distinct from the last thoracic segment, bearing the sixth pair of limbs, only from the fold of the epimeral element, and from its difference in shape; the second segment is very short, but quite distinct; the third is four or five times as long as the second, and bears at the end two little appendages, each consisting of two segments, the lower one with a single spine, and the upper one with three, very long, plumose spines, like those on the rami of the thoracic limbs. The abdomen contains only the rectum and two delicate muscles running into the two appendages, between the bases of which the anus is seated.

Internal Viscera.—Within the body, in front of the mouth, it was easy to find the stomach (with two pear-shaped cæca at the upper end), running first anteriorly, and then curving back and reaching the anus by a long rectum, difficult to be followed: it appeared, however, to me, that this stomach had more relation to the young Cirripede, of which every part could now generally be traced, than to the larva, with its closed and rudimentary

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mouth: the fact, however, of its being prolonged to the anus, which is in a different position in the larva and mature state, shows that the stomach serves, at least, as an excretory channel. Besides the stomach, the several muscles already alluded to, and much pulpy and oily matter, the only other internal organs consist of two long, rather thick, gut-formed masses, into the anterior ends of which the cement-ducts running from the prehensile antennæ could be traced. These masses are formed of irregular orange balls, about ·001 of an inch in diameter, made up of rather large cells, so to have a grape-like appearance, held together by a transparent pale yellowish substance, but apparently not enclosed in a membrane: these masses lie rather obliquely, and approach each other at their anterior ends; they extend from above the compound eyes, to the cæca of the stomach to which they cohere, but in young specimens, they extend some way beyond the cæca, between the folds of the carapace. The two cement-ducts, at the points where they enter these bodies, expand and are lost; at this point, also, the little orange-coloured masses of cells have the appearance of being broken down into a finer substance. Within the cement-ducts I saw a distinct chord of rather opaque cellular matter. We shall presently see, that these gut-formed masses are the incipient ovaria.

The Young Cirripede within the Larva.—Several times I succeeded in dissecting off the integuments of the lately-attached larva, and in diplaying the young Lepas australis entire. The following description applies to the Cirripede in this state; but for convenience sake, I shall occasionally refer to its condition when a little more advanced. I may premise, and the fact in itself is curious, that the bivalve-like shell of the larva, together with the compound eyes, is first moulted, and some time afterwards, the inner lining of the sack, together with the integuments of the thorax and of the natatory legs: hence, I often found specimens, which externally seemed to have perfected their metamorphoses, but which, within their

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sacks, retained all the characters of the natatory larva. According to Mr. King, the larva of Lepas throws off its external shell five days after becoming attached. Whilst the young Lepas is closely packed within the larva, the capitulum, as known by the five valves, about equals in length the peduncle. The peduncle occupies the anterior half of the larva; when fully stretched, it becomes narrower and slightly longer than the capitulum; the separation between the capitulum and peduncle is almost arbitrary in the mature animal, and corresponds with no particular line in the larva. Even at this early period, the muscles of the peduncle are quite distinct. No vestige is preserved in the outer integument, of the sternal and dorsal sutures of the larval carapace; but in the corium of the peduncle, three coloured marks which occur near the eyes, and two little curled marks which occur near the acoustic orifices of the larva, are all preserved for some time after maturity. The compound eyes, as we have seen, are attached to apodemes, springing from the sternal surface of the larval carapace, and are consequently cast off with it: whilst the young Cirripede is packed within the larva, the outer integument of its peduncle necessarily forms a deep transverse fold passing over the eyes and apodemes, and this, as we shall presently see, plays an important part in the future position of the animal. Tha antennæ are not moulted with the carapace, but left cemented to the surface of attachment; their muscles are converted into sinewy fibres, the corium after a short period is absorbed, and they are then preserved in a functionless condition. No trace of the two acoustic sacks can be perceived in the corium of the young Cirripede, excepting the coloured marks above alluded to.

In the young capitulum, the five valves stand some way apart from each other; they are elegant objects under the microscope; they are not calcified, but consist exclusively of chitine; they are rather thick, composed of an outer membrane lined by hexagonal prisms,

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quite unlike any other membrane in the animal. These valves, which I have called primordial valves, resemble pretty closely in shape the valves of the mature animal; the fork of the carina, however, is indicated only by a slight constriction above the lower end. After the exuviation of the larval integuments, and when calcification commences, the first layer of shell is deposited under, and then round these primordial valves. The latter, in well preserved old specimens, may often be detected on the umbones of the scuta, terga, and carina, but not on the umbones of any other valves.

The mouth seems one of the earliest parts developed: in the youngest larva dissected, I could make out at least points corresponding with each organ; and, at the period when the young Cirripede could be dissected out of its larval envelopes, their general details were quite plain. The labrum, however, had not become bullate. The mouth, as we have seen, is formed under the rudimentary mouth of the larva, and at the same relative spot occupied by the probosciformed mouth of the larva in the second stage. Thus far, in the young Cirripede and larva, there has been no great change in the relative positions of the parts: the rudimentary eyes, however, of the former are developed posteriorly to (or above, as applied to a Cirripede,) the cast-off compound eyes of the larva; but the position of the mouth, of the antennæ and of the several coloured marks in the corium, prove to demonstration, the correspondence in both of part to part. The case is rather different with what follows.

The Cirri are developed at first of considerable length, so that the young animal may soon provide itself with food; in Lepas australis they are of great length, the sixth pair consisting of seventeen or eighteen obscure segments. The extreme tips of the twenty-four rami of the six pair of cirri, are formed within the twenty-four, corresponding, little, bi-segmental rami of the six pair of natatory legs; but as the cirri are many times longer than these legs, they occupy in a bundle the

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whole thorax of the larva; no part whatever of the thorax of the Cirripede is formed within the thorax of the larva, but (together with the pedicels of the anterior cirri) within the cephalic cavity. As a consequence of this, the longitudinal axis of the thorax of the young Cirripede lies almost transversely to the longitudinal axis of the larva; and the Cirripede, from this transverse position of its thorax, comes to be, as it were, internally, almost cut in twain, and the sack thus produced. As soon as the young Cirripede is free and can move itself, the cirri are curled up, and the thorax is advanced towards the orifice of the capitulum, its longitudinal axis resuming the position of approximate parallelism to the longitudinal axis of the whole body, which it had in the larval condition. The reader will, perhaps, understand what I mean, if he will look at the mature Cirripede, figured in Pl. IX, fig. 4. In this, he will see that the body or thorax is united to the peduncle only by a small part below the mouth; on the other hand, if he imagines the whole bottom of the body (as high up as the letter h) united and blended into the peduncle, he will see the state in which these parts exist in the larva. Now, let him greatly shorten the cirri, so as to resemble the natatory legs of the larva, and then imagine a young Cirripede, with cirri of full length, formed within the old one, he will see that the new thorax supporting the cirri will have to be developed in an almost transverse position,—the animal consequently being internally almost separated into twain.

Of the internal organs, whilst the Cirripede is still within the larva, I have already mentioned the stomach with its pair of cæca: from the retracted position of the thorax and rudimentary abdomen, and consequently of the anus, compared with these parts in the larva, the alimentary canal is not above half its former length. There is, as yet, no trace of the filaments supposed by some to act as branchiæ, at the base of the first pair of cirri. Nor could I perceive a trace of the testes or

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vesiculæ seminales: the penis is represented by a minute, apparently imperforate projection. I have already briefly described the pair of large, gut-formed bodies in the larva, into the anterior ends of which the cement-ducts ran, and evidently derived their slightly opaque, cellular contents. At a very early age, before the young Cirripede can be distinctly made out, the posterior ends of these gut-formed bodies are absorbed, so as not to pass beyond the cæca of the stomach;. When the young Cirripede is plainly developed within the larva, these bodies in a relatively reduced condition are still distinct near the cæca, and at the opposite or anterior end (i.e. lower, in the position in which Cirripedes are usually figured), they have branched out into a sheet of delicate inosculating tubes; these could be traced by every stage, until, in the young perfected Cirripede, they filled the peduncle as ordinary ovarian tubes. In the larva, the two gut-formed bodies or incipient ovaria keep of equal thickness from one to the other end, but in the mature Cirripede, the ovarian tubes in the peduncle and the small, glandular, grape-like masses, near the stomach-cæca, are connected only by a delicate tube; this I failed in tracing in specimens in the very immature condition of those now under description.

The larva fixes itself with its sternal surface parallel and close to the surface of attachment, and the antennæ become cemented to it: if the Cirripede, after its metamorphosis had remained in this position, the cirri could not have been exserted, or only against the surface of attachment; but there is a special provision, that the young Cirripede shall immediately assume its proper position at right angles to the position which it held whilst within the larva, namely with its posterior end upwards. This is effected in a singular manner by the exuviation of the great compound eyes, which we have seen are fastened to the outer arms of the double ·UU·-like, sternal apodemes: these together with the eyes stretch transversely across, and internally far up into,

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the body of the larva; and, as the whole has to be rejected or moulted, the membrane of the peduncle of the young Cirripede has necessarily to be formed with a wide and deep inward fold, extending transversely across it; this when stretched open, after the exuviation of the larval carapace and apodemes, necessarily causes the sternal side of the peduncle to be longer than the dorsal, and, as a consequence, gives to the young Cirripede its normal position, at right angles to that of the larva when first attached.


I may here state, that I have examined the larvæ in this the final or perfect stage in four species of Lepas, in Conchodermavirgata, Ibla quadrivalvis, and, though rather less minutely, in Balanus balanoides, and I find all essential points of organisation similar. With the exception of diversities in the proportional sizes of the different parts, and in the patterns on the carapace, the differences, even in the arrangement of the spines on the limbs and antennæ, are less than I should have anticipated.

I have in this abstract treated the metamorphoses at greater length than I should otherwise have done, on account of the great importance of arriving at a correct homological interpretation of the different parts of the mature animal. In Crustacea, according to the ordinary view, there are twenty-one segments; of these I can recognise in the Cirripede, on evidence as good as can generally be obtained, all with the exception of the four terminal abdominal segments; these do not occur in any species known to me, in any stage of its development. If that part of the larva in front of the mouth, bearing the eyes, the prehensile antennæ, and in an earlier stage two pair of antennæ, be formed, as is admitted in all other Crustacea, of three segments, then beyond a doubt, from the absolute correspondence of every part, and even every coloured mark, the peduncle of the Lepadidæ is likewise thus formed. The peduncle being filled by the branching ovarian tubes is no objection to this view, for I am

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informed on the high authority of Mr. J. D. Dana,* that this is the case with the cephalo-thorax in some true Crustaceans, for instance, in Sapphirina. To proceed the mouth, formed of mandibles, maxillæ, and outer maxilleæ, correspond with the fourth, fifth, and sixth segments of the archetype Crustacean. Posteriorly to the mouth, we come, in the larva, to a rather wide interspace without any apparent articulation or organ, and then to the thorax, formed of six segments, bearing the six pair of limbs, of which the first pair differs slightly from the others. The thorax is succeeded by three small segments, differently shaped, with the posterior one alone bearing appendages; these segments, I cannot doubt, from their appearance alone, and from their apparent function of steering the body, are abdominal segments. If this latter view be correct, the thoracic segments are the six posterior ones of the normal seven segments, and there must be two segments missing between the outer maxillæ and first thoracic pair of legs, which latter on this view springs from the ninth segment. Now, in a very singular Cirripede, already alluded to under the name of Proteolepas, the two missing segments are present, the mouth being actually succeeded by eight segments, and these by the three usual abdominal segments,—every segment in the body being as distinct as in an Annelid: hence in Proteolepas, adding the three segments for the mouth and three for the carapace, we have altogether

* This distinguished naturalist has given his opinion in the 'American Journal of Science,' March, 1846, that "the pedicel of Anatifa corresponds to a pair of antennæ in the young;" although the peduncle or pedicel is undoubtedly thus terminated, even in mature individuals, I think it has been shown that it is the whole of the anterior part of the larva in front of the mouth, which is directly converted into the peduncle. Professor E. Forbes, in his Lectures, and Professor Steenstrup, in his 'Untersuchungen über das vorkommen des Hermaphroditismus in der Natur,' ch. v, have considered the peduncle as a pair of fused legs. Lovén has taken, judging from a single sentence, the same view of the homologies of the external parts as I have done; in his description of Alepas squalicola, (Ofversigt of Kongl. Vetens., &c, Stockholm, 1844, pp. 192-4), he uses the following words: "Capitis reliquæ partes, ut in Lepadibus semper, in pedunculum mutatæ et invoucrum," &c.; his involucrum is the same as the capitulum of this work.

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seventeen segments, which, as I stated, is the full number ever observed in any Cirripede, the four missing ones being abdominal, and, I presume, the four terminal segments. That the cavity in which the thorax is lodged, in the larva and therefore in the mature Cirripede, is simply formed by the backward production of the carapace, does not require any discussion. The valves have no homological signification.

As we have just seen that the first pair of natatory legs is borne on the ninth segment of the body, so it must be with the first pair of cirri, which consequently correspond to the outer maxillipods (the two inner pair of maxillipods or pied-machoires being here aborted) of the higher Crustacea, and hence their difference from the five posterior pair, which correspond with the five, ordinary pair of ambulatory legs in these same Crustacea. The part of the body, which I have called the prosoma, that is the protuberant, non-articulated, lower part of the thorax (Pl. IX, fig. 4 n), is a special development, either of the ninth segment, bearing the first pair of cirri, or of the segments corresponding with the organs of the mouth. The three abdominal segments of the larva are represented in the mature Cirripede, in the Order containing the Lepadidæ, only by a minute, triangular gusset, let in between the V-shaped tergal arches of the last thoracic segment: in this gusset, small as it is, is seated the anus, and on each side the caudal appendages, often rudimentary and sometimes absent. In another order, I may remark, (including, probably, the Alcippe of Mr. Hancock,) the cirri, of which there are only three pair, are abdominal.

I feel much confidence, that the homologies here given are correct. The cause of their having been generally overlooked arises, I believe, from the peculiar manner, already described, in which the animal, during its last metamorphosis, is internally almost intersected: even for some little time after discovering that the larval antennæ were always embedded in the centre of the surface of attachment, I did not perceive, that this was the anterior

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end of the whole animal. The accompanying woodcut gives at a glance, a view of the homologies of the external parts: the upper figure (from Milne Edwards) is a


Stomapod Crustacean, Leucifer of Vaughan Thompson, and the abdomen, which we know becomes in Cirripedes, after the metamorphosis, rudimentary, and therefore does not fairly enter into the comparison, is given only in faint lines: the lower figure is a mature Lepas, with the antennæ and eyes, which are actually present in the larva, retained and supposed to have gone on growing. All that we externally see of a Cirripede, whether pedunculated or sessile, is the three anterior segments of the head of a Crustacean, with its anterior end permanently cemented to a surface of attachment, and with its posterior end projecting vertically from it.



I will now proceed to a general description of the different parts and organs in the Lepadidæ. The Capitulum is usually much flattened, but sometimes broadly oval in section. It is generally formed of five or more valves, connected together by very narrow or broad strips of membrane; sometimes the valves are rudimental or absent, when the whole consists of membrane. When the valves are numerous, and they occassionally ex-

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ceed a hundred in number, they are arranged in whorls, with each valve generally so placed as to cover the interval between the two valves above. Of all the valves, the scuta are the most persistent; then come the terga, and then the carina; the rostrum and latera occur only in Scalpellum and Pollicipes, and in a rudimentary condition in Lithotrya, and, perhaps, in the fossil genus Loricula. The valves are formed sometimes of chitine (as in Ibla and Alepas), but usually of shell, which varies from transparency to entire opacity. The shell is generally white, occassionally reddish or purple; exteriorly, the valves are covered by more or less persistent, generally yellow, strong membrane. The scuta and terga are always considerably larger than the other valves: in the different genera the valves differ so much in shape that little can be predicated of them in common; even the direction of their lines of growth differs,—thus, in Lepas and some allied genera, the chief growth of the scuta and of the carina is upwards, whereas in Pollicipes and Lithotrya, it is entirely downwards; in Oxynaspis, and some species of Scalpellum, it is both upwards and downwards. Even in the same species, there is often very considerable variation in the exact shape of the valves, more especially of the terga. The adductor muscle is always attached to a point not far from the middle of the scuta, and it generally has a pit for its attachment. In several genera, namely, Pæcilasma, Dichelaspis, Conchoderma, and Alepas, the scuta show a tendency to be bilobed or trilobed. The valves are placed either at some distance from each other, or close together; but their growing margins very rarely overlap each other, though this is sometimes the case with their upper, free, tile-like apices; in a few species the scuta and terga are articulated together, or united by a fold. The membrane connecting the valves, where they do not touch each other, is like that forming the peduncle, and is sometimes brilliantly coloured crimson-red; generally, it appears blueish-gray, from the corium being seen through. Small pointed

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spines, connected with the underlying corium by tubuli, are not unfrequently articulated on this membrane: the tubuli, however, are often present where there are no spines. To allow of the growth of the capitulum, the membrane between the valves splits at each period of exuviation, when a new strip of membrane is formed beneath, connected on each side with a fresh layer of shell,—the old and outer slips of membrane disintegrating and disappearing: when there are many valves, the line of splitting is singularly complicated. This membrane consists of chitine,* and is composed of numerous fine laminæ. After the valves have been placed in acid, a residue, very different in bulk in different genera, is left, also composed of successive laminæ of chitine. It appears to me that each single lamina of calcified chitine, composing the shell, must once have been continuous with a non-calcified lamina in the membrane connecting the several valves: at the line where this change in calcification supervenes; the chitine generally assumes some

* Chitine is confined to the Articulata. It was Dr. C. Schmidt (Contributions, &c., being a Physiologico-Chemical investigation: in Taylor's 'Scientific Memoirs,' vol. v), who discovered that the membrane connecting the valves and forming the peduncle, and the tissues of the internal animal, were composed of this substance. But Dr. Schmidt says that the valves in Lepas are composed of 3·09 of albuminates, and 96·81 of incombustible residue; I cannot but think that the existence of the albuminates is an error caused by Dr. Schmidt's belief that the Cirripedia were intermediate between Crustacea and Mollusca, in the shells of which latter, the animal basis consists of albuminates. For after placing the valves of Lepas and Pollicipes in cold acid, I found that the membrane left could not be dissolved in boiling caustic potash, but could, though slowly, (and without change of colour,) in boiling muriatic acid; and these are the main diagnostic characters of Chitine, compared with albuminous substances. I may add, that Schmidt was also induced to consider the shells of Cirripedia as having the same nature with those of Mollusca, from finding that in the above 96·81 of incombustible matter, 99·3 consisted of carbonate and only 0·7 of phosphate of lime; but Dr. Schmidt's own analyses prove how extremely variable the proportions of these salts are in the Crustacea, as the following instance shows:—

Lobster. Squilla.

Phosphate of Lime . . 12·06 . . 47·52
Carbonate of Lime . . 87·94 . . 52·48

And, therefore, it is not very surprising that Cirripedia should have still less phosphate of lime in their shells, than has a lobster compared with a squilla.

