A Monograph on the Sub-class Cirripedia (Volume 1 of 2) - The Lepadidae; or, Pedunculated Cirripedes
by Charles Darwin
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My duty, in acknowledging the great obligations under which I lie to many naturalists, affords me most sincere pleasure. I had originally intended to have described only a single abnormal Cirripede, from the shores of South America, and was led, for the sake of comparison, to examine the internal parts of as many genera as I could procure. Under these circumstances, Mr. J. E. Gray, in the most disinterested manner, suggested to me making a Monograph on the entire class, although he himself had already collected materials for this same object. Furthermore, Mr. Gray most kindly gave me his strong support, when I applied to the Trustees of the British Museum for the use of the public collection; and I here most respectfully beg to offer my grateful acknowledgments to the Trustees, for their most liberal and unfettered permission of examining, and when necessary, disarticulating the specimens in the magnificent collection of Cirripedes, commenced by Dr. Leach, and steadily added to, during many years, by Mr. Gray. Considering the difficulty in determining the species in this class, had it not been for this most liberal permission by the Trustees, the public collection would have been of no use to me, or to any other naturalist, in systematically classifying the Cirripedes.

Previously to Mr. Gray suggesting to me the present Monograph, Mr. Stutchbury, of Bristol, had offered to intrust to me his truly beautiful collection, the fruit of many years' labour. At that time I refused this most generous offer, intending to confine myself to anatomical observations; but I have since accepted it, and still have the entire splendid collection for my free use. Mr. Stutchbury, with unwearied kindness, further supplied me with fresh specimens for dissection, and with much valuable information. At about the same period, Mr. Cuming strongly urged me to take up the subject, and his advice had more weight with me than that of almost any other person. He placed his whole magnificent collection at my disposal, and urged me to treat it as if it were my own: whenever I told him that I thought it necessary, he permitted me to open unique specimens of great value, and dissect the included animal. I shall always feel deeply honoured by the confidence reposed in me by Mr. Cuming and Mr. Stutchbury.

I lie under obligations to so many naturalists, that I am, in truth, at a loss how to express my gratitude. Mr. Peach, over and over again, sent me fresh specimens of several species, and more especially of Scalpellum vulgare, which were of invaluable assistance to me in making out the singular sexual relations in that species. Mr. Peach, furthermore, made for me observations on several living individuals. Mr. W. Thompson, the distinguished Natural Historian of Ireland, has sent me the finest collection of British species, and their varieties, which I have seen, together with many very valuable MS. observations, and the results of experiments. Prof. Owen procured for me the loan of some very interesting specimens in the College of Surgeons, and has always given me his invaluable advice and opinion, when consulted by me. Professor E. Forbes has been, as usual, most kind in obtaining for me specimens and information of all kinds. To the Rev. R. T. Lowe I am indebted for his particularly interesting collection of Cirripedes from the Island of Madeira—a collection offering a singular proof what treasures skill and industry can discover in the most confined locality. The well-known conchologist, Mr. J. G. Jeffreys, has sent for my examination a very fine collection of British specimens, together with a copious MS. list of synonyms, with the authorities quoted. To the kindness of Messrs. M^c Andrew, Lovell Reeve, G. Busk, G. B. Sowerby, Sen., D. Sharpe, Bowerbank, Hancock, Adam White, Dr. Baird, Sir John Richardson, and several other gentlemen, I am greatly indebted for specimens and information: to Mr. Hancock I am further indebted for several long and interesting letters on the burrowing of Cirripedes.