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colour, and becomes much harder and more persistent; and as the whole valve is formed of component laminæ thus edged (the once continuous laminæ of non-calcified chitine connecting the valves, having disintegrated and disappeared) the surfaces of the valves are generally left covered by a persistent membrane, constituted of these edgings: this membrane has been called the epidermis. In some genera, as in Lepas, this so-called epidermis is seldom preserved, excepting on the last zone of growth: is Scalpellum and Pollicipes it usually covers the whole valves. It appears to me that the laminæ of chitine, and of calcified chitine composing the valves, are both formed not by secretion, but by the metamorphosis of an outer layer of corium into these substances.

Within the capitulum is the sack, which, together with the upper internal part of the peduncle, encloses the animal's body. The sack is lined by a most delicate membrane of chitine, under which there is a double layer of corium; this double layer is united together by short, strong, transverse bundles of fibres, branched at both ends:* in some genera, the ovarian tubes extend between these two layers. We have seen, under the head of the Metamorphoses, that the delicate tunic lining the sack is simply a duplicature of the thick membrane and valves forming the capitulum, the whole being the posterior portion of the carapace of the larva slightly modified.

Peduncle.—Its length varies greatly in different species, and even in the same species, according to the situation occupied by the individual; its lower end is sometimes pointed, but generally only a little narrower than the upper end. In outline, the peduncle is usually flattened, but sometimes quite cylindrical. It is composed of very strong, generally thick, transparent membrane, rarely coloured reddish, and often penetrated by numerous tubuli. The underlying corium is sometimes

* I am much indebted to Mr. Inman of Liverpool for having kindly sent me excellent specimens illustrating this structure.

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coloured in longitudinal bands. At each period of growth a new and larger integument is formed under the old one, which gradually disintegrates and disappears; the extreme lower point is often deserted by the corium, and ceases to grow, whilst the whole upper part still continues increasing in diameter: in length the chief addition is made (as is clearly seen in those genera having calcified scales), round the upper margin, at the base of the capitulum. The surface of the membrane is either naked or superficially clothed with minute, pointed, articulated spines, or it is penetrated by calcified scales or styles, (in Ibla alone formed of chitine,) which pass through it to the corium, and are added to at their bases, like the valves, at each period of growth. In Lithotrya alone the scales of the peduncle are moulted together with the connecting membrane. These scales on the peduncle are generally placed symmetrically in whorls, with each scale corresponding with the junctions of two scales, both above and below. Except in Scalpellum ornatum and the fossil Loricula pulchella, they are very small compared with the valves of the capitulum. When the scales are symmetrical, new ones are first formed only round the summit of the peduncle, and only those in the few uppermost whorls continue to grow or to be added to at their bases; afterwards membrane is deposited under them. The shelly matter of the scales resembles that of the valves, and the manner of growth is the same; tubuli generally run to and through them from the corium. From the continued enlargement of the membrane of the peduncle, the scales come to stand, in the lower portion, some way apart. In Ibla, new horny styles are formed indifferently in all parts of the peduncle. In some species of Pollicipes, the calcareous styles are not symmetrical or symmetrically arranged; and besides those first formed round the top of the peduncle, there are other and larger ones formed near its base. Lastly, in Lithorya we have a row of calcareous discs or an irregular, basal cup, formed in the same

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manner as the valves of the capitulum: in this genus alone (as already stated,) the calcified scales are moulted, and here alone their edges are serrated.

The peduncle is lined within by three layers of muscles, longitudinal, transverse, and oblique, all destitute of the transverse striæ, characteristic of voluntary muscles; they run from the bottom of the peduncle to the base of the capitulum, as in Lepas, or half way up it, as in Conchoderma; in Alepas alone they surround the whole capitulum up to its summit. In Lithotrya there are two little, fan-like, transverse muscles (involuntary), extending from the basal points of the terga to a central line on the under side of the carina. The gentle swaying to and fro movements, and the great power of longitudinal contraction,—movements apparently common, as I infer from facts communicated to me by Mr. Peach, to all the Pedunculata,—are produced by these muscles. The interior of the peduncle is filled up with a great mass of branching ovarian tubes; but in Ibla and Lithotrya, the upper part of the peduncle is occupied by the animal's body.

Means of Attachment.—If the peduncle be very carefully removed (Tab. IX, fig. 7 and Tab. I, fig. 6b), from the surface of attachment, quite close to the end, but not at the actual apex, the larval prehensile antennæ can always be found: these have been sufficiently described for our present purpose under the head of the Metamorphoses; but I may add, that the diagnostic differences between them in the several genera are briefly given, for a special purpose, in a discussion on the sexes of Scalpellum at the end of that genus. We have seen in the larva, that the cement-ducts, with their opaque cellular contents, can be traced from within the discs of the antennæ to the anterior or lower ends of the two gutformed bodies, which it can be demonstrated are the incipient ovaria.

In mature Cirripedes these ducts can be followed, in a slightly sinuous course, along the muscles on each side


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within the peduncle, till they expand into two small organs, which I have called cement-glands. These glands are found with great difficulty, except in Conchoderma aurita, where they are placed on each side under the inner layer of corium, at the bottom of the sack, so as to be just above the top of the peduncle; they resemble in shape a retort, (Pl. IX, fig. 3.) In Pollicipes mitella and polymerus they lie half way down the peduncle, close together, and apparently enclosed within a common membrane; in these two species the broad end of the gland is bent towards the neck of the retort. In Scalpellum the position is the same, but the shape is more globular. In Ibla the structure is more simple, namely, a tube slightly enlarged, running downwards, bent a little upwards, and then resuming its former downward course, the lower portion forming the duct. The gland contains a strongly coherent, pulpy, opaque, cellular mass, like that in the cement-ducts; but in some instances, presently to be mentioned, this cellular mass becomes converted within either the ducts or gland, or within both, into transparent, yellow, tough cement. Generally in Conchoderma, Pollicipes, and Scalpellum, two ovarian tubes, but in one specimen of Conchoderma aurita, three tubes, and in Ibla one tube could be seen running into or forming the gland; of the nature of the tubes there could not be the least doubt, for at a little distance from the glands they gave out branches (Pl. IX, fig. 3), containing ova in every state of development. In some specimens as in that figured of Conchoderma aurita, the ovarian tube on one side of the gland is larger than on the other, and has rather the appearance of being deeply embedded in the gland than of forming it; but, in other specimens, the two ovarian tubes first formed a little pouch, into which their cellular contents could be clearly seen to enter; and then this pouch expanded into the gland; thus quite removing a doubt which I had sometimes felt, whether the ovarian tube was not simply attached to or embedded in the gland, without any further connection. By dissection

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the multiple external coats of the gland and ovarian tubes could be seen to be continuous. The cellular contents of the tubes passed into the more opaque cellular contents of the gland, by a layer of transparent, pulpy, pale, yellowish substance. There appeared in several instances to be a relation, between the state of fulness and condition of the contents of the gland, and of the immediately adjoining portions of the ovarian tubes. In one specimen of Pollicipes mitella it was clear that the altered, tough, yellow, transparent, non-cellular contents of the two glands and ducts, had actually invaded for some little distance, the two ovarian tubes which ran into them, thus showing the continuity of the whole. From these facts I conclude, without hesitation, that the gland itself is a part of an ovarian tube specially modified; and further, that the cellular matter, which in the ovarian tubes serves for the development of the ova, is, by the special action of the walls of the gland, changed into the opaquer cellular matter in the ducts, and this again subsequently into that tissue or substance, which cements the Cirripede to its surface of attachment.

As the individuals grow and increase in size, so do the glands and cement-ducts; but it seems often to happen, that when a specimen is immovably attached, the cementing apparatus ceases to act, and the cellular contents of the duct become converted into a thread of transparent tough cement; the investing membrane, also, of the ducts, in Conchoderma sometimes becomes hard and mamillated. I have already alluded to the case of a Pollicipes, in which both glands and ducts, and even a small portion of the two adjoining ovarian tubes, had become thus filled up. As in sessile Cirripedes, at every fresh period of growth a new cement gland is formed, it has occurred to me, that possibly in Pollicipes something similar may take place. In sessile Cirripedes, the old cement-glands are all preserved in a functionless condition, adhering to the membranous or calcareous basis, each new larger one attached to that last formed, and

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each giving out cement-ducts, which, bifurcating in the most complicated manner, pass outside the shell and thus attach it to some foreign body.

The cement, removed from the outside of a Cirripede, consists of a thin layer of very tough, bright-brown, transparent, laminated substance, exhibiting no structure under the highest powers, or at most a very fine dotted appearance, like a mezzotinto drawing. It is of the nature of chitine; but boiling caustic potash has rather more effect on it than on true chitine; and I think boiling nitric acid rather less effect. In one single instance, namely, in Coronula, the cement comes out of the four orifices of the two bifurcating ducts, in the shape of distinct cells, which, between the whale's skin and the basal membrane, arrange themselves so as to make a circular, continous slip of cement; then the cells blend together, and are converted into transparent, structureless cement. Cementing tissue or membrane would, perhaps, have been a more correct title than cement; but, in ordinary cases, its appearance is so little like that of an organized tissue, that I have for this reason, and for brevity-sake, preferred the simple term of Cement.

In the larva the cement always escapes through the prehensile antennæ; and it thus continues to do throughout life in most or all of the species of Lepas, Conchoderma, Dischelaspis and Ibla. In the first two of these genera, the cement escapes from the borders of the lower side of the disc of penultimate segment of the antennæ and can be there seen radiating out like spokes, which at their ends divide into finer and finer branches, till a uniform sheet of cement is formed, fastening the antennæ and the adjoining part of the peduncle down to the surface of attachment. In Dichelaspis Warwickii and Scalpellum Peronii, the cement, or part at least, comes out of the ultimate segment of the antennæ, in the shape of one tube, within another tube of considerable diameter and lenght. In Scalpellum vulgare, and probably in some of the other

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species, which live attached to corallines, the cement soon ceases to debouch from the antennæ but instead, bursts through a row of orifices on the rostral margin of the peduncle (Pl. IX, fig. 7), by which means this margin is symmetrically fastened down to the delicate, horny branches of the zoophyte. In Pollicipes, the two cement-ducts, either together or separately (Pl. IX, fig. 2, 2 a'), wind about the bottom of the peduncle in the most tortuous course, at each bend pouring out cement through a hole in the membrane of the peduncle. In Ibla the lower part of the peduncle is internally filled by cement, and thus rendered rigid. In Lepas fascicullaris a vesicular ball of cement surrounding the peduncle is thus formed (Pl. I, fig. 6), and serves as a float! All these curious, special adaptations are described under the respective genera. How the cement forces its way through the antennæ and often through apertures in the thick membrane of the peduncle, I do not understand. I do not believe, though some appearances favoured the notion, that the duct itself debouches and divides, at least this is not the case in Coronula, but only that the internal chord of cellular matter thus acts and spreads itself out; nor do I understand how, when the antennæ and immediately adjoining parts are once cemented down, any more cement can escape; yet this must take place, as may be inferred from the breadth of the cemented, terminal portion of the peduncle in Lepas and Conchoderma; and from the often active condition in old individuals of the cementing organs.

I have entered on this subject at some length, (and I wish I had space for more illustrations,) from its offering, perhaps, the most curious point in the natural history of the Cirripedia. It is the one chief character of the Subclass. I am well aware how extremely improbable it must appear, that part of an ovarian tube should be converted into a gland, in which cellular matter is modified, so that instead of aiding in the development of new beings, it forms itself into a tissue or substance, which leaves the

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body* in order to fasten it to a foreign support. But on no other view can be structure, clearly seen by me both in the mature Cirripede and in the larva, be explained, and I feel no hesitation in advancing it. I may here venture to quote the substance of a remark made by Professor Owen, when I communicated to him the foregoing facts, namely, that there was a new problem to solve,—new work to perform,—to attach permanently a crustacean to a foreign body; and that hence no one could, a priori, tell by what singular and novel means this would be effected.

Filamentary Appendages.—These have generally been considered to act as branchiæ; they occur at the bases of the first pair of cirri in Lepas, Alepas, Conchoderma, and in three species of Pollicipes: in Conchoderma there are similar appendages attached to the pedicels of the cirri (Pl. IX, fig. 4, g—k); and in the above three species of Pollicipes there is a double row of them on the prosoma: their numbers differ in different species (in some there being none) of the same genus, and even in different individuals of the same species; they are entirely absent in the majority of the genera. These facts would indicate that they are not of high functional importance; and they seem so generally occupied by testes (Pl. IV, fig. 5), that I suspect their function is quite a much to give room for the development of these glands, as to serve for respiratory purposes. With the exception of the four above-named genera, the mere surface of the body and of the sack must be sufficient for respiration: in Conchoderma aurita the two great expansions of surface, afforded by the folded, tubular, ear-like projections, aid, as I believe, towards this end.

* The protrusion of the egg-bearing pouches in Cyclops and its kindred genera, outside the body, offers a feeble analogy with what takes place in Cirripedes. Professor Allman ('Annals of Natural History,' vol. xx, p. 7,) who has attended to the subject, says that the external egg-bearing pouches are "a portion of the membrane of the true ovaries:" if the membrane of these pouches had been specially made adhesive, the analogy would have been closer.

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The shape of the body varies, owing to the greater or less development of the lower part of the prosoma, the greater or less distance of the first from the second pair of cirri, and of the mouth from the adductor scutorum muscle, (Pl. IX, fig. 4, and Pl. IV, 8a'). In all the genera, the body is much flattened. I may here mention a few particulars about the muscular system. One of the largest muscular masses is formed by the adductor scutorum, and by the muscles which surround in a double layer (the fasciæ being oblique to each other) the whole of the upper part of the prosoma. From under the adductor, a pair of delicate muscles runs to the basal edge of the labrum, so as to retract the whole mouth, and two other pair to the integument between the mouth and the adductor, so as to fold it: again, there are other delicate muscles in some (for instance in Lepas Hillii) if not in all the Lepadidæ, crossing each other in the most singular loops, and serving apparently to fold the membrane between the occludent edges of the scuta. Within the prosoma there is a strong adductor muscle, running straight from side to side, for the purpose, as it appears, of flattening the body. The thorax, on the dorsal and ventral surfaces, is well furnished with straight and oblique muscles (without striæ), which straighten and curl up this part of the body. The muscles running into the pedicels of the cirri, cross each other on the ventral surface of the thorax; the muscles within the rami are attached to the upper segments of the pedicels. Finally, I may remark that the whole of the body and the cirri are capable of many diversified movements.

Mouth.—This is prominent, and almost probosciformed (Pl. IX, fig. 4 b), and in the abnormal Anelasma (Pl. IV, fig. 2 d), quite probosciformed,—such, also, was its character in the larval condition. In outline, it is either sub-triangular, or oval with the longer axis transverse; the whole is capable, as well as the separate organs, of considerable movement, as I have seen in living sessile Cirripedes. It is composed (Tab. V, fig. 2) of a labrum,

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swollen or bullate, often to such an extent as to equal in its longitudinal axis the rest of the mouth; of palpi soldered to the labrum; of mandibles, maxillæ and outer maxillæ, the latter serving as a lower lip. These organs have only their upper segments free, but there are traces, clearly seen in the mandibles (Pl. X, fig. 1, a, b), of their being formed of three segments. The two lower segments are laterally united, and open into each other, the prominence of the mouth being thus caused: this condition appears to me curious, and is, to a certain limited extent, intermediate between those articulated animals which have their trophi soldered into a proboscis, and those furnished with entirely free masticatory or prehensile organs. The palpi adhere to the corners of the labrum; and I call them palpi only from seeing that they spring laterally from above the upper articulation of the mandibles. The prominence of the mouth, measured from the basal fold by which the whole is separated from the body, is much greater on the half formed by the labrum and mandibles, than on the other half facing the cirri. The trophi surround a cavity—the supra-oesophageal cavity—in the middle of which, between the mandibles is seated the orifice of the oesophagus. The oesophagus is surrounded by long, fine, muscular fasciæ, radiating in all directions, opposing the constrictor muscles, and is capable of violent swallowing movements,—constriction after constriction being seen to run down its whole course: there are also some fine muscles attached to the membrane forming the supra-oesophageal cavity. The trophi serve merely for the prehension of prey, and not for mastication.

The Labrum, as stated, is always bullate or swollen; and sometimes the upper exterior part forms, as in Ibla (Pl. IV, fig. 8 a, c), and Dichelaspis, an overhanging blunt point. The object, I suspect, of this bullate form is to give, in the upper part, attachment to longer muscles running to the lateral surfaces of the mandibles, and lower down to the oesophagus. The crest close over the

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supra-oesophageal cavity, is generally furnished with small, often bead-like teeth. The Palpi are small, their apices never actually touching each other; they are more or less blunt, not differing much in shape in the different genera (Pl. X, figs. 6 to 8), and clothed with spines. They are not capable of movement; their function seems to be to prevent prey, brought by the cirri, escaping over the labrum; I infer this from finding in Anelasma and in the male of Ibla, which have the cirri functionless, that the palpi are rudimentary.

The Mandibles (Pl. X, figs. 1—5) have from two to ten strong teeth in a single row; where the number exceeds five, several of the teeth are small; the inferior angle is generally pectinated with fine spines; in Lithotrya (fig. 2), the interspaces between the teeth are also pectinated. In the same individual there is not unfrequently one tooth, more or less, on opposite sides of the mouth. Internally, the mandibles are furnished on their outer and inner sides with several ligamentous apodemes, (in Lithotrya roughened with points (Pl. X, fig. 2), for the attachment of the muscles; of these (fig. 1), there is a chief depressor and elevator, attached at their lower ends to near the basal fold of the mouth, and a lateral muscle, attached to the broad basal end of the palpi, and serving, apparently, to oppose the edge of mandible to mandible. The Maxillæ in the different genera (Pl. X, figs. 9 to 15) differ considerably in outline; they are generally about half the size of the mandibles; at the upper corner, there are always two or three spines larger than the others, and often separated from them by a notch; the rest of the spinose edge is straight, or irregular, or step-formed, or with the lowest part projecting, or with one or two narrow prominences bearing fine spines. All these spines, quite differently from the teeth of the mandibles, are articulated on the edge of the organ, and stand in a double row. At a point corresponding with the upper articulation of the mandibles, a long, thin, narrow, rigid apodeme, projects inwards (fig. 10), and running

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down nearly parallel to the thin, outer, flexible membrane of the mouth, is attached to the corium, and thus serves as a support to the whole organ. This apodeme is embedded in muscles (Pl. X, fig. 10); there are other large muscles attached to the inner side of the organ, and again others running laterally towards the mandibles. The apodeme, of course, is moulted with the integuments of the mouth. The Outer Maxillæ (Pl. X, figs. 16, 17) serve as a lower lip; they are thicker than the other trophi; they have their inner surfaces clothed with spines, sometimes divided into an upper and lower group, and occasionally separated by a deep notch: there are often long bristles outside. They are furnished with at least two muscles; in sessile Cirripedes I have seen that they are capable of a rapid to and fro movement, and I have no doubt that their function is to brush any small creature, caught by the cirri, towards the maxillæ, which are well adapted to aid in securing the prey, and to hand it over to the mandibles, by them to be forced down the oesophagus. On the exterior face of the outer maxillæ, above a trace of an upper articulation, either two small orifices or two large tubular projections can always be discovered; and these, as will presently be mentioned, I believe to be olfactory organs.