Nor are my obligations confined to British naturalists. Dr. Aug. Gould, of Boston, has most kindly transmitted to me some very interesting specimens; as has Prof. Agassiz other specimens collected by himself in the Southern States. To Mr. J. D. Dana, I am much indebted for several long letters, containing original and valuable information on points connected with the anatomy of the Cirripedia. To Mr. Conrad I am likewise indebted for information and assistance. Both the celebrated Professors, Milne Edwards and Mueller, have lent me, from the great public collections under their charge, specimens which I should not otherwise have seen. To Professor W. Dunker, of Cassel, I am indebted for the examination of his whole collection. I have, in a former publication, expressed my thanks to Professor Steenstrup, but I must be permitted here to repeat them, for a truly valuable present of a specimen of the Anelasma squalicola of this work. I will conclude my thanks to all the above British and foreign naturalists, by stating my firm conviction, that if a person wants to ascertain how much true kindness exists amongst the disciples of Natural History, he should undertake, as I have done, a Monograph on some tribe of animals, and let his wish for assistance be generally known.

Had it not been for the Ray Society, I know not how the present volume could have been published; and therefore I beg to return my most sincere thanks to the Council of this distinguished Institution. To Mr. G. B. Sowerby, Junr., I am under obligations for the great care he has taken in making preparatory drawings, and in subsequently engraving them. I believe naturalists will find that the ten plates here given are faithful delineations of nature.

In Monographs, it is the usual and excellent custom to give a history of the subject, but this has been so fully done by Burmeister, in his 'Beitraege zur Naturgeschichte der Rankenfuesser,' and by M. G. Martin St. Ange, in his 'Memoire sur l'Organisation des Cirripedes,' that it would be superfluous here to repeat the same list of authors. I will only add, that since the date, 1834, of the above works, the only important papers with which I am acquainted, are, 1st. Dr. Coldstream 'On the Structure of the Shell in Sessile Cirripedes,' in the 'Encyclopaedia of Anatomy and Physiology;' 2d. Dr. Loven 'On the Alepas squalicola,' ('Ofversigt of Kongl. Vetens.,' &c. Stockholm, 1844, p. 192,) giving a short but excellent account of this abnormal Cirripede; 3d. Professor Leidy's very interesting discovery, ('Proceedings of the Academy of Natural Sciences,' Philadelphia, vol. iv, No. I, Jan. 1848,) of eyes in a mature Balanus; 4th. Mr. A. Hancock's Memoir, ('Annals of Natural History, 2d series, Nov. 1849,) on his Alcippe lampas, the type of a new order of Cirripedes; 5th. Mr. Goodsir's Paper, ('Edinburgh New Philosoph. Journal,' July 1843,) on the Larvae in the First Stage of Development in Balanus; 6th. Mr. C. Spence Bate's valuable Paper on the same subject, lately published, (Oct. 1851,) in the 'Annals of Natural History;' and lastly, M. Reinhardt has described, in the 'Copenhagen Journal of Natural History, Jan. 1851,' the Lithotrya Nicobarica, and has discussed its powers of burrowing into rocks.

I have given the specific or diagnostic characters, deduced from the external parts alone, in both Latin and English. As I found, during the progress of this work, that a similarly abbreviated character of the softer internal parts, was very useful in discriminating the species, I have inserted it after the ordinary specific character.

In those cases in which a genus includes only a single species, I have followed the practice of some botanists, and given only the generic character, believing it to be impossible, before a second species is discovered, to know which characters will prove of specific, in contradistinction to generic, value.

In accordance with the Rules of the British Association, I have faithfully endeavoured to give to each species the first name attached to it, subsequently to the introduction of the binomial system, in 1758, in the tenth edition[1] of the 'Systema Naturae.' In accordance with the Rules, I have rejected all names before this date, and all MS. names. In one single instance, for reasons fully assigned in the proper place, I have broken through the great law of priority. I have given much fewer synonyms than is usual in conchological works; this partly arises from my conviction that giving references to works, in which there is not any original matter, or in which the Plates are not of a high order of excellence, is absolutely injurious to the progress of natural history, and partly, from the impossibility of feeling certain to which species the short descriptions given in most works are applicable;—thus, to take the commonest species, the Lepas anatifera, I have not found a single description (with the exception of the anatomical description by M. Martin St. Ange) by which this species can be certainly discriminated from the almost equally common Lepas Hillii. I have, however, been fortunate in having been permitted to examine a considerable number of authentically named specimens, (to which I have attached the sign (!) used by botanists,) so that several of my synonyms are certainly correct.