Cirri.—The five posterior pair are seated close to each other and equidistant; the first pair is generally seated at a little distance, and sometimes at a considerable distance from the second pair. The first pair is the shortest; the others, proceeding backwards, increase gradually in length. The rami of each pair are either equal in length or slightly unequal: those of the first pair are oftenest unequal. The number of segments in the posterior cirri is sometimes very great; in one species of Alepas, there were above sixty segments in one ramus, the other ramus being in this unique case (Pl. X, fig. 28) small and rudimentary. The pedicels consist of two segments, a lower, longer, and upper short one (fig. 18, c, d.) In the usual arrangement of the spines on the segments of the three posterior pair of cirri, there are (figs. 26, 27) from three to six pair of

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long spines on the anterior face, with generally some minute spines (occasionally forming a tuft) intermediate between them: on the dorsal surface, in the uppermost part of each segment, there is a tuft of short spines generally mingled with some longer, finer ones: on the inner side of each segment, on the upper rim, there are generally a few extremely minute and short spines. From the increase of these latter and of the intermediate spines, the antero-lateral faces of the segments of the first cirrus, and of the lower segments of the anterior ramus of the second cirrus (Pl. X, fig. 25), are almost always thickly paved with brush-like masses of spines. The lower segments of the anterior ramus of the third cirrus is generally, though not always, thus paved: these paved segments are much broader than the others. The posterior rami of the second and third cirri are often in some slight degree paved, though in other cases they resemble the three posterior pair of cirri. The two segments of the pedicels have bristles on their anterior faces, essentially arranged on the same plan as on the segments of the rami: the bristles are generally not so symmetrically arranged on the pedicels of the second and third cirri, as on the three posterior pair. There are some exceptions to the foregoing general rules: in the posterior cirri of Alepas cornuta, there is only one pair of long spines to each segment (fig. 28); in Dichelaspis Lowei, there are eight pair; in Lepas fascicularis, in old specimens, the segments are paved with a triangular brush of spines; the upper segments in Pæcilasma eburnea support small oblong brushes; and, lastly, in Pæcilasma fissa (fig. 29), and crassa, the spines form a single circle round each segment, interrupted on the two sides. These spines are often doubly serrated or plumose: many of them on the protuberant segments of the first three pair of cirri, are sometimes coarsely and doubly pectinated.

Caudal Appendages.—These are present (Pl. X, figs. 18 to 24) seated on each side of the anus, in all the genera, except in Conchoderma, Anelasma, and Scalpellum

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villosum; they consist of a very small single segment, destitute of spines in Lepas, and spinose in Pæcilasma, Dichelaspis, Oxynaspis, Scalpellum, and some species of Pollicipes; they consist of several segments in Alepas, Ibla, Lithotrya, and in some species of Pollicipes. In the latter genus, some species have their caudal appendages multiarticulate, though so obscurely articulated, that the passage (fig. 22) from several to one segment is seen to be easily effected. When the appendage consists of many articulations, it is generally about as long as the pedicel of the sixth cirrus; but in Ibla quadrivalvis, it is four times as long. The segments are narrow, slightly flattened, much tapering; each (fig. 24) is surmounted by a ring of short spines, which are generally longest on the apex of the terminal segment. I could never trace muscles into these appendages.

Alimentary Canal.—The oesophagus is of considerable length: it is formed of strong, transparent, much folded membrane, continuous with the outer integuments, and moulted with them: it is surrounded by corium, and as already stated, by numerous muscles: at its lower end it expands into a bell, with the edges reflexed, and sometimes sinuous: this bell lies within the stomach, and keeps the upper broad end expanded. According to the less or greater distance of the mouth from the adductor muscle, the oesophagus runs in a more or less parallel course to the abdominal surface between the first and succeeding pairs of cirri, and enters the stomach more or less obliquely. In Ibla alone, it passes exteriorly to, and over the adductor scutorum muscle. The stomach lies in a much curved, almost doubled course; it is often a little constricted where most bent; it is broadest at the upper end, and here, in Lepas and Conchoderma, there are some deep branching cæca; in the latter of these two genera, the whole surface is, in addition, pitted in transverse lines. The stomach is coated by small, opaque, pulpy, slightly arborescent glands, believed to be hepatic; these are arranged in longitudinal lines, in all

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the genera, except in Alepas, in which they are transverse and reticulated: the whole stomach is thus coated. There is, also, a coating of excessively delicate, longitudinal and transverse muscles without striæ. The rectum varies in length, extending inwards from the anus to between the bases of the second and fifth pair of cirri: it is narrow, and formed of much folded transparent membrane, resembling the oesophagus, continuous with the outer integuments, with which it is periodically moulted. The anus is a small longitudinal slit, in the triangular piece of membrane representing the abdomen, let in between the last thoracic tergal arches, as already mentioned under the head of the Metamorphoses; it lies almost between the caudal appendages, and opens on the dorsal surface. Within the stomach, there can generally be plainly seen, in accordance with the period of digestion when the specimen was taken, a thin, yet strong, perfectly transparent epithelial membrane, not exhibiting under the highest power of the microscope any structure: it enters the branching cæca, and extends from the edge of the bell of the oesophagus to the commencement of the closed rectum, and consequently terminates in a point: it consists of chitine, like the outer integuments of the animal, and by placing the whole body in caustic potash, I have dissolved the outer coats of the stomach, and seen the bag open at its upper end, perfectly preserved, floating in the middle of the body, and full of the debris of the food. In most of the specimens which I have examined, preserved in spirits of wine, this epithelial lining was some little way distant and separate from the coats of the stomach; and hence was thought by M. Martin St. Ange to be a distinct organ, like the closed tube in certain Annelids. Occasionally, I have seen one imperfect epithelial bag or tube within another and later-formed one. Digestion seems to go on at the same rate throughout the whole length of the stomach; if there be any difference, the least digested portions lie in the lower and narrower part. The prey, consisting generally of crustacea, infu-

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soria, minute spiral univalves, and often of the larvæ of Cirripedes, is not triturated: when the nutritious juices have been absorbed, the rejectamenta are cast out through the anus, all kept together in the epithelial bag, which is excluded like a model of the whole stomach, with the exception of that part coated by the bell of the oesophagus. I have sometimes thought that the bag was formed so strong, for the sake of thus carrying out the excrement entire, so as not to befoul the sack. I believe Lepas can throw up food by its oesophagus; at least, I found in one case, many half-digested small Crustaceans in the sack, and others of the same kind in the stomach.

Circulatory System.—I can add hardly anything to what little has been given by M. Martin St. Ange: like others, I have failed, as yet, in discovering a heart. The whole body is permeated by channels, which have not any proper coat: there is one main channel along the ventral surface of the thorax, dividing and surrounding the mouth, and giving out branches which enter the inner of the two channels in each cirrus: as Burmeister has shown, there are also two channels in the penis. There are two dorso-lateral channels in the prosoma, which are in direct connection with the great main channel, running down the rostral (i.e., ventral) side of the peduncle. This latter main channel branches out in the lower part, and transmits the fluid through the ovarian tubes, whence, I believe, it flows upwards and round the sack, re-entering the body near the sides of the adductor scutorum muscle. The main rostral channel (or artery?) in the uppermost part of the peduncle, has a depending curtain, which, I think, must act as a valve, so as to prevent the circulating fluid regurgitating into the animal's body during the contractions of the peduncle.

Nervous System and Organs of Sense.—In most of the genera, there are six main ganglia, namely, the supra-oesophageal, and five thoracic ganglia; but in Pollicipes mitella there are only four thoracic ganglia. Of these, the first thoracic or infra-oesophageal ganglion is con-

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siderably the largest and most massive; it is squarish, or oval, or heart-shaped; it presents no trace of being formed by the union of two lateral ganglia. Two great nerves spring from its under side (A), represented in the woodcut on page 49, by dotted lines), and run straight down amongst the viscera in the prosoma: these nerves are about as large as those forming the collar and those running to the second ganglion; hence, six great nerves meet here, two in front, two behind, and two on the under side. At the anterior end, over the junction with the collar chord, three equal-sized nerves rise on each side, with a fourth, smaller one, outside; these go to the trophi and to the two olfactory sacks. At the posterior end, on each side, a pair of nerves branch out rectangularly, one of which (a,) goes to the first cirrus, and there divides into two branches; of these, the upper runs up the cirrus, and the lower one downwards. The other nerve (b), proceeding on each side from this first thoracic ganglion, runs to the muscles beneath the basal articulation of the first cirrus. The collar surrounding the oesophagus is generally very long, sometimes equalling the whole thoracic chord; at a middle point, a small branch is sent off, and at the anterior end (e, e), close to the supra-oesophageal ganglia, double or treble fine branches run to the true ovaria, lying close to the upper end of the stomach. The four (or only three) other thoracic ganglia, when viewed as transparent bodies, are seen to be solid; but in some of the genera, as in Conchoderma, the outline plainly shows, that each consists of a lateral pair fused together. The second thoracic ganglion (B) is rather small; it is either close to the first, as in Pollicipes mitella and Lepas fascicularis, or far distant, as in Ibla. The third (C) and fourth are of about the same size with the second: these three ganglia send large branches to the second, third, and fourth pair of cirri: other minute branches spring from their under sides, and from the intermediate double chords. The fifth ganglion is larger and longer than the three preceding ones, and gives off

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nerves to the fifth and sixth pair of cirri; it is clearly formed by the union of the fifth, with what ought to have formed a sixth ganglion. The two nerves going to the sixth cirrus give off on their inner sides, each a great branch to the penis. In Pollicipes mitella, in which there are only four instead of five thoracic ganglia, it is evident from the outline and position of the nerves going to the fourth pair of cirri, that the fourth ganglion is fused into the fifth, itself, as we have just seen normally composed of two consecutive ganglia. In this Pollicipes there is other evidence of concentration in the nervous system, for none of the ganglia show signs of being formed of lateral pairs; the second is close to the first; and the abdominal double chord is in part separated by a mere cleft; lastly, as we shall immediately see, the same remark is applicable to the supra-oesophageal ganglia.

The latter (D) alone remain to be described; they present far more diversity in shape than do the thoracic ganglia; they are almost always seen in outline to be laterally distinct, and usually resemble two pears with their tapering ends cut off and united; in a transverse line they are as long as the infra-oesophageal ganglion, but are much less massive. In Lepas fascicularis (D), they are pear-shaped; in Pollicipes mitella they are globular, and separated by a third globular ganglion, which I believe is the ophthalmic ganglion, presently to be described; in Pollicipes spinosus, however, the ophthalmic ganglion is, as usual, placed in advance of the supra-oesophageal ganglion, which latter, in this one species, shows no sign of being formed of a lateral pair fused together. In Alepas cornuta the supra-oesophageal ganglion consists of two quite distinct ganglia, elongated in the longitudinal axis of the body, and separated from each other by the whole width of the mouth; the chord which unites them is of the same thickness as the rest of the collar. In all the genera, from the front of each of two supra-oesophageal ganglia, a pair of nerves,

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(f, f,) united and together as large as the collar nerve, rises, and can be traced running unbranched, in a nearly straight line, for a length equalling the whole rest of the nervous chord, so as to supply the peduncle and the inside of the capitulum or sack. At the inner ends of these two same ganglia, from a central point where they are united, a little central branch runs in front to the adductor scutorum and other adjoining muscles; and still smaller fibrils run behind to the oesophageal muscles.

Opthalmic Ganglia and Eyes.—Owing to Professor Leidy's* discovery of eyes in a Balanus, I was led to look for them in the Lepadidæ. Extending from the front of the two supra-oesophageal ganglia, two chords may be seen in Lepas fascicularis (of which a rude diagram is here given), to run into two small, perfectly distinct oval

Diagram of the anterior portion of the nervous system in Lepas fascicularis. A. First thoracic or infra-oesophageal ganglion. B. Second thoracic. C. Third thoracic ganglion. D. Supra-oesophageal ganglion. E. The two ophthalmic ganglia. F. Double eye. a. Nerve going to first cirrus; b, to the muscles below the first cirrus; c, to the second cirrus; d, to the third; e, nerves running to the ovaria; f, double nerves supplying the sack and peduncle.

* Proceedings of the Academy of Natural Sciences, Philadelphia. No. i, vol. iv, Jan. 1848.


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ganglia (E), which are not united by any transverse commissure.

From the opposite ends of these two ganglia smaller nerves run, and, bending inwards at right angles, enter, beyond the middle, an elongated (F), almost black, eye, composed of two eyes united together. Although in outline the eye appears single, two lenses can be distinctly seen at the end, directed upwards and towards the ganglia; two pigment-capsules can also be distinguished; these are deep and cup-formed, and of a dark reddish-purple. The following measurements will show the proportions of the parts in a specimen of the Lepas fascicularis having a capitulum 4/10ths of an inch in length.

Double eye length . . 26/6000
Double eye width . . 13/6000
Diameter of single lens . . 6/6000
Opthamalic ganglion length . . 16/6000
Opthamalic ganglion width . . 11/6000

Supra-œsophagal ganglion, transverse or longest axis of both together . . 126/6000
Supra-œsophagal ganglion, longitudinal axis of . . 45/6000
Infra-œsophagal ganglion, transverse axis of . . 120/6000
Infra-œsophagal ganglion, longitudinal axis of . . 114/6000

In Conchoderma aurita the ophthalmic ganglia are much smaller, and nearer to the supra-oesophageal ganglion, than in L. fascicularis. In Alepas cornuta the ophthalmic chords run towards each other from the two distant and separate supra-oesophageal ganglia; and the ophthalmic ganglia, (instead of being quite separate, as in L. fascicularis,) are united by their front ends, and the two eyes instead of standing some way in front, with nerves running to them, are embedded on the double ophthalmic ganglion; the pigment-capsules here, also, have the shape of mere saucers, and are joined back to back, with the two lenses projecting far out of them. In neither sex of Ibla could I perceive that the eye was double. In Pollicipes spinosus the ophthalmic ganglion stands in front of the single supra-oesophageal ganglion, and shows no signs of being formed of a lateral pair; the eyes themselves, however, differently from, in all the foregoing cases, are, though approximate, quite distinct. In Pollicipes

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mitella I did not see the eyes; but the ophthalmic ganglion consists, as I believe, of a single globular one, placed exactly between the two globular, supra-oesophageal ganglia, all three being of nearly equal size. Professor Leidy does not mention the ophthalmic ganglia; hence I infer that in Balanus, which is a more highly organised Cirripede, they are fused into the supra-oesophageal ganglion.

In all the genera, the double eye is seated deep within the body; it is attached by fibrous tissue to the radiating muscles of the lowest part of the oesophagus, and lies actually on the upper part of the stomach; consequently, a ray of light, to reach the eye, has to pass through the exterior membrane and underlying corium connecting the two scuta, and to penetrate deeply into the body. In living sessile Cirripedes, vision seems confined to the perception of the shadow of an object passing between them and the light; they instantly perceived a hand passed quickly at the distance of several feet between a candle and the basin in which they were placed.

As the infra-oesophageal ganglion sends nerves to the trophi and to the first pair of cirri, it must correspond to the segments, from the fourth to the ninth inclusive, of the archetype crustacean. The state of the supra-oesophageal and opthalmic ganglia appears to me very interesting: I do not believe that in any mature ordinary crustacean, the first or ophthalmic ganglion can be shown to be distinct from the two succeeding ganglia, or to be itself composed of a pair laterally distinct. The ganglia, corresponding with the second and third segments of the body, which should normally support two pair of antennæ, are in the Lepadidæ united together; but laterally they are generally distinct in outline, and are actually separate in Alepas: the supra-oesophageal ganglion shows also its double nature, by giving rise to a pair of large double nerves evidently corresponding with the two pair of antennular nerves in ordinary crustaceans. The embryonic condition of the whole supra-oesophageal portion of the nervous system in the Lepadidæ, corresponds with the

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rudimentary state of the only organ of sense supplied by it, namely, the eye, which in size and general appearance has retrograded to the state in which it was in, during the first stage of development of the larva;—I have used the term embryonic, because, in the embryos of ordinary crustacea, all the ganglia are at first longitudinally distinct, and laterally quite separate. The conclusion at which we before arrived from studying the metamorphoses, namely, that the whole peduncle and capitulum consisted of the first three segments of the head, is beautifully supported by the structure of the nervous system, in which these parts are seen to be supplied with nerves exclusively from the supra-oesophageal ganglion: now in ordinary curstacea the supra-oesophageal ganglion sends nerves to the eyes and the two pair of antennæ corresponding, as is known by embryological dissections, to the first three segments of the body. Moreover, it is asserted that the carapace which covers the thorax in crustacea, is not formed by the development of the first segment; and this, likewise, may be inferred to be the case with the peduncle and capitulum in the Lepadidæ, as the nerves of the ophthalmic ganglia go exclusively to the eyes. Finally, I may remark that in Pollicipes, looking to the whole nervous system, the state of concentration nearly equals that in certain macrourous decapod crustaceans, for instance the Astacus marinus, of which a figure is given by Milne Edwards.

Olfactory Organs.—In the outer maxillæ, at their bases where united together, but above the basal fold separating the mouth from the body, there are, in all the genera, a pair of orifices (Pl. X, fig. 16); these are sometimes seated on a slight prominence, as in Lithotrya, or on the summit of flattened tubes (Pl. X, fig. 17), projecting upwards and towards each other, as in Ibla, Scalpellum, and Pollicipes. In Ibla these tubular projections rise from almost between the outer and inner maxillæ. It is impossible to behold these organs, and doubt that they are of high functional importance to the animal. The orifice leads

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into a deep sack lined by pulpy corium, and closed at the bottom. The outer integument is inflected inwards, (hence periodically moulted,) and becoming of excessive tenuity, runs to near the bottom of the sack, where it ends in an open tube: so excessively thin is this inflected membrane, that, until examining Anelasma, I was not quite certain that I was right in believing that the outer integument did not extend over the whole bottom. I several times saw a nerve of considerable size entering and blending into a pulpy layer at the bottom of the sack of corium; but I failed in tracing to which of the three pair of nerves, springing from the front end of the infra-oesophageal ganglion, it joined. I can hardly avoid concluding, that this closed sack, with its naked bottom, is an organ of sense; and, considering that the outer maxillæ serve to carry the prey entangled by the cirri towards the maxillæ and mandibles, the position seems so admirably adapted for an olfactory organ, whereby the animal could at once perceive the nature of any floating object thus caught, that I have ventured provisionally to designate the two orifices and sacks as olfactory.