[1] In the Rules published by the British Association, the 12th edition, (1766,) is specified, but I am informed by Mr. Strickland that this is an error, and that the binomial method was followed in the 10th edition.

The Lepadidae, or pedunculated Cirripedes, have been neglected under a systematic point of view, to a degree which I cannot quite understand: no doubt they are subject to considerable variation, and as long as the internal surfaces of the valves and all the organs of the animal's body, are passed over as unimportant, there will occasionally be some difficulty in the identification of the several forms, and still more in settling the limits of the variability of the species. But I suspect the pedunculated Cirripedes have, in fact, been neglected owing to their close affinity, and the consequent necessity of their being included in the same Work with the Sessile Cirripedes; for these latter will ever present, I am fully convinced, insuperable difficulties in their identification by external characters alone.

I will here only further remark, that in the Introduction I have given my reasons for assigning distinct names to the several Valves, and to some parts of the included animal's body; and that in the Introductory Remarks, under the general description of the Lepadidae, I have given an abstract of my Anatomical Observations.



12, twenty lines from bottom, for "hinder pair of true thoracic limbs," read "pair of true thoracic limbs."

42, 43. I should have added, that the number of the segments in the cirri increases with the age of the specimen; but that the relative numbers in the different cirri keep, as far as I have seen, nearly constant; hence the numbers are often given in the descriptions.

99 et passim, for Paecilasma, read Poecilasma.

156. In a foot-note, I have alluded to a new genus of sessile Cirripedes, under the name of Siphonicella, I now find that this species has been called, by Professor Steenstrup, Xenobalanus globicipitis.





I should have been enabled to have made this Volume more complete, had I deferred its publication until I had finished my examination of all the other known Cirripedes; but my work would thus have been rendered inconveniently large. Until this examination is completed, it will be more prudent not to discuss, in detail, the position of the Lepadidae amongst the Cirripedia, or of these latter in the great class of Crustacea, to which they now, by almost universal consent, have been assigned. I may, however, remark that I believe the Cirripedia do not approach, by a single character, any animal beyond the confines of the Crustacea: where such an approach has been imagined, it has been founded on erroneous observations; for instance, the closed tube within the stomach, described by M. Martin St. Ange (to whose excellent paper I am greatly indebted), as indicating an affinity to the Annelides, is, I am convinced, nothing but a strong epithelial lining, which I have often seen ejected with the excrement. Again, a most distinguished author has stated that the Cirripedia differ from the Crustacea:—1st. In having "a calcareous shell and true mantle;" but there is no essential difference, as shown by Burmeister, in the shells in these two classes; and Cirripedes certainly have no more claim to a mantle than have the bivalve entomostraca. 2d. "In the sexes joined in one individual;" but this, as we shall see, is not constant, nor of very much weight, even if constant. 3d. "In the body not being ringed;" but if the outer integument of the thorax of any Cirripede be well cleaned, it will be seen, (as was long ago shown by Martin St. Ange), to be most distinctly articulated. 4th. "In having salivary glands;" but these glands are, in truth, the ovaria. 5th. "In the liver being formed on the molluscous type;" I do not think this is the case, but I do not quite understand the point in question. 6th. "In not having a head or organs of sense;" this is singularly erroneous: Professor Leidy has shown the existence of eyes in the mature Cirripede; the antennae, though preserved, certainly become functionless soon after the last metamorphosis; but there exist other organs of sense, which I believe serve for smelling and hearing: and lastly, so far from there being no head, the whole of the Cirripede externally visible, consists exclusively of the three anterior segments of the head.

The sub-class, Cirripedia, can be divided into three Orders; the first of which, mainly characterised by having six pair of thoracic cirri, includes all common Cirripedes: these latter may be divided into three families,—the Lepadidae, or pedunculated Cirripedes, the subject of the present memoir; the Verrucidae containing the single genus Verruca or Clisia; and, lastly, the Balanidae, which consist of two very distinct sub-families, the Balaninae and Chthamalinae. Of the other two Orders above alluded to, one will, I believe, contain the remarkable burrowing genus Alcippe, lately described by Mr. Hancock, and a second burrowing genus, or rather family, obtained by me on the coast of South America. The third Order is highly singular, and differs as much from all other Cirripedes as does a Lernaea from other crustaceans; it has a suctorial mouth, but is destitute of an anus; it has not any limbs, and is as plainly articulated as the larva of a fly; it is entirely naked, without valves, carapace, or capitulum, and is attached to the Cirripede, in the sack of which it is parasitic, by two distinct threads, terminating in the usual larval, prehensile antennae. I intend to call this Cirripede, Proteolepas. I mention it here for the sake of calling attention to any parasite at all answering to this description.


Although the present volume is strictly systematic, I will, under the general description of the Lepadidae, give a very brief abstract of some of the most interesting points in their internal anatomy, and in the metamorphoses of the whole class, which I hope hereafter to treat, with the necessary illustrations, in detail. I enter on the subject of the metamorphoses the more readily, as by this means alone can the homologies of the different parts be clearly understood.

On the Names given to the different parts of Cirripedes.

I have unwillingly found it indispensable to give names to several valves, and to some few of the softer parts of Cirripedes. The accompanying figure of an imaginary Scalpellum includes every valve; the two most important valves of Lepas are also given, in which the direction of the lines of growth and general shape differ from those of Scalpellum as much as they do in any genus. The names which I have imposed will, I hope, be thus acquired without much difficulty.

Whoever will refer to the published descriptions of recent and fossil Cirripedia, will find the utmost confusion in the existing nomenclature: thus, the valve named in the woodcut the Scutum, has been designated by various well-known naturalists as the "ventral," the "anterior," the "inferior," the "ante-lateral," and the "latero-inferior" valve; the first two of these titles have, moreover, been applied to the rostrum or rostral valve of sessile Cirripedes. The Tergum has been called the "dorsal," the "posterior," the "superior," the "central," the "terminal," the "postero-lateral," and the "latero-superior" valve. The Carina has received the first two of these identical epithets, viz. the "dorsal" and the "posterior;" and likewise has been called the "keel-valve." The confusion, however, becomes far worse, when any individual valve is described, for the very same margin which is anterior or inferior in the eyes of one author, is the posterior or superior in those of another; it has often happened to me that I have been quite unable even to conjecture to which margin or part of a valve an author was referring. Moreover, the length of these double titles is inconvenient. Hence, as I have to describe all the recent and fossil species, I trust I may be thought justified in giving short names to each of the more important valves, these being common to the pedunculated and sessile Cirripedes.

The part supported by the peduncle, and which is generally, though not always, protected by valves, I have designated the Capitulum.

The title of Peduncle, which is either naked or squamiferous, requires no explanation; the scales on it, and the lower valves of the capitulum, are arranged in whorls, which, in the Latin specific descriptions, I have called by the botanical term of verticillus.

I have applied the term Scutum to the most important and persistent of the valves, and which can generally be recognised by the hollow giving attachment to the adductor scutorum muscle, from the resemblance which the two valves taken together bear to a shield, and from their office of protecting the front side of the body. From the protection afforded by the two Terga to the dorso-lateral surface of the animal, these valves have been thus called. The term Carina[2] is a mere translation of the name already used by some authors, of Keel-Valve.

[2] In the Carina of Fossil Species of Scalpellum, I have found it necessary to distinguish different parts, viz., A, the tectum, of which half is seen; B, the parietes; and C, the intra-parietes.