Acoustic (?) Organs.—A little way beneath the basal articulation of the first cirrus (Pl. IX, fig. 4d, and Pl. IV, fig. 2e), on each side, there may be seen a slight swelling, and on the under side of this, a transverse slit-like orifice, 1/20th of an inch in length in Conchoderma, but often only half that size. In Ibla this orifice is seated lower down (Pl. IV, fig. 8a', e), between the bases of the first and second cirri, which are here far apart: in Alepas cornuta it is placed rather nearer to the adductor scutorum muscle, namely, beneath the mandibles. The orifice leads into a rather deep and wide meatus; the external integument is turned in for a short distance, widening a little, and then ends abruptly. The meatus, enlarging upwards, is lined by thick pulpy corium, and is closed at the upper end; from its summit is suspended a flattened sack of singular and different shapes in the different genera. This, the so-called acoustic sack of Conchoderma virgata,

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is figured Pl. IX, fig. 6. The deep and wide notch faces towards the posterior end of the animal; the inferior lobe, thus almost cut off, is flattened in a different plane from the upper part; the lobe is lodged in a little pouch of corresponding form, leading from the open meatus in which the upper part is included. In Conchoderma aurita, the top of the acoustic sack is narrower and more constricted, the whole more rounded, and the lobe more turned down. In Lepas fascicularis the notch is not so deep or wide, and the lobe larger. In Ibla Cumingii the sack is of the shape of a vase, with one corner folded over. In Scalpellum vulgare it is small, oval, with the lower end much pushed in, and furnished with a little crest. Lastly, in Pollicipes mitella it is simply oval. In all cases the sack is empty, or contains only a little pulpy matter: it consists of brownish, thick, and remarkably elastic tissue, formed, apparently, of transverse little pillars, becoming fibrous on the outside, and with their inner ends appearing like hyaline points. The mouth of the acoustic sack (removed in the drawing) is closed by a tender diaphragm, through which I saw what I believe was a moderately-sized nerve enter; I have not yet succeeded in tracing this nerve. The first pair of cirri seem, to a certain extent, to serve as antennæ, and therefore the position of an acoustic organ at their bases, is analogous to what takes place in crustacea; but there are not here any otolites, or the siliceous particles and hairs, as described by Dr. Farre, in that class. Nevertheless, the sack is so highly elastic, and its suspension in a meatus freely open to the water, seems so well adapted for an acoustic organ, that I have provisionally thus called it. In the larva, as I have shown, a pouch, certainly serving for some sense, I believe for hearing, is seated in quite a different position at the anterior end of the carapace. I may mention that I found sessile Cirripedes very sensitive of vibrations in objects adjoining them, though not, apparently, of noises in the air or water. In a group of specimens, I could not touch one even most delicately

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with a needle, without all the adjoing ones instantly withdrawing their cirri; it made no difference if the one touched had its operculum closed and motionless.

Reproductive System,—Male Organs.—All the Cirripedia which I have hitherto examined, with the exception of certain species of Ibla and Scalpellum, are hermaphrodite or bisexual.* I shall so fully describe the sexual relations of the several species of these two genera, under their respective headings, and at the end of the genus of Scalpellum, that I will not here give even an abstract of the grounds on which my firm belief is based, that the masculine power of certain hermaphrodite species of Ibla and Scalpellum, is rendered more efficient by certain parasitic males, which, from their not pairing, as in all hitherto known cases, with females, but with hermaphrodites, I have designated Complemental Males.

The male organs have been well described by M. Martin St. Ange, whose observations have since been confirmed by R. Wagner.† The testes are small, often leaden-coloured, either pear or finger-shaped, or branched like club-moss,—these several forms sometimes occurring in the same individual; they coat the stomach, enter the pedicels, and even the basal segments of the rami of the cirri, and in some genera occupy certain

* I am compelled to differ greatly from the account given by Prof Steenstrup of the reproductive system in the Cirripedia, in his 'Untersuchungen über das Vorkommen des Hermaphroditismus,' ch. v, 1846;—a translation of which I have seen, owing to the great kindness of Mr. Busk. Mr. Goodsir has described ('Edin. New Phil. Journal, 'July 1843,) what he considers the male of Balanus; but I have seen this same parasitic creature charged with ova, including larvæ! From the resemblance of the larvæ to the little crustacean described by Mr. Goodsir, in the same paper, as a distinct parasite, I believe the latter to be the male of his so-called male Balanus, and that all belong to the same species, allied to Bopyrus. This genus, as is well known, is parasitic on other crustacea; and it is a rather interesting fact thus to find, that this new parasite which is allied to Bopyrus, in structure, is likewise allied to it in habits, living attached to Cirripedia, a sub-class of the crustacea.

† In 'Müller's Archiv,' 1834, p. 467. I have already several times referred to M. Martin St. Ange's excellent Memoir, read before the Academy of Sciences, and subsequently, in 1835, published separately.

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swellings on the thorax and prosoma, and in others the filamentary appendages: the testes seen in the apex in one of these appendages in Conchoderma, is represented in Pl. IX, fig. 5. The two vesiculæ seminales are very large; they lie along the abdominal surface of the thorax, and generally (but not in some species of Scalpellum) enter the prosoma, where their broad ends are often reflexed; here the branched vessels leading from the testes enter. The membrane of the vesiculæ seminales is formed of circular fibres; and is, I presume, contractile, for I have seen the spermatozoa expelled with force from the cut end of a living specimen. The two canals leading from the vesiculæ generally unite in a single duct at the base of the penis; but in Conchoderma aurita, half-way up it. The probosciformed penis, except in certain species of Scalpellum, is very long; it is capable of the most varied movements; it is generally hairy, especially at the end; it is supported on a straight unarticulated basis, which in Ibla quadrivalvis alone (Pl. IV, fig. 9a), is of considerable length; in this species, the upper part is seen to be as plainly articulated as one of the cirri; in Alepas, the articulations are somewhat less plain, and in the other genera, the organ can be said only to be finely ringed, but these rings no doubt are in fact obscure articulations. In the females of Ibla Cumingii and Scalpellum ornatum, there is, of course, no penis.

Female Organs.—M. Martin St. Ange has described how the peduncle* is gorged with an inextricable mass of branching ovarian tubes, filled with granular matter and immature ova. In Conchoderma and Alepas, the ovarian tubes run up in a single plane (Pl. IX, fig. 3,) between the two folds of corium round the sack. Here the development of the ova can be well followed: a minute point first branches out from one of the tubes; its

* I may here mention, that in all sessile Cirripedes, the ovarian branching tubes lie between the calcareous or membranous basis and the inner basal lining of the sack, and to a certain height upwards round the sack: the true ovaria and the two ducts occupy the same position as in the Lepadidæ.

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head then enlarges, like the bud of a tulip on a footstalk; becomes globular; shows traces of dividing, and at last splits into three, four, or five egg-shaped balls, which finally separate as perfect ova. Within the peduncle, the ovarian tubes branch out in all directions, and within the footstalks of the branches (differently from what takes place round the sack), ova are developed, as well as at their ends. Close together, along the rostral (i.e., ventral) edge of the peduncle, two nearly straight, main ovarian tubes or ducts may be detected, which do not give out any branches till about half way down the peduncle, where they subdivide into branches, which inosculate together, and give rise to the mass filling the peduncle, and sometimes, as we have just seen, sending up branches round the sack. These two main unbranched ovarian ducts, followed up the peduncle, are seen to enter the body of the Cirripede (close along side the great double peduncular nerves), and then separating, they sweep in a large curve along each flank of the prosoma, under the superficial muscles, towards the bases of the first pair of cirri; and then rising up, they run into two glandular masses. These latter rest on the upper edge of the stomach, and touch the cæca were such exist; they were thought by Cuvier to be salivary glands. They are of an orange colour, and form two, parallel, gut-formed masses, having, in Conchoderma, a great flexure, and generally dividing at the end near the mouth into a few blunt branches. I was not able to ascertain whether the two main ducts, coming from the peduncle, expanded to envelope them, or what the precise connection was. The state of these two masses varied much; sometimes they were hollow, with only their walls spotted with a few cellular little masses; at other times they contained or rather were formed of, more or less globular or finger-shaped aggregations of pulpy matter; and lastly, the whole consisted of separate pointed little balls, each with a large inner cell, and this again with two or three included granules. These so closely

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resembled, in general appearance and size, the ovigerms with their germinal vesicles and spots, which I have often seen at the first commencement of the formation of the ova in the ovarian tubes in the peduncle, that I cannot doubt that such is their nature. Hence I conclude, that these two gut-formed masses are the true ovaria. I may add, that several times I have seen in the two long, unbranched ducts, connecting the true ovaria and the ovarian tubes in the peduncle, pellets of orange-coloured cellular matter (i.e., ovigerms) forming at short intervals little enlargements in the ducts, and apparently travelling into the peduncle.

The structure here described is quite comformable with that which we have seen in the larva; in the latter, two gut-formed masses of equal thickness extended from the cæca of the stomach to within the future peduncle, where the cement-ducts entered them, and where, after a short period, they were seen to expand into a mass of ovarian tubes. In the mature Cirripede, the cement-ducts can still be found united to the ovarian tubes in the middle of peduncle; and the cause of the wide separation of the true ovaria and ovarian tubes, can be simply accounted for by the internal, almost complete intersection of the animal, which takes place during the last metamorphosis.

The ova, when excluded, remain in the sack of the animal until the larvæ are hatched; they are very numerous, and generally form two concave, nearly circular, leaves, which I have called after Steenstrup and other authors, the ovigerous lamellæ (Pl. IV, fig. 2b). These lamellæ lie low down on each side of the sack: in Conchoderma virgata, however, there is often only a single lamella, forming a deeply concave cup: in C. aurita there are generally on each side four lamellæ, one under the other. The ova lie in a layer from two to four deep; and all are held together by a most delicate transparent membrane, which separately enfolds each ovum: this membrane is often thicker and stronger round the margins of the lamellæ, where they are united, in a peculiar manner, presently to be

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described, to a fold of skin, on each side of the sack: these two folds, I have called the ovigerous fræna (Pl. IV, fig. 2f).

M. Martin St. Ange describes an orifice under the carina, by which he supposes the ova to enter the sack; this, after repeated and most careful examinations, I venture to affirm does not exist; on the contrary, I have every reason to believe that the ova enter the sack in the following curious manner. Immediately before one of the periods of exuviation, the ova burst forth from the the ovarian tubes in the peduncle and round the sack, and, carried along the open circulatory channels, are collected (by means unknown to me) beneath the chitinetunic of the sack, in the corium, which is at this period remarkably spongy and full of cavities. The corium then forms or rather (as I believe) resolves itself into the very delicate membrane separately enveloping each ovum, and uniting them together into two lamellæ; the corium having thus far retreated, then forms under the lamellæ the chitine-tunic of the sack, which will of course be of larger size than the last-formed one, now immediately to be moulted with the other integuments of the body. As soon as this exuviation is effected, the tender ova, united into two lamellæ, and adhering, as yet, to the bottom of the sack, are exposed: as the membranes harden, the lamellæ become detached from the bottom of the sack, and are attached to the ovigerous fræna. To demonstrate this view, an individual should have been found, with both the old and new chitine tunic of the sack, and with the lamellæ lying between them; this, I believe, I have seen, but it was before I understood the full importance of the fact: a great number of specimens would have to be examined in order to succeed again, for the changes connected with exuviation supervene very quickly. I have, however, several times found the ova so loose under the sack, as to be detached with a touch from the ovarian tubes; and I have twice carefully examined specimens, which had just moulted, as shown by even the mandibles being flexible, in which the lamellæ had not become united to

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the fræna, but still adhered to the newly-formed chitine tunic of the sack; in these, the ova were so tender, that they broke into pieces rather than be separated from the membrane of the lamella, itself hardly perfectly developed, for pulpy cellular matter adhered outside some of the ova. These and other facts are quite inexplicable on any other view than that advanced.

As the lamellæ are formed without organic union with the parent, they would be liable to be washed out of the widely open sack of the Lepadidæ, if they had not been specially attached to the fræna. These fræna consist of a pair of more or less semicircular folds of skin, depending inside the sack, on each side of the point of attachment of the body. The fræna are often of considerable size, but in Ibla, they are very minute; they are formed of chitine tunic with underlying corium, like the rest of the sack; on their crests, there is a row, or a set of circular groups, or a broad surface, covered, either with minute, pointed, bead-like bodies mounted on long hair-like footstalks, or with staff-formed bodies on very short footstalks. I measured some of the bead-like bodies, in Lepas anserifera, and they were 1/2000 of an inch in diameter, and the footstalks three or four times as long as the elongated heads. These heads, of whatever shape they may be, have an opaque, and, I believe, glandular centre; I could not make out with certainty an aperture at their ends, but, I believe, such exists, and they seem to secrete a substance, which hardens into a strong membrane, serving to unite the crest of the frænum to the edges of the lamellæ. In one case, this bit of membrane seemed formed of a woven mass of threads. These little glandular bodies, with the membrane formed by them, are cast off at each exuviation, and new glands formed on the crest of the frænum underneath. In some species of Pollicipes, (viz., P. cornucopia and elegans,) the fræna, though present and large, are functionless and destitute of the glands: I believe, they exist in this same functionless condition, and in rather a different position in the

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sessile Cirripedes, and that in this family they serve as Branchiæ.

The above-described method by which Cirripedia lay their eggs, namely, united together in a common membrane, placed between their old outer and new inner integuments, and the manner in which the lamellæ, when thus formed, are retained for a time fastened to the fræna, and are then cast off, appears to me very curious. In some of the lower Crustacea, it is known, that the ova escape by rupturing the ovisacs formed by the protruded ovarian tubes, and this is the nearest analogy with which I am acquainted. The ova are impregnated (as I infer from the state of the vesiculæ seminales), when first brought into the sack, and whilst the membrane of the lamellæ is very tender: the long probosciformed penis seems well adapted for this end. In the male of Ibla Cumingii, which has not a probosci-formed penis, the whole flexible body, probably, performs the function of the penis: in Scalpellum ornatum, however, the spermatozoa must be brought in by the action of the cirri, or of the currents produced by them. That cross impregnation may and sometimes does take place, I infer from the singular case of an individual, in a group of Balani, in which the penis had been cutt off, and had healed without any perforation; notwithstanding which fact, larvæ were included in the ova.

Exuviation; Rate of Growth; Size.—I have had occasion repeatedly to allude to the exuviation of the Lepadidæ: with the exception of the genus Lithotrya,* in which the calcareous scales on the peduncle, together with the membrane connecting them, is cast off, neither the valves nor the membrane uniting them, nor that forming the peduncle with its scales and styles, are

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

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moulted; but the surface gradually disintegrates and is removed, perhaps sometimes in flakes, whilst new and larger layers are formed beneath. In Scalpellum, I ascertained that the new membrane, connecting together the newly-formed calcified rims under the valves of the capitulum, was formed as a fold, with the articulated spines which it bears, all adpressed in certain definite directions. This fold of new membrane, when the old membrane splits and yields, of course expands, and thus the size of the capitulum is increased. In the peduncle, lines of splitting can seldom be perceived, except, indeed, in the sub-globular, embedded, downward-growing peduncle of Anelasma, as described under that genus. I do not understand what determines the complicated lines of splitting of the old membrane between the several valves of the capitulum,—without it be simply, that along these lines alone, the old membrane is not strengthened by the new membrane being closely applied under it, the new being formed, as we have just said, in a fold, in order to allow of increase in size. Although, as I believe, there is strictly no exuviation in the outer membranes of mature Lepadidæ, it seems that narrow strips of membrane are cast off from between the valves, for the few first moults, after the final metamorphosis of the larva. I may here remark that, in most sessile Cirripedes, the outside membrane connecting the operculum and shell, is regularly moulted.

The delicate tunic lining the sack, (a mere duplicature of that thick one, forming the outside of the capitulum, and generally transformed into valves,) and the integuments of the whole body, are regularly moulted. With these integuments, the membrane lining the oesophagus, the rectum, and the deep olfactory pouches, and the horny apodemes of the maxillæ, are all cast together. I have seen a specimen of Lepas, in which, from some morbid adhesion, the old membrane lining one of the olfactory pouches had not been moulted, but remained projecting from the orifice as a brown shrivelled scroll. The new

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spines on the cirri (and on the maxillæ) are formed within the old ones; but as they have to be a little longer than the latter, and as they cannot enter these up to their very points, their basal portions are not thus included, but are formed, running obliquely across the segments of the cirri; and what is curious, these same basal portions are turned inside out, like the fingers of a glove when hastily drawn off. After the exuviation of the old spines, the new spines have their inverted basal portions drawn out from within the segments, and turned outside in, so as to assume their proper positions.

All Cirripedia grow rapidly: the yawl of H.M.S. Beagle was lowered into the water, at the Galapagos Archipelago, on the 15th of September, and, after an interval of exactly thirty-three days, was hauled in: I found on her bottom, a specimen of Conchoderma virgata with the capitulum and peduncle, each half an inch in length, and the former 7/20ths in width: this is half the size of the largest specimen I have seen of this species: several other individuals, not half the size of the above, contained numerous ova in their lamellæ, ready to burst forth. Supporting the larva of the largest specimen became attached the first day the boat was put into the water, we have the metamorphosis, an increase of length from about ·05, the size of the larva, to an whole inch, and the laying of probably several sets of eggs, all effected in thirty-three days. From this rapid growth, repeated exuviations must be requisite. Mr. W. Thompson, of Belfast, kept twenty specimens of Balanus balanoides, a form of much slower growth, alive, and on the twelfth day he found the twenty-first integument, showing that all had moulted once, and one individual twice within this period. I may here add, that the pedunculated Cirripedes never attain so large a bulk as the sessile; Lepas anatifera is sometimes sixteen inches in length, but of this, the far greater portion consists of the peduncle. Pollicipes mitella is the most massive kind; I have seen a specimen with a capitulum 2·3 of an inch in width.

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

Comparing the pedunculated and sessile Cirripedes, it is, I think, impossible to assign them a higher rank than that of Families. The chief difference between them consists, in the Lepadidæ, in the presence of three layers of striæless muscles, longitudinal, transverse and oblique, continuously surrounding the peduncle, but not specially attached to the scuta and terga; and on the other hand, in the Bala-

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nidæ, of five longitudinal bundles of voluntary muscles, with transverse striæ, fixed to the scuta and terga, and giving them powers of independent movement. In the Lepadidæ, the lower valves, or when such are absent, the membranous walls of the capitulum, move with the scuta and terga when opened or shut; and the lower part of the capitulum is separated by a moveable peduncle from the surface of attachment; in the sessile Cirripedes, the lower valves are firmly united together into an immovable ring, fixed immovably on the surface of attachment. I will not compare the softer parts, such as the cirri and trophi, of the Lepadidæ with those of the Balanidæ, as my examination of this latter family is not fully completed: I will only remark, that there is a very close general resemblance, more especially with the sub-family Chthamalinæ.