The Rostrum has been so called from its relative position to the carina or keel. There is often a Sub-carina and a Sub-rostrum.

The remaining valves, when present, have been called Latera; there is always one large upper one inserted between the lower halves of the scuta and terga, and this I have named the Upper Latus or Latera; the other latera in Pollicipes are numerous, and require no special names; in Scalpellum, where there are at most only three pair beneath the Upper Latera, it is convenient to speak of them (vide Woodcut, I,) as the Carinal, Infra-median, and Rostral Latera.

As each valve often requires (especially amongst the fossil species) a distinct description, I have found it indispensable to give names to each margin. These have mostly been taken from the name of the adjoining valve, (see fig. I.) In Lepas, Pollicipes, &c., the margin of the scutum adjoining the tergum and upper latus, is not divided (fig. II) into two distinct lines, as it is in Scalpellum, and is therefore called the Tergo-lateral margin. In Scalpellum (fig. I) these two margins are separately named Tergal and Lateral. The angle formed by the meeting of the basal and lateral or tergo-lateral margins, I call the Baso-lateral angle; that formed by the basal and occludent margins, I call, from its closeness to the Rostrum, the Rostral angle. In Pollicipes the carinal margin of the tergum can be divided into an upper and lower carinal margin; of this there is only a trace (fig. I) in Scalpellum.

That margin in the scuta and terga which opens and shuts for the exsertion and retraction of the cirri, I have called the Occludent margin. In the terga of Lepas (fig. III) and some other genera, the occludent margin is highly protuberant and arched, or even formed of two distinct sides.

Occasionally, I have referred to what I have called the primordial valves: these are not calcified; they are formed at the first exuviation, when the larval integuments are shed: in mature Cirripedes they are always seated, when not worn away, on the umbones of the valves.

The membrane connecting the valves, and forming the peduncle, and sometimes in a harder condition replacing the valves, I have often found it convenient to designate by its proper chemical name of Chitine, instead of by horny, or other such equivalents. When this membrane at any articulation sends in rigid projections or crests, for the attachment of muscles or any other purpose, I call them, after Audouin, apodemes. For the underlying true skin, I use the term corium.

The animal's body is included within the capitulum, within what I call the sack (see Pl. IV, figs. 2 and 8' a, and Pl. IX, fig. 4). The body consists of the thorax supporting the cirri, and of an especial enlargement, or downward prolongation of the thorax, which includes the stomach, and which I have called the prosoma. (Pl. IX, fig. 4 n). The cirri are composed of two arms or rami, supported on a common segment or support, which I call the pedicel. The caudal appendages are two little projections, either uni-or multi-articulate (Pl. IV, fig. 8' a), on each side of the anus, and just above the long proboscis-like penis. On the thorax and prosoma, or on the pedicels of the cirri, there are in several genera, long, thin, tapering filaments, which have generally been supposed to serve as branchiae; these I call simply filaments, or filamentary appendages (Pl. IX, fig. 4 g-l). The mouth (fig. 4 b) is prominent, and consists of palpi soldered to the labrum; mandibles, maxillae, and outer maxillae, these latter serve as an under lip; to these several organs I sometimes apply the title used by Entomologists, of "trophi." Beneath the outer maxillae, there are either two simple orifices or tubular projections; these, I believe, serve as organs of smell, and have hence called them the olfactory orifices. Within the sack, there are often two sheets of ova (Pl. IV, fig. 2 b), these I call (after Steenstrup, and other authors) the ovigerous Lamellae; they are united to two little folds of skin (Pl. IV, fig. 2 f), which I call the ovigerous Fraena.