Geographical Range; Habitats.—The Pedunculated Cirripedes extend over the whole world; and most of the individual species have large ranges, more especially, as might have been expected, those attached to floating objects; excepting these latter, the greater number inhabit the warmer temperate, and tropical seas. Of those attached to fixed objects, or to littoral animals, it is rare to find more than three or four species in the same locality. On the shores of Europe I know of only three, viz., a Scalpellum, Pollicipes, and Alepas. At Madeira (owing to the admirable researches of the Rev. R. T. Lowe), two Pæcilasmas, a Dichelaspis, and an Oxynaspis are known. In New Zealand, there are two Pollicipes and an Alepas, and, perhaps, a fourth form. From the Philippine Archipelago, in the great collection made by Mr. Cuming, there are a Pæcilasma, an Ibla, a Scalpellum, Pollicipes, and Lithotrya. Of all the Lepadidæ, nearly half are attached to floating objects, or to animals which are able to change their positions; the other half are generally attached to fixed organic or inorganic bodies, and more frequently to the former than to latter. Most of the species of Scalpellum are in-


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habitants of deep water; on the other hand, most of Pollicipes,* of Ibla, and Lithotrya are littoral forms. The species of Lithotrya have the power of excavating burrows in calcareous rocks, shells, and corals; and the singular manner in which this is effected, is described under that genus. Anelasma has its sub-globular peduncle deeply embedded in the flesh of Northern Sharks; and I have seen instances of the basal end of the peduncle of Conchoderma aurita, being sunk into the skin of Cetacea; in the same way the point of the peduncle in the male of Ibla, is generally deeply embedded in the sack of the female. I believe in all these cases, the cementing substance affects and injures the corium or true skin of the animal on which the creature is parasitic, whilst the surrounding parts, being not injured, continue to grow upwards, thus causing the partial embedment of the Cirripede. In the case of Anelasma, we have growth at the end of the peduncle, and consequently downward pressure, and this may possibly cause absorption to take place in the skin of the shark at the spot pressed on.

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

* I am informed by Mr. L. Reeve that Pollicipes mitella is eaten on the coast of China; and Ellis states ('Phil. Trans.,' 1758) that this is the case with P. cornucopia on the shores of Brittany. It is well known that the gigantic Balanus psittacus, on the Chilian coast, is sought after as a delicacy; and I am assured, by Mr. Cuming, that it deserves its reputation.

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Genus—LEPAS. Plate I.

LEPAS. Linnæus.* Systema Naturæ, 1767.

ANATIFA. Brugière.† Encyclop. Method. (des Vers), 1789.

ANATIFERA. (Lister) et plerumque Auctorum Anglicorum.

PENTALASMIS. (Hill.) Leach. Journal de Physique, July, 1817.

PENTALEPAS. De Blainville. Dict. des Sci. Nat., 1824.

DOSIMA. J. E. Gray. Annals of Philosophy, vol. x, 1825.

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

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

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

Distribution.—Mundane; attached to floating objects.

Description.—Capitulum flattened, sub-triangular, composed of five approximate valves. The valves are

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

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

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

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

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

Carina.—This is always very narrow and curved, concave within, often carinated and barbed exteriorly; it extends upwards between the terga for one half or two thirds of their length: at the lower extremity it ends (with the exception of L. fascicularis), in a small fork (Pl. I, fig. 1, a, b) rectangularly inflected and embedded in the membrane, beneath the basal margin of the scuta. From comparing this lower part of the carina in L. australis (fig. 5a), with the same part in some of the species of the allied genus Pæcilasma, it would appear that the fork is formed by an oblong disc, more and more notched at the end, and with the rim between the two points more or less folded backwards: conformably with this view, in very young specimens of L. australis, instead of a large and sharp fork, there is a small disc. The only use of the fork appears to be to give firm attachment to the membrane uniting the valves and peduncle. In L. fascicularis, instead of a fork, there is a broad, oblong disc (figs. 6, 6a), rectangularly inflected; it is much longer than the fork, in proportion to the upper part of the carina; the disc is not more deeply embedded than the upper part. The umbo (and primordial valve when

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distinguishable,) of the carina is seated just above the embedded fork (or disc in L. Fascicularis), at the point where the inflection takes place; hence the main growth of the carina is upwards,—the fork, however, being of course, likewise added to at its point: in L. fascicularis, the growth is both upwards and downwards.

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

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

Filamentary Appendages.—These are attached to beneath the basal articulation of first pair of cirri; they vary in the several species, from one to five or six on each side, the lowest being always the longest. Several of them are occupied by testes. In L. pectinata, generally, not even one is developed. They are subject to great

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variation in their proportional lengths, and in number, in the same species. These organs have generally been considered to serve as branchiæ; I see no reason to believe that they are more especially designed for this end, than is the general surface of the body.

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

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

Caudal Appendages (Pl. X, fig. 18 b), very small, either blunt or pointed, and quite destitute of spines.

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

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

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

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1. LEPAS ANATIFERA. Tab. I. fig. 1. (var.)

L. ANATIFERA. Linnæus. Systema Naturæ, 1767.

ANATIFA vel ANATIFERA vel PENTALASMIS lævis*, plerumque auctorum.

—ENGONATA (!).† Conrad. Journal Acad. Nat. Sc. Philadelphia, vol. vii, 1837, p. 262, Pl. XX, fig. 15.

—DENTATA (var.) Brugière. Encyclop. Meth. (des Vers), 1789.

PENTALASMIS DENTATUS (var.) Brown. Illust. Conch., Pl. lii, fig. 5.

ANATIFA . . . . . Martin St. Ange. Mem. sur l'organisation des Cirripedes, 1835.

L. valvis aut lævibus aut delicate striatis: è duobus scutis, dextro solùm dente interno umbonali instructo; pedunculi parte superiore fuscâ.

Valves smooth, or delicately striated. Right-hand scutum alone furnished with an internal umbonal tooth: uppermost part of peduncle dark-coloured.

Filaments, two on each side.

Var. (a). Fig. 1. Scuta and terga with one or more diagonal lines of dark greenish-brown, square, slightly depressed marks.

Var. (b). (Fig. 1 b.) Carina strongly barbed.

Extremely common; attached to floating timber, vessels, seaweed, bottles, &c., and to each other, in the Atlantic Ocean, Mediterranean, West Indies, Indian Ocean, Philippine Archipelago, Sandwich Islands, Bass's Straits, Van Diemen's Land.

General Appearance.—Valves white, more or less translucent and thick, with a tinge of blueish-grey, from the underlying corium; sometimes brownish cream-coloured, rarely with a tint of purple. Surfaces smooth,

* As this, though the commonest species, has never been defined, I give only a few synonyms and references, it being quite impossible to distinguish, in any published description, this species from A. Hillii of Leach; this latter species I recognise under this name only from having authentic specimeus from the British Museum, as Leach overlooked every one of the real diagnostic characters.

† I have used, in conformity with botanists, the mark of interjection, to show that I have seen an authentic specimen.

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with traces of very fine lines radiating from the umbones, sometimes rather plain on the basal part of the scuta. Length in proportion to the breadth of the capitulum variable, owing to the varying degree to which the scuta and terga have their apices produced. Scuta with the occludent margin either considerably curved or nearly straight. The internal tooth of the right-hand scutum, close to the umbo, varies in size and form, being either pointed, square, or obliquely truncated on either side, or it has a notch on the summit; internal basal rim of the scuta either plainly developed or nearly absent. In many specimens (Pl. I, fig. 1), on the scuta, or on the scuta and terga, (and sometimes more on one side of the individual than on the other,) a nearly straight line, running diagonally across the capitulum, of slight, quadrilateral depressions, of a dirty greenish colour, with the edges blending away, is either conspicuously developed, or can only just be discerned. These marks increase in size from the umbones to the margins of the valves. There are sometimes two or even three rows on the scuta. They are formed by the retention of a portion of the chitine membrane, which is cast off the rest of the surface; the margins of the valves are occasionally notched slightly on the line of marks; there is no difference along this line in the underlying corium. Specimens both with and without a barbed carina are thus characterised. Carina; the interspace between the carina and the scuta and terga is not wide. The carina exteriorly, is either convex and smooth, or furnished with knobs or with extremely sharp, long teeth (Pl. I, fig. 1 b); small specimens, with the capitulum under half an inch in length, are generally most strongly barbed.* Apex more or less acuminated; width and thickness variable; sides strongly furrowed. Fork (fig. 1 a) generally less wide than the widest upper part of the valve, with the two prongs diverging from each

* Mr. W. Thompson found that 15 specimens, out of about 200, attached to a vessel which came from New Orleans into Belfast, had their carinas barbed.

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other at less than a right angle; their sharpness and precise form variable; rim between them reflexed (figs. 1 a and b), making a slight notch behind. Peduncle smooth, wrinkled, length in proportion to that of the capitulum varying, from barely equalling it, to six or seven times as long. I have noticed a specimen including mature ova, with a capitulum under half an inch long.

Filamentary Appendages;—never more than two on each side, with sometimes only one developed; of variable length; one seated on the flank of the prosoma, under the first cirrus; the second close under the basal articulation of this cirrus, on the posterior face of a slight swelling: these appendages correspond with g and h in Fig. 4, Pl. IX.

Mouth.—Mandibles (Pl. IX, fig. 5), with, as usual, five teeth, all pointing downwards. Maxillæ (Pl. IX, fig. 9), with the lower step of variable width compared to the two upper steps. Cirri; posterior cirri with segments (fig. 26) bearing six pair of spines; intermediate fine spines rather long; first cirrus, anterior ramus longer by only about two segments than the posterior ramus; second cirrus with anterior ramus, with very broad transverse rows of bristles; spine-bearing surfaces considerably protuberant; caudal prominences smooth, rounded.

Size.—The largest specimen which I have seen had a capitulum two inches in length; the longest, including the peduncle, was sixteen inches.

Colours.—Calcareous valves already described. Edges of the orifice bright scarlet orange; basal edges of the scuta, and sometimes of all the valves, with a torn border of orange membrane. Interspaces between the valves dull orange-brown. Peduncle darkish purplish-brown, with the lower part sometimes pale; chitine membrane itself tinted orange; in young specimens, peduncle pale, the colour first appearing in the uppermost part, close under the capitulum; this upper part is often darker than the other parts, and never orange-coloured, as in L. Hillii and L. anserifera. Sack internally dark purplish lead-

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colour, sometimes with a tinge of orange, darkest under the growing edges of the valves; body of animal pale purplish lead-colour. The four posterior cirri blackish purple; the second, and often the third cirrus, appear as if the colour had been laterally abraded off; these latter cirri have sometimes a tinge of orange. In very young specimens, the cirri are only barred with purple. The ova and the contents of the ovarian tubes are of a beautiful azure blue, becoming yellow in spirits.

In museums a vast amount of difference is seen in the colours of this species, caused by the method of preparation: if dried without having been in spirits, and subsequently kept dry, the orange tint round the orifice is preserved; if kept long in spirits, this is quite lost; but sometimes in specimens in spirits the colour of the membrane of peduncle is preserved and rendered pinker. The colours of the sack and animal are either quite discharged or rendered extremely dark. The valves themselves also often become more opaque. In some specimens well preserved in spirits, the sack and cirri were purplish-brown or lead-colour, tinted with dirty green, or orange, or bright yellow, or brick-red.

General Remarks.—From the foregoing description it will be seen how extremely variable almost every part of this species is. I find, in the British Museum, ten distinct specific names given by Dr. Leach to different varieties, or rather to different specimens, for some of them are undistinguishable. A specimen from the Sandwich Islands, sent by Mr. Conrad to Mr. Cuming, is marked A. engonata.

In looking over a large collection of specimens in a museum, the most distinctive characters appear at first to be the colours, the dentation or barbed condition of the carina, the row of square marks on the scuta and terga, and the more or less produced form of the whole capitulum: all these characters are absolutely worthless as distinctive characters, and blend into each other. In a fresh condition, the colours of this species, and of

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L. anserifera and L. Hillii are surprisingly alike, though in L. anatifera alone, the uppermost part of the peduncle is dark. As far as I have seen, the smoothness of the valves, together with the presence of a tooth beneath the umbo, on the right-hand scutum, and its entire absence on the left side, (in other species it is smaller on this, than on the right-hand side,) is an unfailing diagnostic mark. I believe this species is always attached to floating objects, though there are some very young specimens in the British Museum, collected by Sir G. Grey, adhering to sandstone, but this may have been buoyed up by some large sea-weed. Mr. Peach has given me the particulars of two instances, in which, after gales of wind, this species, of nearly full size, adhering to apparently freshly broken-off Laminariæ, has been cast upon the coast of England and Scotland.

2. LEPAS HILLII. (Pl. I. fig. 2).

ANATIFA vel PENTALASMIS LæVIS (!) plerumque auctorum.

PENTALASMIS H ILLII (!). Leach. Tuckey's Congo Expedit. p. 413, 1818.

—CHELONIæ (!) Ib. Ib.

ANATIFA TRICOLOR (?). Quoy et Gaimard. Ann. des Sc. Nat., 1st series, tom. x, 1827, Pl. VII, fig. 7, et Voyage de l'Astrolabe, Pl. Xciii, fig. 4.

—SUBSTRIATA (!). Conrad. Journal Acad. Nat. Sc., Philadelphia, vol. vii, 1837, p. 262, Pl. X, fig. 14.

L. valvis lævibus; scutorum dentibus internis umbonalibus nullis; carinâ à cæteris valvis, furcâ etiam a scutorum basali margine, paululum distante; pedunculi parte superiore aut pallidâ aut aurantiacâ.

Valves smooth; scuta destitute of internal umbonal teeth; carina standing a little separate from the other valves, with the fork not close to the basal margin of the scuta; uppermost part of peduncle either pale or orange-coloured.

Filaments three on each side.

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Extremely common; attached to ships' bottoms, from all parts of the world; on floating timber; associated with L. anatifera and L. anserifera. Mediterranean. Attached to turtles, in the Atlantic, lat. 30° north. West Indies. Falkland Islands. "South Seas," collected by A. Menzies. Port Stephen, Australia.

General Appearance.—Capitulum laterally flat; length varies in proportion to the breadth; valves white, somewhat translucent, moderately thick, very smooth, but with faint traces of radiating lines; in some varieties, surface rather irregular along the zones of growth. Scuta without any internal teeth, and with scarcely any trace of the internal basal rim; upper angle little acuminated; the occludent margins of the two scuta stand rather separate from each other, showing a wide space of corium between them: these margins are arched and protuberant, but with the lower part a little hollowed out; basal margin a little curved. In one specimen alone, I saw a trace of a diagonal line of square coloured marks, like those common in L. anatifera. Terga rather broad, with the basal angle not much acuminated. The degree of prominence and outline of the double occludent margin varies very much. Carina, separated by a rather wide space from the scuta and terga; of very varying shape, the upper part not much acuminated, generally very flat, sometimes exteriorly marked by a central depressed line; never barbed; occasionally, (in a specimen from Australia,) middle part so wide as almost to become spoon-shaped; on the other hand occasionally of nearly the same width throughout; somewhat constricted above the fork. Fork deeply embedded as usual; situated, in fresh specimens, a little way beneath the basal margins of the scuta, instead of touching them, as in the other species; forks of varying width, not so abruptly inflected as in many species; sometimes much narrower than the upper widest part of the valve, sometimes nearly twice as wide; prongs of fork not very sharp, diverging at about a right angle, with the rim between them reflexed. The apex of the carina extends up between the terga for barely half their length,

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instead of up fully three fourths of their length, as in L. anatifera.

The chitine membrane at the base of the capitulum, especially at the anterior and posterior ends, is covered with beautiful, little, embedded, yellowish beads, about 3/2000 of an inch in diameter; above this, on each side of the carina, there is a space with similar but smaller little spheres, and still higher up still minuter ones; others occur on different parts of the capitulum; these spaces are seen to be distinctly separated from each other, and present a beautiful appearance under a high power.

Peduncle, as long as, or rather longer than, the capitulum: in one set of specimens, however, it was thrice or four times as long as the capitulum. The peduncle, in some specimens, was conspicuously covered with transverse plates of yellowish hard chitine.

Filamentary Appendages.—Three on each side; one on the flank of the prosoma, with a pair beneath the basal articulation of the first cirrus; relative lengths various, but the posterior filament of the pair under the cirrus, is the shortest. Mouth; palpi not much acuminated; maxillæ step-formed, but with the upper or first step in some specimens indistinct, or forming a curve. Cirri; the segments of the first cirrus and of the posterior arm of the second cirrus are highly protuberant, the protuberances sometimes equalling half the thickness of the segments themselves. Caudal appendages smooth, rounded.

Size.—The largest specimen which I have seen, in the collection of Mr. Cuming, had a capitulum 1 1/10th of an inch long, and 1 1/14 wide; therefore not quite equalling in size the largest specimens of L. anatifera.

Colours.—When fresh, valves blueish-grey from the underlying corium, edges of all the valves and round the orifice, and round the top of the peduncle, bright orange-yellow, passing into the finest scarlet, and varying slightly in tint in different specimens. Space between the carina and the other valves, and between the occludent margins of the scuta, rich purplish-brown; peduncle

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either pale or purplish-brown, or only clouded on the sides with the same. In young specimens, peduncle nearly colourless; and in those under a quarter of an inch long in the capitulum, the top of the peduncle has not acquired its orange tint. Sack pale, leaden-purple, body the same, but paler and more reddish; cirri (but only the tips of first pair) tinted with fine golden orange. Immature ova in peduncle beautiful blue. After being long kept in spirits, the colours are changed, weakened, or discharged, as in L. anatifera and L. anserifera, and the valves become opaque. In some long-kept specimens the corium everywhere had become pale brown; more usually it assumes a dirty purplish lead-colour.

Monstrous Variety.—Amongst a set of ordinary specimens from a ship from Genoa, sent me by Mr. Stutchbury, there were three, one full-grown and two very young, with the whole capitulum, (and likewise with the scuta and terga taken separately,) not above half the usual length in proportion to the breadth. Neither the colours nor animal in this variety presented any difference.

General Remarks.—This species is almost universally confounded with L. anatifera. Quoy and Gaimard, however, appear to have distinguished it, under the name of A. tricolor, from its colours. Leach named it accidentally, for he specifies not one distinctive character, and besides his two published names, he has appended two other names to specimens in the British Museum. A specimen, from the Sandwich Islands, sent by Mr. Conrad to Mr. Cuming, is marked A. substriata. In a dry state, from the shrinking of the membranes, and consequent approach of the carina to the other valves, and of the fork to the basal margin of the scuta, it is most difficult to distinguish this species, though so decidedly distinct, from L. anatifera; the absence, however, of a tooth on the under side of the right-hand scutum is at once characteristic. Even in specimens kept in spirits, in which there has been no shrinking, but in which the colours have changed, and taking into

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account the variation in the carina and upper part of the terga, this species is not always readily distinguished from L. anatifera, without opening the valves and looking for the right-hand tooth of the latter. In fresh specimens, the orange ring at the top of the peduncle, and the broad purplish interspace between the carina and other valves, are characteristic. In all states, the filamentary appendages offer a good character.

3. LEPAS ANSERIFERA. Pl. I, fig. 4.

L. ANSERIFERA, Linnæus. Syst. Naturæ, 1767.

ANATIFA STRIATA. Brug. Encyclop. Meth. (des vers), Pl. clxvi, fig. 3.

PENTALASMIS DILATATA! (young). Leach. Tuckey's Congo Expedit., p. 413, 1818.

ANATIFA SESSILIS (?). Quoy et Gaimard. Voyage de l'Astrolabe, Pl.xciii, fig. 11.

LEPAS NAUTA.* Macgillivray. Edin. New Phil. Journ., vol. xxxviii, p. 300.

P ENTALASMIS ANSERIFERUS. Brown. Illust. Conch., 1844, Pl. li, fig. 1.

L. valvis approximatis leviter sulcatis (tergis præcipuè) ; scuto dextro dente forti interno umbonali, lævo aut dente exigno, aut merâ cristâ instructo; margine occludente arcuato, prominente: pedunculi parte superiore aurantiacâ.

Valves approximate, slightly furrowed, especially the terga; right-hand scutum with a sytrong internal umbonal tooth; left-hand with a small tooth, or mere ridge; occludent margin arched, protuberant: uppermost part of peduncle orange-coloured.

* Professor Macgillivray does not consider the species, which he has described under L. nauta, and which I cannot doubt is the same with the present species, as the L. anserifera of Linnæus; but I find it so named in all old collections, and it seems to agree very well with Linnæus's description. There has been much groundless confusion about this species; I have no hesitation in giving A. striata, of Brugière, as a synonym, though I have received from Paris the Lepas pectinata of this volume, named as the A, striata; and on the other hand, Poli has incorrectly called a common variety of L. pectinata by the name of L.anserifera.


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Filaments five or six on each side.

Var. (dilatata, young); valves rather thin, finely furrowed, often strongly pectinated; scuta broad, with the occludent margins much arched, making the space wide between this margin and the ridge connecting the umbo and the apex: carina often barbed.

Common on ships' bottoms from the Mediterranean, West Indies, South America, Mauritius, Coast of Africa and the East-Indian Archipelago. Central Pacific Ocean. China Sea. Chusan. Sydney. Attached to pumice, various species of fuci, Janthinæ, Spirulæ; often associated with L. anatifera and L. Hillii, and, in a young state, with L. fascicularis.

General Appearance.—Capitulum more or less elongated relatively to its breadth; in two specimens, with scuta of equal width, one was longer than the other by the whole of the occludent margin of the terga. Valves white, thick, (in young specimens sometimes diaphanous and thin,) closely approximate to each other; surfaces furrowed to a very variable amount. Terga generally more plainly furrowed than the scuta, of which the basal portion is generally less furrowed than the upper part; ridges, often rough, generally much narrower than the furrows: in half-grown specimens (var., dilatata of Leach,) the ridges are frequently denticulated, and there is even sometimes a row of bead-like teeth along the basal margins of the scuta. The ridges vary much, sometimes alternately wide and narrow; in two specimens of equal size, there were, in one, thirty-two ridges, and in the other only eighteen, on the scutum.

Scuta, with the occludent margin rounded and protuberant to a variable degree, but always leaving a rather wide space between the margin, and the ridge which runs from the umbo to the apex; apex pointed. Right-hand internal tooth considerably larger than that on the left, which is often reduced to a mere ridge; internal basal rim thick, sometimes furrowed along its upper edge, but of variable thickness, sometimes not extending as far as the baso-carinal angle. Terga, some-

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times equalling, sometimes only two-thirds of, the length of the scuta; in young specimens, the two occludent margins form of right-angle with each other; in older specimens they form less than a right-angle, and hence the portion of valve thus bounded is unusually protuberant. Carina, within deeply concave; exterior sides finely furrowed longitudinally, generally denticulated; valve only slightly narrowed in above the fork, of which the prongs diverge at an angle of 90°, or rather more, and are wider than the widest upper part of the valve; rim between the prongs reflexed; the heel or external angle, just above the fork, sometimes considerably prominent. I have seen only a single large specimen with its carina barbed. In half-grown specimens, (var. dilatata, Leach,) the carina is often strongly barbed, with the upper point much acuminated, the fork about twice as wide as the widest upper part, and the prongs diverging at rather more than a right-angle. In some specimens, especially very young ones, there are at the base of the carina, above the fork, some strong, downward-pointed, inwardly-hooked, calcareous teeth; such occur also in some specimens along the basal margins of the scuta, two of these hooked teeth under the umbones of the scuta being larger than the rest: specimens conspicuously thus characterised came from the Navigator Islands; in these, I may add, the acutely triangular primordial valves were quite plan.

Peduncle, generally about as long as the capitulum; in young specimens generally short.

Filamentary Appendages, generally five, sometimes six, on each side; one is seated on the side of the prosoma, and the four others placed in pairs beneath the basal articulation of the first cirrus; the lowest posterior filament of the four generally is the largest. In young specimens, having a capitulum only half an inch long, the upper pair of the four often is not developed, or is represented by mere knobs. The mouth presents no distinctive characters. Cirri, with the longer ramus of the first pair almost equal to the shorter arms of the

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second pair; spine-bearing surfaces only slightly protuberant. Caudal appendages smooth, curved, pointed.

Size.—The largest specimen which I have seen, had a capitulum one inch and a half in length.

Colours—The white valves are edged with bright orange membrane; and are so close to each other that no interspaces, coloured from the underlying corium, are left. Peduncle, dark orange-brown, with the uppermost part under the capitulum bright orange all round; the chitine membrane itself being thus coloured. Sack, internally, dark purplish lead-colour. Body and cirri, either nearly white or pale purplish-lead colour, with the arms of the second, third, and fourth cirri, and pedicels of the fifth and sixth, more or less tinted with orange. A specimen preserved during fourteen months in good spirits had only a tinge of orange left round the orifice and round the upper part of peduncle, and on the cirri. In some other specimens, badly preserved, the chitine membrane was quite colourless, and sack and cirri dirty lead-colour. Fresh ova, peach-blossom-red; immature ova, in ovarian tubes, pale pink.

Monstrous Variety.—In Mr. Stutchbury's collection, there was a specimen, with the scuta, broad, smooth, thin, and fragile, without any ridge running from the umbo to the apex, and with the occludent margin reflexed. This seemed caused by the shell having been attacked by some boring animal, and from having supported Balani. In the same specimen the first cirrus on one side was monstrously thick and curled; the second cirrus had its posterior ramus in a rudimentary condition. In Mr. Cuming 's Collection, there are small specimens with the zones of growth overlapping each other, with thick irregular margins, and with the carina distorted.

This species has cost me much trouble: I have examined vast numbers of specimens, from a tenth to half an inch in length, attached to light floating objects, such as Janthinæ and Spirulæ from the tropical oceans, which all resembled each other, and slightly differed

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from the common appearance of L. anserifera: this variety is the Pentalasmis dilatata of Leach; and for a long time I considered it as a distinct species. It differs from L. anserifera, in the less thickness of the valves, in their being more finely and yet plainly furrowed; in the greater width of the scuta; and more especially, of that part of the valve lying between the occludent margin, and the ridge running from the umbo to the apex; in the less elongation of the area in the terga, bounded by the two occludent margins; and, lastly, in the less size of the whole individual. The trophi and cirri are absolutely identical. Lately, however, in carefully going over a great suite of specimens, all the above few disctinctive characters broke down and insensibly graduated away; and I am convinced that this form is only a variety of L. anserifera; its different aspect being caused partly by youth, but chiefly, I suspect, from being attached to light objects floating close to the surface of the sea.

The Lepas anserifera can be distinguished by the slight furrows on its valves from all the other species excepting L. pectinata: this latter species can be readily known, by the close proximity in the scuta of the occludent margin, and the ridge extending from the umbo to the apex; by its carina being very narrow above the fork; by the prongs of the fork diverging at an angle of from 135° to 180°; by the thinness of its valves; by the coarseness of the furrows on them; and lastly, by there being at most in L. pectinata only one filamentary appendage beneath the first cirrus.

4. LEPAS PECTINATA. Pl. I, fig. 3.

LEPAS PECTINATA. Spengler. Skrifter Naturhist. Selbskabet, 2, B. 2. H., 1793, Tab. X, fig. 2

—MURICATA (var.) Poli. Test. Utriusque Scicil., vol. i, Pl. VI, figs. 23, 29, 1795.

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LEPAS ANSERIFERA. Poli. Test. Utriusque Scicil., vol. i. Pl. VI, figs. 25-27.

—SULCATA. Montagu. Test. Brit., Pl. I, fig. 6, 1803.

PENTALASMIS SULCATA. Leach. Encyclop. Brit. Suppl., tom. iii, Pl. lvii, 1824.

—SPIRULE (!) (var.) Leach. Tuckey's Congo Expedit. Appendix, 1818.

—RADULA (var.) et SULCATUS. Brown. Illust. of Conchology, Pl. li, figs. 3—6, 1844.

—INVERSUS. Chenu. Illust. Conchy., Pl. I, fig. 14.

ANATIFA SULCATA. Quoy et Gaimard. Voyage de l'Astrolabe, Pl. xciii, figs. 18, 20.*

L. valvis tenuibus, crassè sulcatis, sæpe pectinatis; scutorum cristâ prominente ab umbone ad apicem juxta marginem occludentem pertinente: furcæ carinalis cruribus inter angulos 135° et 180° divergentibus.

Valves thin, coarsely furrowed, often pectinated. Scuta with a prominent ridge extending, from the umbo to the apex, close to the occludent margin; fork of the carina wit the prongs diverging at an angle of from 135° to 180°.

Filaments absent, or only one on each side.

Var. (Pl. I, fig. 3a), upper part of the terga (bounded by the two occludent margins) produced and sharp; surface of all the valves often coarsely pectinated, and with the carina barbed.

Atlantic Ocean, from the North of Ireland to off Cape Horn; common, under the tropics; Mediterranean: attached to wood, cork, charcoal, seaweed, a reed-like leaf, spirulæ cuttle-fish bones, to a bottle together with L. anatifera; to a ship's bottom, Belfast, (W. Thompson.) Often associated with L. fascicularis. Montagu states ('Test, Brit.,' p. 18) that this species is sometimes attached to the fixed Gorgonia flabellum.

General Appearance.—The capitulum varies considerably in length compared to its breadth, caused chiefly by the greater or less production of the occludent portion of the terga; valves thin, brittle; the furrowed surface varies

* I may add, that I have received many specimens incorrectly labelled A. striata, which is properly a synonym of L. anserifera.

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much in character, narrow and broad ridges often alternating; frequently each ridge (buth more especially the ridge running from the umbo to the apex of each scutum, and sometimes that alone,) is covered with prominent, curled, flat, calcareous spines, giving the shell an appearance like that of many mollusca. Other specimens show no trace of these calcified projections. From the thinness of the valves and the depth of the furrows, the margins of the valves are sinuous. Scuta: the ridge running from the umbo to the apex is unusually prominent and curved; it runs very close to the occludent margin, so that, differently from in all the other species, only a very narrow space is left between this margin and the ridge. Internal teeth, under the umbones, either sharp and prominent, or mere knobs; sometimes that on the right side is much larger than that on the left; sometimes they are nearly equal; sometimes that on the left is scarcely distinguishable. Internal basal rim absent, or barely developed.

Terga: these valves have a conspicuous notch to receive the apex of the scuta; the two occludent margins either meet each other at a rectangle, or at a much smaller angle, causing the portion thus bounded to vary much in outline, area, and degree of prominence. This at first led me to think that the P. spirulæ of Leach, in which the point is very sharp and prominent, was a distinct species; but there are so many intermediate forms, that the idea must be given up. I may remark, that in all the species of Lepas, the upper part of the tergum seems particularly variable. The degree of acumination of the basal portion of the tergum also varies; the internal surface sometimes has small crests radiating from the umbo.

Carina, broad, within deeply concave; edges sinuous, externally sometimes strongly barbed; narrow above the fork, which latter is wider than the widest upper part of the valve; prongs sharp, thin, diverging at an angle of from 135° to 180°; the rim connecting the prongs not, or only slightly, reflexed.

Peduncle, narrow, shorter than the capitulum.

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Filamentary Appendages, none, or only one, short, obtuse projection on each side, on the posterior face of the swelling under the first cirrus.

Mouth.—Mandibles, with the inferior point produced into a single pectinated tooth, rarely into two pectinated teeth; on one side of one specimen, there were only four instead of five teeth. Palpi very narrow. Maxillæ highly variable; they may be described as formed of five steps, of which the two lower ones are generally united into a single one, divided by a mere trace of a notch; or with the three lower steps blended into an irregular, projecting surface, and with even the fourth step indistinct. I have seen these two extreme forms on opposite sides of the mouth of the same individual,-on one side the maxillæ being regularly step-form, on the other the whole inferior part forming an almost straight edge, standing high up above the first notch or step which bears the two upper great spines.

Cirri.—First pair rather far removed from the second pair, with the longer ramus about three-fourths of the length of shorter ramus of second cirrus; spine-bearing surfaces, hardly at all protuberant; lateral marginal spines on the posterior cirri rather long; caudal appendages smooth, rounded, extremely minute: penis very spinose.

Size.—Capitulum in the largest specimen, six-tenths of an inch long; only a few arrive at this size.

Colours, after having been kept in spirits,-sack and cirri, especially first cirrus, clouded with pale purple; peduncle brownish; valves appear blueish in specimens not long preserved, but in specimens kept longer they become perfectly and delicately white.

General Remarks.—Under the head of L. anserifera, I have made some remarks on the diagnostic characters of this species. In the thinness of the valves,-form of the carina, with the rim connecting the prongs being not, or scarcely, reflexed,-and in the shortness and narrowness of the peduncle, there is some approach to L. australis, and thence to L. fascicularis. In the form of the maxillæ,

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—in one specimen having the mandible on one side bearing only four teeth,-and in the frequent absence of filamentary appendages, there is some approach to the genus Pæcilasma; but there is no such approach in the characters derived from the capitulum. We have seen that, as in so many other species of this genus, most of the parts are variable, and this is the case to a most unusual extent in the form of the maxillæ. Dr. Leach has attached eight specific names to the specimens preserved in the British Museum.

5. LEPAS AUSTRALIS. Pl. I. fig. 5.

L. valvis glabris, tenuibus, fragilibus, scutorum dentibus umbonalibus utrinque internis; carinæ parte superiore latâ, planâ, suprâ furcam valdè constrictâ; furcæ cruribus latis, planis, tenuibus, acuminatis, intermedio margine non reflexo.

Valves smooth, thin, brittle; scuta with internal umbonal teeth on both sides. Carina with the upper part broad, flat; much constricted above the fork, which has wide, flat, thin, pointed prongs, with the intermediate rim not reflexed.

Filaments, two on each side.

Common on Laminariæ in the whole Antarctic Ocean: Bass's Straits, Van Diemen's Land: Bay of Islands, New Zealand, lat. 35° S.: lat. 50° S., 172° W. coast of Patagonia, lat. 45° S.: attached to bottom of H. M. S. Beagle, lat. 50° S., Patagonia: attached to a Nullipora, (I presume a drift piece) British Museum.

General Appearance.—Capitulum rather obtuse and thick; valves thin, brittle, approximate, either white and transparent, or dirty-brown and opaque; or sometimes tinted internally with purple (perhaps the effects of being preserved in spirits); surface plainly marked by lines of growth, rarely marked with traces of lines radiating from the umbones. Scuta with teeth on both sides,

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nearly equal; internal basal rim rather wide, sometimes furrowed; basal margin considerably curved inwards. Terga rather wide; basal angle blunt; angle formed by the two occludent margins blunt and rounded. Carina (fig. 5 a) with the apex blunt, flat; the middle part generally very broad; much constricted above the fork, where it is internally deeply concave, and externally carinated; fork twice as broad as the broadest upper part of the valve; with the prongs flat, broad, thin, pointed, diverging at about an angle of 75°, with the intermediate rim not at all reflexed; the fork generally not deeply imbedded in the chitine membrane of the peduncle, so as to be quite easily visible externally; sometimes there is an internal, transverse, depressed line on the fork. In young specimens, with the capitulum about a quarter of an inch long, the fork of the carina is not developed, the lower slightly inflected portion consisting simply of an oval plate, twice as wide as the upper part. Until I had carefully examined a perfect series, showing the gradual changes in this part, I did not doubt that the young specimens formed a distinct species, and named it accordingly: the shortness of the penis first made me perceive that the specimens were immature. At this early age, I may add, the filamentary appendages were not developed. Peduncle either quite short, or as long as the capitulum, close under which it is considerably constricted all round.

Filamentary Appendages.—Two on each side; one long, tapering, placed on the prosoma (in one specimen represented by a mere knob), and the second shorter, situated on the posterior margin of the swelling beneath the first cirrus.

Mouth.—Maxillæ, with three large spines at the upper angle, and with the first step distinct, but narrow; mandibles with five teeth; in young specimens the inferior point ends in a single spine; sides of the supra-oral cavity very hairy; the membrane, forming the inner fold of the labrum, yellow and thickened in the form of a spoon.

Cirri.—In the posterior cirri there are, at the upper

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lateral edges of the segments on both sides, small spines; the segments in the first cirrus, and in the broad anterior ramus of the second cirrus, are hemispherically and considerably protuberant. Caudal appendages smooth.

Size.—The largest specimen had a capitulum one inch long.

The Colours (after having been long in spirit) of the valves have already been given; sack and peduncle dirty yellowish-brown, with the parts corresponding to the margins of the valves much darker brown, or almost black; segments of the cirri clouded with dark brown; body and pedicels of the cirri dirty yellowish. I have reason to believe that the colours are totally different in living specimens.

Monstrous Varieties.—Most of the specimens from lat. 50° S., on the coast of Patagonia, were more or less deformed, with the successive zones of growth overlapping each other, and forming coarse concentric ridges. The carina in several specimens was laterally distorted.

I have already remarked that this species has some affinity to L. pectinata; but it is much more closely related to L. fascicularis, the affinity being clearly shown by the thinness and translucency of the valves, their convexity, by the width and little acumination of the upper part of the carina, by the width of the fork, and by its not being deeply imbedded. In young specimens, moreover, before the fork is fully developed, there is a remarkable similarity between the two species, in the form of this lower part of the carina. Again, the narrowness and inflection of the peduncle under the capitulum in L. australis, and lastly, the lateral marginal spines on both sides of the segments of the posterior cirri, all clearly indicate this same affinity to L. fascicularis.

I believe this species is confined to the southern ocean; and perhaps there represents L. fascicularis of the northern and tropical seas. It must, judging from the number of specimens brought home by Captain Sir J. Ross, and from those previously in the British Museum, and from those collected by myself, be a very common species.

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LEPAS FASCICULARIS. Ellis and Solander. Zoophytes, 1786, Tab. xv, fig. 5.

Montagu. Test. Brit. Suppl., 1808, pp. 5, 164.

-CYGNEA. Spengler. Skrifter Naturhist. Selbskabet, Bd. i, 1790, Tab. vi, fig. 8.

-DILATA. Donovan. British Shells, 1804.

PENTALASMIS FASCICULARIS. Brown. Illust. Conch., 1844, Pl. li, fig. 2.

-SPIRULICOLA (!) et D ONOVANI (!) Leach. Tuckey's Congo Expedit., p. 413, 1818.

ANATIFA VITREA. Lamarck. Animaux sans Vertebres.

DOSIMA FASCICULARIS. (!) J. E. Gray. Annals of Philosophy, vol. x, 1825.