From the peculiar curved position which the animal's body occupies within the capitulum, I have found it far more convenient (not to mention the confusion of nomenclature already existing) to apply the term Rostral instead of ventral, and Carinal instead of dorsal, to almost all the external and internal parts of the animal. Cirripedes have generally been figured with their surfaces of attachment downwards, hence I speak of the lower or Basal margins and angles, and of those pointing in an opposite direction as the Upper; strictly speaking, as we shall presently see, the exact centre of the usually broad and flat surface of attachment is the anterior end of the animal, and the upper tips of the Terga, the posterior end of that part of the animal which is externally visible; but in some cases, for instance in Coronula, where the base is deeply concave, and where the width of the shell far exceeds the depth, it seemed almost ridiculous to call this, the anterior extremity; as likewise does it in Balanus to call the united tips of the Terga, lying deeply within the shell, the most posterior point of the animal, as seen externally.

I have followed the example of Botanists, and added the interjection [!] to synonyms, when I have seen an authentic specimen bearing the name in question.

Every locality, under each species, is given from specimens ticketed in a manner and under circumstances appearing to me worthy of full confidence,—the specific determination being in each case made by myself.



Cirripedia pedunculo flexili, musculis instructo: scutis[3] musculo adductore solummodo instructis: valvis caeteris, siquae adsunt, in annulum immobilem haud conjunctis.

Cirripedia having a peduncle, flexible, and provided with muscles. Scuta[3] 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; antennae, ib.; eyes, p. 16; mouth, p. 17; thorax and limbs, p. 18; abdomen, p. 19; viscera, ib.; immature cirripede, p. 20; homologies of parts, p. 25.

Description of mature Lepadidae, 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 lamellae, p. 58; ovigerous fraena, ib.; exuviation, p. 61; rate of growth, ib.; size, ib.; affinities of family, p. 64; range and habitats, p. 65; geological history, p. 66.

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

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 larvae in the last stage of development in Balanus, much has been done on this subject: this same author subsequently published[4] in the 'Philosophical Transactions,' an account of the larvae 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 difference in the two families, instead of to the stage of development. Burmeister[5] first showed, and the discovery is an important one, that in Lepas the larvae pass through two totally different stages. This has subsequently been proved by implication to be the case in Balanus, by Goodsir,[6] 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 larvae of five species in this same early stage, and has most kindly permitted me to quote from his unpublished paper[7]. I am enabled to confirm and generalise these observations, in all the Cirripedes in the Order containing the Balanidae and Lepadidae.

[4] Philosophical Transactions, 1835, p. 355, Pl. vi.

[5] Beitraege zur Naturgeschichte der Rankenfuesser, 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.

[6] Edinburgh New Philosophical Journal, July 1843, Pls. iii and iv.

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

The ova, and consequently the larvae of the Lepadidae, 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 larvae 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 antennae of the larvae in the last stage; that I have not the least doubt, that these horns are the cases in which antennae are in process of formation. Posteriorly 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 antennae (without a case), directed forwards: hence, before the first moult in Scalpellum, we have two pair of antennae in process of formation. Anteriorly to the bases of these smaller antennae 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 larvae, 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 larvae 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 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 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 larvae, like the mandibles and maxillae 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 antennae within the lateral horns, though standing more in front than the 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,[8] adhering to sea-weed in the midst of other larvae 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 antennae 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 antennae, 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 antennae, the eggs have two great projecting horns including 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.

[8] Beitraege zur Naturgeschichte der Rankenfuesser, s. 16, Tab. i, figs. 3, 4.

Larva, Last Stage.—My chief examination has been directed, at this stage of development, to the larvae of Lepas australis, which are of unusual size, namely, from .065 to even almost .1 of an inch in length; I examined, however, the larvae 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 antennae, preserved in the bases of all mature Cirripedes, and which I have carefully inspected in almost every genus. The larvae 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 antennae 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.

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.

Antennae.—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;[9] they can (at least in Ibla) 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 antennae, 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.[10] 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 antennae up to the discs.