PENTALEPAS VITREA. Lesson. Voyage de la Coquille. Mollusca, Pl. xvi, fig. 7, 1830.

ANATIFA OCEANÍCA (!) Quoy et Gaimard. Voyage de l'Astrolabe, Pl. xciii.

L. valvis glabris, tenuibus, pellucidis; carinâ rectangulè flexâ, parte inferiore in discum planum oblongum expansâ.

Valves smooth, thin, transparent; carina rectangularly bent, with the lower part expanded into a flat oblong disc.

Filaments, five on each side; segments of the three posterior cirri with triangular brushes of spines.

Var. (Donovani, of Leach.) Carina with the upper part flat, spear-shaped, externally with a narrow central ridge.

Var. (Villosa. Pl. I, figs. 6 b, c.) Valves placed rather distant from each other; carina extremely narrow, with the upper part of nearly the same width throughout; terga with the lower part much acuminated; body of animaly finely villose.

Coasts of Great Britain and France; Baltic Sea, according to Montagu Southern United States (from Agassiz); tropical Atlantic Ocean; East-

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Indian Archipelago, off Borneo and Celebes; Pacific Ocean, between the Sandwich and Mariana Archipelagos; New Zealand: attached to fuci, Spirulæ Janthinæ, Velellas, often to feathers and cork; often associated with the young of L. anserifera, (var. dilatata,) and L. pectinata.

General Appearance.—Capitulum highly variable in all its characters; thick and broad in proportion to its length, but the breadth is variable, —in some specimens, the capitulum being longer by one-fifth of its total length than broad; in others, one-fifth broader than long. Valves generally approximate; in some varieties, however, from the narrowness of the carina and terga, the valves stand far apart, there being an interval between the carina and scuta of nearly half the breadth of the latter. Valves excessively thin, brittle, transparent, colourless, smooth, but generally sinuous along the zones of growth, which are conspicuous: valves generally covered throughout by thin chitine membrane, which is thickly clothed, especially in the interspaces between the valves, with minute spines, barely visible to the naked eye. Scuta with the lower part of the tergo-carinal margin extremely protuberant; occludent margin, more or less, but slightly reflexed, with a depressed line running from the umbo to the apex; basal margin much reflexed, but to a variable extent and at a varying angle, event up to a right angle,-an external rim or collar being thus formed. There are no distinct internal teeth, but the basal margin under the umbones, is more or less distinctly produced into a rounded disc or projection, which is generally not so much outwardly reflexed as the rest of the basal margin: there is no distinct internal basal rim. The primordial valves are generally visible, but they do not lie, as in all other species, close to the basal margin, but a little above it,-the lower reflexed portion having been subsequently developed. Terga flat, with the occludent margin slightly arched, and not, as in the foregoing species, formed of two sides; apex bent towards the carina; width of the lower half highly variable, owing to the varying extent to which the scutal margin is hollowed

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out; in some specimens, the whole lower half beneath the apex of the scuta is of nearly the same width throughout; in other specimens this lower part is spear-shaped. The widest part of the tergum either equals in width, or is only two-thirds of the width of the widest part of the carina beneath its umbo. Carina (Pl. I, fig. 6 a) highly variable in shape, with the part above the umbo either spear-shape and slightly concave within, or nearly flat and furnished with a central external ridge; or the upper part (fig. 6 c) is of equal and extreme narrowness throughout, and deeply concave within, appearing as if only the central ridge had been developed. The part below the umbo, (answering to the fork in the foregoing species,) is about one-third of the length of the whole valve, and generally twice as wide as the upper part, but in the variety with the upper part of the carina equally narrow throughout, the lower part is thrice as wide as the upper; the disc, or lower part, is generally slightly concave within, exteriorly either with or without a central ridge; basal margin rounded; lateral margin more or less curved, according to the form of the upper part. The disc is not more deeply imbedded in membrane than is the upper part of the valve. The heel or umbo is either angular and prominent, or rounded. In very young specimens the carina is simply bowed, instead of being rectangularly bent.

Peduncle,—short, narrow, being abruptly inflected all round under the basal edges of the capitulum; lower part of very variable shape, being often suddenly contracted into a mere thread (fig. 6 b), which sometimes widens again at the extreme end. The external membrane is very thin, and is penetrated by the usual fine tubuli leading to the corium; its surface is wrinkled and destitute of spines, or with extremely few. The peduncle is often completely surrounded by a yellowish ball, (of which I have seen specimens from the coast of England, and from off Borneo), sometimes half as wide as the capitulum, composed of very tender, vesicular, structure-

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less membrane, and of a pulpy substance: perhaps the yellow colour may be owing to long immersion in spirits. Some authors have supposed that the ball was the ovisac of the animal; and for the first few minutes, deceived by the numerous included spores of, as I believe, Bacillariæ, I thought that this was the case; others have supposed that it consisted of some encrusting algæ or other foreign organism; but it is, in reality, a most singular development of the cement-tissue, which ordinarily serves to attach Cirripedes by their bases to some extraneous object, but here surrounding that object and the peduncle, gives buoyancy, by its vesicular structure, to the whole. The membrane of the ball falls to pieces in caustic potash, differently from the chitine membrane of the enclosed peduncle, and this shows that there is some difference in composition from ordinary cement. The ball, when cut in two, exhibits an obscure concentric structure. The whole is excreted by the two cement-ducts, through two rows of orifices, one on each side of the surrounded portion of the peduncle; and I actually traced, in one case, the yellow pulpy substance coming out of the cement-ducts. The upper apertures are in gradation larger than those below them, and they stand a little further apart from each other; these are figured as seen from the outside, much magnified, at Pl. I, fig. 6 d. I did not succeed in finding the cement-glands, but I followed the ducts, of rather large size, running for a considerable distance as usual along and within the longitudinal muscles of the peduncle. Nearly opposite the uppermost aperture, on each side, the duct passes out through the corium, and becomes laterally attached to the outer membrane of the peduncle, at which point an aperture is formed (as in other cases, by some unknown process), thus giving exit to the contents of the duct. Beneath this upper aperture the duct runs down the peduncle, between the corium and the outer membrane, till it comes to the next aperture, to which it is also attached, and so on to all the lower ones; but I

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believe no cement tissue continues to pass out through these lower apertures. Beneath the lowest aperture the two ducts run into the two prehensile antennæ of the larva, which, as usual, terminate the peduncle. The antennæ are attached to some small foreign body in the centre of the vesicular ball, by the usual tough, light brown, transparent cement. The two upper apertures are nearly on a level with the outside surface of the ball; and it was evident that as the animal grows, new apertures are formed higher and higher up on the sides of the peduncle, and that out of these, fresh vesicular membrane proceeds, and grows over the old ball in a continuous layer. It appears that the growth of the vesicular ball is not regular,-that it is not always formed,-and that when formed the whole, or the lower part, sometimes disintegrates and is washed away. As that portion of the peduncle which is enclosed ceases to grow, and has its muscles absorbed, retaining only the underlying corium, whereas the upper unenclosed portion, and likewise, (as it appears) lower portions once enclosed but since denuded, continue to increase in diameter, the peduncle, when the vesicular ball is removed, often has the most irregular outline, contracting suddenly into a mere thread, and then occasionally expanding again at the basal point.

Frequently two or three specimens have their peduncles imbedded in one common ball, of which there is a fine specimen in the College of Surgeons (Pl. I, fig. 6), the ball being about one inch and a quarter in diameter, with a slice cut off. In this specimen, it is seen that the vesicular membrane proceeding from several individuals, unites to form one more or less symmetrical whole, and that the original common object of attachment is entirely hidden. Dr. Coates* gives a curious account of the infinite number of specimens, through which he sailed during several days, in the Southern Atlantic Ocean: the balls appeared like bird's eggs, and were mistaken for some

* Journal of the Acad. Nat. Sc., Philadelphia, vol. vi, p. 138, 1829.

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fucus, which was supposed to have encrusted the scales of the Velellæ, to which the Cirripede had originally become attached. Several individuals had their peduncles imbedded in the same ball, "which floated like a cork on the water." As this species grows into an unusually bulky animal, we here see a beautiful and unique contrivance, in the cement forming a vesicular membranous mass, serving as a buoy to float the individuals, which, when young and light, were supported on the small objects to which they originally had been cemented in the usual manner.

Filamentary Appendages.—Five on each side, of which four lie in pairs at the base of the first cirrus (of these, only three are sometimes developed), and one on the flank of the prosoma.

Mouth.—Palpi much acuminated. Mandibles with five teeth; the first not far remote from the second; inferior point rather broad and finely pectinated. Maxillæ with two large, unequal, upper spines, and four regular steps.

Cirri.—Posterior cirri, with the upper parts of the segments slightly protuberant; in young specimens, the spines can be seen to consist of five pairs, placed in two converging lines in the upper half of each segment, with numerous minute, latero-marginal, and intermediate little bristles: in large specimens, all these latter have so increased in number, that the normal five pair cannot be distinguished, and the front of each segment is covered by a triangular thick brush of bristles, all pointing in the same direction, thus giving a very unusual character to the posterior cirri: the dorsal tuft on each segment consists of six or seven large spines, with from one to three dozen fine ones. First cirrus and anterior ramus of second cirrus with broad brushes of bristles. The pedicels of all the cirri are thickly covered with bristles. Caudal appendages smooth, with rounded summits.

Penis very hairy: vesiculæ seminales purple, much convoluted, lying within the prosoma; testes dendritic, scarcely enlarged at their terminal points, purplish;


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ovigerous fræna large with sinuous margins, the glandular beads being arranged in groups.

Size.—The largest specimen (from the coast of Devonshire) had a capitulum 1·6 of an inch long, and 1·2 broad, and of unusual thickness.

Colours, after having been in spirits: front surfaces of the segments of the cirri and of the pedicels purple. In some specimens from off Borneo, parts of the sack and the interspaces between the two scuta, were of a fine purple. Montagu states, that the whole shell and body of animal, when fresh, are pale blue, with the cirri spotted with brown.

General Remarks.—The extreme variability of this species is remarkable. In the College of Surgeons, there is a group of specimens collected by Mr. Bennett, I believe, in the Atlantic, in which the extreme narrowness of the carina and of the terga (Pl. I, fig. 6, b, c) (with consequent wide spaces of membrane left between these valves), led me, at first, to entertain no doubt, that it was quite a distinct species, which was strengthened by finding that the whole surface of the cirri were villose, with very minute spines; hence I called this variety, villosa. On the closest examination, however, I could detect no other differences, and the narrowness of the carina and terga varied very considerably: moreover, in one of the specimens, which was about intermediate in the form of its valves between this variety and the common form, the surfaces of the cirri were not in the least degree villose. Again, in some other specimens, the terga were as narrow as in Mr. Bennett's, whilst the carina had its usual outline.

In a var. (called by Leach, P. Donovani,) from the Atlantic, under the Equator, the carina is remarkable from the extreme flatness of the upper part, and from the presence of an exterior, narrow, central ridge. In one specimen from Jersey, in the British Museum, the carina made an extremely near approach to this same form.

Affinities.—This species is certainly much the most


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distinct of any in the genus, and Mr. Gray has proposed to separate it under the name of Dosima; but considering the close similarity of the whole organisation of the internal parts, together with the transitional characters afforded by L. australis, I think the grounds for this separation are not quite sufficient. I have remarked, under L. australis, on the affinity between that and the present species. In the carina terminating in a disc (though here not imbedded), there is some slight affinity to Pæcilasma eburnea and crassa, and markedly so in the arrangement of the bristles on the posterior cirri. In the valves being covered with villose membrane, and to a certain extent in the form of the carina and of the occludent margin of the terga, and especially in the two rows of cement-orifices in the peduncle, there is some affinity to Scalpellum.



ANATIFA. J. E. Gray. Proc. Zoolog. Soc., 1848, p. 44.

TRILASMIS. Hinds. Voyage of the Sulphur. Mollusca, 1844.

Valvæ, 3, 5, aut 7, approximatæ: carina solùm ad basales apices tergorum extensa, termino basali aut truncato aut in discum profunde infossum producto: scuta pænè ovalia, umbonibus ad angulum rostralem positis.

Valves, 3, 5, or 7, approximate: carina extending only to the basal points of the terga; with its lower end either truncated or produced into a deeply imbedded disc. Scuta nearly oval, with their umbones at the rostral angle.

Mandibles with four teeth; maxillæ notched, with the lower part of edge prominent; anterior ramus of the

* Ποκιλοσ, various, and ελανμα, plate or valve. I have not been able to adopt Mr. Hinds' name for this genus, as it would be too glaringly incorrect to call a five-valved species, a Trilasmis.

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second cirrus not thicker than the posterior ramus; caudal appendages uniarticulate, spinose.

Generally attached to Crustacea.

I have already given my reasons for instituting and separating this genus from Lepas; as far as the capitulum is concerned, the differences between these genera certainly appear trivial; they consist in the carina not extending up between the terga, and in the lower end being either truncated, or produced into an imbedded disc: the terga have a single occludent margin. The included animal's body differs in more important respects; for both mandibles and maxillæ are very distinct; the cirri of some of the species also differ; and the caudal appendages are here always spinose: there are no filamentary appendages: and lastly, the habits are different.

The genus may be divided into two sections, firstly, P. Kæmpferi and P. aurantia, which have their carinæ basally truncated, the basal angles of their terga cut off, and the anterior rami of their second cirri shorter than the posterior rami; and, secondly, P. crassa, P. fissa, and P. eburnea, which in these several respects are otherwise characterised. The P. eburnea, however, differs rather more from P. crassa and P. fissa, than these two do from each other; but certainly not enough to allow of the retention of Mr. Hinds' genus of Trilasmis. P. crassa, in an especial degree, connects together all the forms.

General Appearance.—Capitulum oval, more or less produced, flat or gibbous; formed of three, five, or seven approximate valves; the lesser number arising from the abortion of the terga, and the greater number from the scuta being divided into two segments. Valves moderately thick, either white or reddish, smooth or striated, and sometimes partly covered by membrane, bearing minute spines. Scuta oval, of varying proportions; the basal margin is generally narrow, and blends into the

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carina-tergal margin; the internal basal rim generally is well developed, sometimes with, and sometimes without internal teeth beneath the umbones. In P. eburnea, and sometimes in P. crassa, there is a line of apparent fissure, and in P. fissa of actual disseverment, running from the umbo to the apex of each scutum, nearly in the line in which a ridge extends in Lepas: the primordial valves of the scuta in these three species, are seated at the basal angles of the lateral and larger segments. The positions of the primordial valves, and the direction of growth in the calcified valves, are, in all the species, the same as in Lepas. In several of the species attached to Crustacea, the two scuta are unequally convex, which is caused, as was pointed out to me by Mr. Gray, by that valve which lies close and nearly parallel to the body of the crab, being least developed. The Terga are either quite absent, or rudimentary as in P. crassa, or pretty well developed as in the other species: the occludent margin is single, and not double as generally in Lepas; the basal angle is either pointed or truncated. The Carina varies considerably in shape, but never extends up between the terga, nor ends downwards in a fork; in the first two species it is truncated; in the others; it terminates in a deeply-imbedded oblong disc, which in P. eburnea seems almost entirely (but of course not quite) to separate the inside of the capitulum from the peduncle; a similar separation is effected in P. fissa, where the imbedded disc is small, by two large teeth on the internal basal rims of the two scuta. The carina is always narrow, and either solid internally or very slightly concave.

Peduncle, is very short and narrow; the membrane is generally ringed with thicker, yellower portions, and often bears very minute spines.

Size.—All the species are small, with a capitulum not exceeding half an inch in length.

Filamentary Appendages.—None.

Mouth.—Labrum generally considerably bullate in

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the upper part, with a row of teeth on the crest. The mandibles have four teeth, with the inferior point narrow and spine-like, or rudimentary and absent. The maxillæ have, under the two or three upper great spines, a deep notch itself bearing spines; beneath this, the lower part is straight and considerably prominent, Pl. X, fig. 15. Outer maxillæ are covered on their inner sides continuously with spines.

Cirri.—The first is sometimes seated very distant from the second. The arrangement of the spines on the posterior cirri varies, to an unusual degree within the limits of the same genus. We have either the ordinary structure of anterior pairs, with single fine intermediate spines (as in P. Kæmpferi and aurantia), or we have the pairs increased by one or two additional longitudinal lateral rows, as in P. eburnea; or we have the front spines forming a single transverse row, as in P. crassa and P. fissa, Pl. X, fig. 29, a. The segments in none of the species are protuberant; the anterior ramus of the second cirrus does not seem to be thicker than the posterior ramus, as is usually the case. The rami of the second, and of most of the other cirri, are unequal in length,—the anterior ramus, contrary to the ordinary rule, being longer in P. eburnea, P. fissa, and P. crassa, than the posterior ramus by several segments; I have hitherto observed this inequality only in the sessile genus Chthamalus.

The Caudal Appendages are small, uniarticulate, and always furnished with bristles.

Distribution.—Four out of the five species live attached to Crustacea in the European and Eastern warmer temperate and tropical oceans; the fifth species was found attached to the dead spines of an Echinus, off New Guinea. It is probable that several more species will be hereafter discovered.

1. PæCILASMA KæMPFERI. Pl. II, Fig. 1.

P. valvis 5; carinæ basi truncatâ et cristatâ: scutorum dentibus internis umbonalibus fortibus: tergo-

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rum acumine basali truncato, margini occludenti pæne parallelo.

Valves 5; carina with a truncated and crested base; scuta with strong internal umbonal teeth; terga with the basal point truncated, almost parallel to the occludent margin.

Maxillæ with short thick spines in the notch under the two upper great spines; caudal appendages with scattered bristles on their summits, and along their whole outer margins.

Japan; attached in great numbers, to the upper and under sides of the Inachus Kæmpferi of De Haan, a slow-moving brachyourous crab, probably from deep water. British Museum.

General Appearance.—Capitulum rather compressed, narrow, and produced. Valves white, tinged with orange, smooth, moderately thin, occasionally with faint traces of striæ radiating from the umbones. Scuta, apex pointed, with a very slight ridge running to the umbo; basal margin equalling two thirds of the length of the terga, with an internal basal rim; on the under side of each valve, beneath the umbo, there is a strong tooth. Out of the numerous specimens, all excepting one had their scuta unequally convex, with their occludent margins unequally curved, that of the more convex valve at the umbo, curling beyond the medial line. The basal end of the carina is; likewise, slightly curved laterally, and always turns towards the more convex valve. This inequality, as Mr. Gray pointed out to me, depends on the position of the specimens; the flatter side lying close to the carapace of the crab. Terga, flat, oblong, nearly rectangular; occludent margin straight; basal angle, truncated, almost parallel to the occludent margin; in width, three of four times as wide as the carina. Carina, (fig. 1, a) short, narrow, slightly curved, upper part broadest, with the apex rounded, only just passing up between the basal broad ends of the terga; externally carinated, internally very slightly concave; basal end

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abruptly truncated, crested, not deeply imbedded in the membrane of the peduncle.

Peduncle, barely as long as the capitulum, apparently (for specimens dry and much shrunk) narrow, surrounded by rings or folds of thicker yellowish membrane, of which the upper ones retain moderately long spines; low down these rings become confluent; whole surface finely dotted, dots largest on the rings.

Mouth.—Labrum highly bullate in the upper part, with a row of teeth on the crest; mandibles with four teeth, the fourth close to the inferior apex, which is very little developed, sometimes making the fourth tooth appear simply bifid. Maxillæ with two large spines on the upper angle, beneath which there is a large depression, bearing one rather long and thick and four short and thick, spines; inferior upraised part with a double row of longer and thinner spines.

Cirri.—Posterior cirri with segments bearing five pairs of spines, of which the lowest pair is very minute; intermediate spines minute; spines of the dorsal tuft thin, of nearly equal size; segments not at all protuberant, elongated. First cirrus, standing far separated from the second (as in Scalpellum), with its nearly equal rami rather above half as long as those of the second cirrus. Second cirrus with anterior ramus not thicker, and scarcely more thickly clothed with spines, than the posterior ramus, but shorter than it by three or four segments; the spines not forming a very thick brush on the anterior ramus. Both rami of third cirrus with a longitudinal row of minute spines, parallel to the main pairs. Between the bases of the pedicels of the first pair of cirri, there are two closely approximate, conical flattened protuberances, like the single one to be described in Ibla.

Caudal Appendages, about one third of the length of the pedicel of the sixth cirrus, with some moderately long and strong spines at the end, and down the whole outer sides.

Ova, much pointed. Penis, hairy.

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Size.—Capitulum in largest specimens half an inch long.


P. valvis 5; carinæ basi truncatâ: scutis ovatis, margine basali perbrevi, dentibus parvis, internis, umbonalibus instructo: tergorum acumine basali perobliquè truncato.

Valves 5; carina with a truncated base; scuta oval, with the basal margin very short, furnished with small internal umbonal teeth; terga, with the basal point very obliquely truncated.

Maxillæ with fine spines in the notch under the three great upper spines; caudal appendages with scattered bristles on their summits, and along only the upper part of their outer margins.

Madeira; found by the Rev. R. T. Lowe, attached to the rare Homola Cuvierii, probably a deep-water crab. British Museum.

General Appearance.—This species so closely resembles P. Kæmpferi, that it is superfluous to describe it in detail; and I will indicate only the points of difference. When the valves have been well preserved, they are of fine pale orange colour, and hence the name above given, which was proposed by the Rev. R. T. Lowe.

Scuta, with the internal umbonal teeth small; basal internal marginal rim very prominent, furrowed within; basal margin short, (only equalling half the length of terga), owing to the great curvature of the lower part of the carino-tergal margin; hence, the outline of the scuta is almost pointed oval. I saw no appearance of inequality in the two sides.

Terga, rather smaller in proportion to the scuta, than in P. Kæmpferi, with the basal end very obliquely truncated, so as to appear at first simply pointed, not parallel to the occludent margin; apex considerably more pointed and produced than in the foregoing species.

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Carina, almost of equal narrowness throughout, barely concave within; lower end triangular, abruptly truncated, and not crested.

Primordial valves very plain, with the usual hexagonal structure: those of the terga, rounded at both ends, instead of being square, as in the mature calcified valves.

Peduncle short, narrow, not half as long as the capitulum; paved with minute equal beads, as in the genus Dichelaspis.

Mouth.—Mandibles with the fourth tooth very small; inferior angle rudimentary. Maxillæ, with three great upper spines, beneath which there is a deep notch bearing some delicate spines; inferior upraised part, as in P. Kæmpferi.

Cirri.—Rami of first cirrus hardly more than one third as long as the rami of the second cirrus, which latter rami are unequal in length by only two segments; the posterior ramus being the longer one.

Caudal Appendages, with only two or three lateral bristles, besides those on the summit.

Size.—Capitulum, three to four tenths of an inch long.

General Remarks.—This species has the closest general resemblance to P. Kæmpferi, and is evidently a representative of it. On close examination, however, almost every part differs slightly; the chief points being the narrowness of the basal margin of the scuta; the obliqueness of the truncated basal end of the terga and the sharpness of the upper end; the rudimentary state of the inferior angle of the mandibles; the character of the spines on the maxillæ; the proportional lengths of the cirri, and the fewness of the spines on the outer sides of the caudal appendages. The fact of Madeira having this Pæcilasma, a representative both in structure and habits of a Japan species, is interesting, inasmuch, as I am informed by Mr. Lowe, that some of the Madeira fishes are analogues of those of Japan.

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3. PæCILASMA CRASSA. Pl. II. Fig. 3.

ANATIFA CRASSA. J. E. Gray. Proc. Zoolog. Soc., 1848, p. 44, Annulosa, Tab. iii, figs. 5, 6.

P. valvis 5; carinæ termino basali in discum parcum infossum producto: scutis convexis, dentibus internis umbonalibus nullis: tergis pæne rudimentalibus, vix carinâ latioribus.

Valves 5; carina with the basal end produced into a small imbedded disc; scuta convex, without internal umbonal teeth; terga almost rudimentary, scarcely broader than the carina.

Spines on the segments of the posterior cirri arranged in single transverse rows.

Madeira; attached to the Homola Cuvierii, Rev. R. T. Lowe. British Museum*

General Appearance.—Capitulum highly bullate, or thick. Valves rather thick, opaque, either pale or dark flesh-red, smooth, yet rather plainly striated from the umbones. There are a few very minute spines on the membranous borders of the valves.

Scuta highly convex, broadly oval, apex broad rounded; basal margin narrow, much curved; no internal, umbonal teeth; basal internal rim strong, running up part of the occludent margin. A slightly prominent ridge, either rounded or angular, but in one specimen a narrow depressed fissure-like line, runs parallel to the occludent margin and ends near the apex in a slight notch; this fact is of interest in relation to the structure of the scuta in P. eburnea and P. fissa. The scuta are either equally or very unequally convex; in the latter case, the occludent margin of one valve is curled, so that its umbo is not quite medial.

* It is stated, in 'Zoolog. Proc.,' (1848, p. 44,) that this species was attached to a gorgonia, from Madeira; I cannot but suspect that there has been some confusion with the Oxynaspis celata from Madeira, which is thus attached.

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Terga, minute, almost rudimentary, scarcely broader than the carina, and half as long as the chord of its arc; carinal margin slightly curved; scutal margin straight, with a slight prominence fitting into a notch in the scuta; basal end bluntly pointed.

Carina, (fig. 3, a) rather shorter than the scuta, extending up only to the basal ends of the terga; moderately curved; apex moderately sharp; middle part broadest, externally carinated; internally not concave, with the inner lamina of shell, at the basal end, produced into a very small oblong disc or tooth, which is only as wide as the narrowest upper part of the valve. The exterior keel does not extend on to this disc, which is slightly constricted at its origin.

Peduncle very short, narrow, ringed, and apparently without spines.

Size.—Capitulum four tenths of an inch long.

The following parts of the animal are described from some small and not well preserved specimens from Madeira, which I owe to the kindness of Mr. Lowe.

Mouth.—Labrum highly bullate in the upper part, with large, inwardly pointed, unequal teeth. Mandibles, with four large, pointed, equal-sized teeth, with the inferior angle very narrow, acuminated like a single spine. Maxillæ, with three (?) large upper spines, of which the middle one is extremely strong and long, beneath which, there is a deep notch with a single strong spine, and with the whole inferior part square and much upraised, so as to stand on a level almost with the tips of the great upper spines.

Cirri in a miserable state of preservation; first cirrus short, second cirrus with rami unequal, and I suspect the anterior one the longest; some of the other cirri also have unequal rami. The segments of the posterior cirri are not protuberant, they have on their anterior faces a single transverse row of bristles: in the upper segments, some of the spines in each dorsal tuft (which is much spread out), are much thicker, though rather

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shorter than those on the anterior face. This peculiar structure is common to all five posterior cirri.

Caudal Appendages.—I can only say that they are spinose on their summits.

Affinities.—This species is allied to P. eburnea in the rudimentary condition of its terga; in the disc-shaped basal end of its carina; and in the presence in some specimens, of a fissure-like line on the scuta parallel to their occludent margins. Its affinity; however, is closer to P. fissa, as is more especially shown by the remarkable arrangement of the spines on the five posterior cirri.

4. PæCILASMA FISSA. Pl. II, Fig. 4.

P. valvis 7; scuto utroque è duobus juxtapositis segmentis formato; segmento altero intus dentalo: tergis brevibus, ter aut quater carinâ latioribus; carinæ termino basali in discum parvum angustum infossum producto.

Valves 7; each scutum being formed of two closely approximate segments; of which one is internally toothed: terga short, three or four times as wide as the carina: carina with the basal end produced into a small, narrow, imbedded disc.

Spines on the segments of the posterior cirri arranged in single transverse rows.

Philippine Archipelago; Island of Bohol; parasitic on a spinose crab, found under a stone at low water; single specimen, in Mus., Cuming.

General Appearance.—Capitulum gibbous, broadly oval, nearly a quarter of an inch long. Valves white, smooth, moderately thick, marked by the lines of growth. The occludent segments of the scuta, and nearly the whole of the terga, and the whole of the carina, enveloped in lemon-yellow membrane, tinged with orange, but the specimen had long been kept dry.

Scuta formed of two, apparently always separate, segments, closely united, so that externally their separa-

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tion is hardly visible, and does not allow of movement; the fissure thus formed runs almost in the line connecting the umbo and apex, (where in most species a ridge extends,) but a little on the carinal side of it. The occludent segment is narrowly bow-shaped, pointed at both ends, with the upper end projecting slightly beyond the apex of the lateral segment, and with the occludent margin regularly curved from end to end. The lateral segment is large, of an oval shape, with a narrow strip cut off on one side. Primordial valves very plain at the umbones of the lateral segments, but none are visible on the occludent segments; and this makes me believe that these two pieces are normally parts of a single valve; having only one specimen of P. fissa, I was not able to make out quite certainly whether the two segments are continuously united at their umbones by a non-calcified portion of valve, as is certainly the case with Dischelaspis. The basal margin of the lateral segment is narrow, inflected, and blends with the carino-tergal margin; it has an internal, prominent, basal rim, and towards the occludent margin a large, prominent, internal tooth. This internal basal rim is not parallel to the outer basal margin, but rises to a point a little way up the occludent margin, in the same manner as in P. eburnea, but in a lesser degree; in this latter species the peduncle is internally almost cut off by the large disc of its carina; here, on the other hand, it is internally almost cut off by these rims and the two large teeth of the lateral segments of the scuta.

Terga sub-triangular, short, nearly half as broad as long; three or four times as wide as the carina, and rather wider than the occludent segment of the scuta; occludent margin single, arched; carinal margin slightly arched; basal angle bluntly pointed.

Carina vary narrow, much arched, running up just between the basal ends of the terga; exterior ridge enveloped in membrane; heel blunt, prominent; internally, not concave, even slightly convex, produced at the lower

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end into a very narrow, short, imbedded disc, (or rather tooth,) which is itself a little curved downwards and blunt at the end.

Peduncle very narrow, about half as long as the capitulum; yellow, finely beaded, plainly ringed, without spines.

Mouth.—Labrum, with a row of minute teeth; palpi narrow. Mandibles with all the lower part narrow; of the four teeth, the second and third are narrow, the fourth is pectinated and placed very close to the inferior angle, which is produced into a long thin tooth. Maxillæ unknown.

Cirri.—First pair lost. The arrangement of the spines on all is most abnormal, Pl. X, fig. 29: dorsal tuft long, arranged in a transverse line and seated in a deep notch; in the sixth cirrus, the spines on the lower segments are fine, those on the upper segments are thick and claw-like, mingled with some fine spines; in the four anterior cirri the spines of the dorsal tufts are even thicker and more claw-like. On the anterior faces, also, of all the segments the spines form a single row; they are shorter than those composing the dorsal tuft; hence the spines on each segment are arranged in a circle, interrupted widely on the two sides: this arrangement is common to all five posterior cirri. Second cirrus, with the anterior ramus one third longer and thinner than the posterior ramus (this is the reverse of the usual arrangement) this longer ramus equals in length the sixth cirrus. Third cirrus, with the anterior ramus considerably longer than the posterior ramus; in the three posterior pair of cirri, also, the anterior rami are a little longer than the posterior: except in length, there is little difference of any kind between the five posterior pair of cirri. Pedicels of the cirri long; rami rather short; segments elongated, not protuberant.

Caudal Appendages nearly as long as the pedicels of the sixth cirrus, thickly clothed with very fine bristles, like a camel's-hair pencil brush.

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Affinities.—In the structure of the carina, and more especially of the scuta, there is a strong affinity between the present and following species; for we shall immediately see that in P. eburnea there is evidence of the scuta being composed of two segments fused together; and the larger segment is furnished with an internal oblique, strong, basal rim. To this same species there is an evident affinity in the form of the mandibles and of the caudal appendages, and in the anterior rami of the cirri being longer than the posterior rami. Notwithstanding these points of affinity, I consider that P. fissa is more closely related in its whole organisation, as more particularly shown in the arrangement of the spines on the cirri and in the presence of terga, to P. crassa than to P. Eburnea. Although in Dichelaspis, the scuta are invariably composed of two almost separate segments, yet P. fissa shows no special affinity to this genus.


TRILASMIS EBURNEA. Hinds. Voyage of Sulphur, 1844, vol. i, Mollusca, Pl. Xi, fig. 5.

P. valvis 3; scutis acuminatis, ovatis; ad pedunculum pæne transversè spectantibus; dentibus internis umbonalibus fortibus: tergis nullis: carinæ termino basali in discum amplum oblongum infossum producto.

Valves 3; scuta pointed, oval, placed almost transversely to the peduncle; internal umbonal teeth strong: terga absent: carina with the basal end produced into a large, oblong, imbedded disc.

Spines on the upper segments of the posterior cirri, arranged in three or four approximate longitudinal rows, making small brushes.

Habitat.—New Guinea, attached to the spines of a dead Echinus. Brit. Mus., and Cuming.

General Appearance.—Capitulum flat, pear-shaped,

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placed almost transversely to the peduncle. Valves white, smooth, moderately thick.

Scuta: the basal margin, as seen externally, is narrow, and can hardly be separated from the carinal margin; but an internal basal rim, (fig. 5, b) (along which the imbedded disc of the carina runs,) shows where, in the other species, the basal and carinal margins are separated. This basal internal rim is not parallel to the external basal margin, but runs upwards to the occludent margin, leaving beneath it a large triangular space, to which the membrane of the peduncle is attached; and this makes it appear as if the rostral umbones of these valves had grown downwards; but, judging from the allied species, P. fissa, I have no doubt that the primordial valves really lie on the umbones, and that the growth has been in the usual direction, that is, exclusively upwards. The occludent margin is curved, and blends by a regular sweep into the carinal margin, so that there is no acute upper angle. A distinct line can be seen, as if two calcareous valves had been united, running from the umbo to the upper end of the valve, thus in appearance separating a slip of the occludent margin; internally this appearance is more conspicuous; this structure is important in relation to that of P. fissa. The pointed umbones are divergent, and internally under each, there is a large tooth. The two valves are equally convex.

Terga, entirely absent.

The Carina (Tab. II, fig. 5, a, c), including the disc, is three fourths as long as the scuta; it is placed almost transversely to the longitudinal axis of the peduncle; it is narrow and internally convex; the imbedded disc is very large, forming a continuous curve with the upper part of the carina; this disc runs along the internal basal rim of the scuta, and hence almost separates, internally, the peduncle from the capitulum; it equals one fourth of the total length of the valve, and is thrice as wide as the upper part; it is oval, externally marked by a central


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line, and with a slight notch at the end, giving a divided appearance to the whole, and indicating how easily a fork might be formed from it. The carina is thick, measured from the inner convex to the exterior surface, which is carinated; heel prominent.

Peduncle, narrow, very short, not nearly so long as the capitulum.

Mouth.—Labrum considerably bullate, with the lower part much produced towards the adductor muscle; crest with small bead-like teeth; palpi small, pointed; mandibles, with the first tooth standing rather distant from the second; inferior angle spine-like and bifid; maxillæ (Pl. X, fig. 15), with two considerable spines (only one is shown in the Plate) beneath the upper large pair; the inferior upraised part bears seven or eight pair of spines, and its edge is not quite straight; close to the main notch, lying under the four upper spines, there are two minute notches, with the interspace bearing a tuft of fine spines and a pair of larger ones.

Cirri.—The rami in all are rather unequal in length, the anterior rami being rather the longest; the anterior rami of the second and third cirri are not thicker than the posterior rami. The segments in the three posterior cirri are not protuberant; the upper segments bear three or four pair of spines, with some minute intermediate ones, and with the lateral marginal spines unusually large and long, so as to form, with the ordinary pairs, a third or fourth longitudinal row; hence a small brush is formed on each segment. The dorsal tuft is large and wide, so as to contain even fourteen spines, of which some are as long as those in front. In the lower segments of these same posterior cirri, the lateral marginal spines are not so much developed (nor is the dorsal tuft), and hence the segments can hardly be said to be brush-like. The first cirrus is placed rather distant from the second pair. The second and third cirri differ from the three posterior pair, only in the bristles being slightly more numerous, and in the dorsal tufts being more spread out.

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Caudal Appendages about half the length of the lower segments of the pedicels on the sixth cirrus; truncated and rounded at their ends; thickly clothed with long excessively fine bristles, so as to resemble camel-hair pencils.

The Stomach, I believe, is destitute of cæca; in it was a small crustacean.

General Remarks.—I was at first unwilling to sacrifice Mr. Hind's genus, Trilasmis, which is so neatly characterised by its three valves; moreover, the present species does differ, in some slight respects, from the other species of Pælasma; but under the head of P. fissa I have shown how that species, P. crassa and P. eburnea are tied together. The absence of terga, which are rudimentary in P. crassa, (and we shall hereafter see, in Conchoderma, how worthless a character their entire absence is,) and the arrangement of the spines in the upper segments of the posterior cirri, are the only characters which could be used for a generic separation.

Darwin, A monograph on the sub-class cirripedia, with figures of all the species. The lepadidae, or, pedunculated cirripedes. 1851.

-Front matter; preface; introduction (pp. -7)
-Family—lepadidæ; capitulum; Genus—lepas; pæcilasma. nov. genus
(pp. 8-115)
-Genus—dichelaspis; oxynaspis. gen. nov.; genus—conchoderma; Genus—alepas; anelasma. gen. nov.; Genus—ibla
(pp. 115- 214)
-Genus—scalpellum; [sub-carinâ nullâ.] 1. scalpellum vulgare; 2. scalpellum ornatum; 3. scalpellum rutilum; [sub-carinâ presente.] 4. scalpellum rostratum; 5. scalpellum peronii; 6. scalpellum villosum
(pp. 215-281)
-Summary on the nature and relations of the males and complemental males in ibla and scalpellum; Genus—pollicipes; Genus—lithotrya; Species mihi non satis notæ, aut dubiæ (pp. 281-375)
-Explanation of the plates
(pp. 377-393)
-Index (pp. 395-400)


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