[9] Mr. J. D. Dana, who has examined these organs in the larvae of Lepas, informs me in a letter, that in his opinion they "correspond with the inferior antennae, the superior being wanting, as in most Daphnidae." 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 antennae 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.

[10] Rev. B. L. King. Annual Report of B. Institution of Cornwall, 1848, p. 55.

Eyes.—Close behind the basal articulations of the antennae, the sternal surface consists of two approximate, elongated, narrow, flat pieces, or segments. These Burmeister considers as the basal segments of the antennae: 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, = deg.UU deg., 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 striae may be found attached to the knobbed tips of the two middle arms of the two = deg.UU deg., and running up to the antero-dorsal surface of the carapace, where they are attached; other muscles (without transverse striae) 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 antennae 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. 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 this, its last stage, cannot eat; it may be called a locomotive Pupa;[11] its whole organisation is apparently adapted for the one great end of finding a proper site for its attachment and final metamorphosis.

[11] 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 larvae, when, under that denomination, have hitherto been comprised animals capable of nourishing themselves?"

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 by 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 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 caeca 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 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 antennae 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 caeca of the stomach to which they cohere, but in young specimens, they extend some way beyond the caeca, 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 displaying 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 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. The antennae 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, 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 antennae, 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 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 caeca: 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 branchiae, at the base of the first pair of cirri. Nor could I perceive a trace of the testes or vesiculae 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 caeca 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 caeca, 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-caeca, 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 antennae 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 = deg.UU deg.=-like, sternal apodemes: these together with the eyes stretch transversely across, and internally far up into, 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 larvae 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 antennae, 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 antennae, and in an earlier stage two pair of antennae, 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 Lepadidae is likewise thus formed. The peduncle being filled by the branching ovarian tubes is no objection to this view, for I am informed on the high authority of Mr. J. D. Dana,[12] that this is the case with the cephalo-thorax in some true Crustaceans, for instance, in Sapphirina. To proceed, the mouth, formed of mandibles, maxillae, and outer maxillae, 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 maxillae 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 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.

[12] 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 antennae 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 ueber das vorkommen des Hermaphroditismus in der Natur,' ch. v, have considered the peduncle as a pair of fused legs. Loven 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 reliquae partes, ut in Lepadibus semper, in pedunculum mutatae et involucrum," &c.; his involucrum is the same as the capitulum of this work.

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 Lepadidae, 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 antennae were always embedded in the centre of the surface of attachment, I did not perceive, that this was the anterior 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 antennae 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 Lepadidae. 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 occasionally exceed 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, occasionally 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, Paecilasma, 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 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,[13] and is composed of numerous fine laminae. After the valves have been placed in acid, a residue, very different in bulk in different genera, is left, also composed of successive laminae 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 colour, and becomes much harder and more persistent; and as the whole valve is formed of component laminae thus edged (the once continuous laminae 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: in Scalpellum and Pollicipes it usually covers the whole valves. It appears to me that the laminae 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.

[13] 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.

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:[14] 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.

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

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 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 Lithotrya we have a row of calcareous discs or an irregular, basal cup, formed in the same 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 striae, 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. 6 b), from the surface of attachment, quite close to the end, but not at the actual apex, the larval prehensile antennae 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 antennae to the anterior or lower ends of the two gut-formed 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 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 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 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, continuous 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 organised 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 antennae; and it thus continues to do throughout life in most or all of the species of Lepas, Conchoderma, Dichelaspis and Ibla. In the first two of these genera, the cement escapes from the borders of the lower side of the disc or penultimate segment of the antennae, 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 antennae 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 antennae, in the shape of one tube, within another tube of considerable diameter and length. In Scalpellum vulgare, and probably in some of the other species, which live attached to corallines, the cement soon ceases to debouch from the antennae, 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 fascicularis 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 antennae, 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 antennae 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 Sub-class. 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 body[15] in order to fasten it to a foreign support. But on no other view can the 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.

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