Transcriber’s Note

Obvious misprints have been fixed. Archaic and unusual words, spellings and styling have been maintained. Details of the changes are in the [Detailed Transcriber’s Notes] at the end of the book. Hovering over a Greek word will display its transliteration.

This book was published in two volumes, of which this is the second. The first volume was released as Project Gutenberg ebook #31558, available at [http://www.gutenberg.org/ebooks/31558]. Links to the first volume have not been created in this volume.

Transcriber Added

Table of Contents.

List of Illustrations.

THE

RAY SOCIETY.

INSTITUTED MDCCCXLIV.

LONDON.

MDCCCLIV.

A MONOGRAPH
ON THE SUB-CLASS
CIRRIPEDIA,
WITH
FIGURES OF ALL THE SPECIES.

BY

CHARLES DARWIN, F.R.S., F.G.S.

THE BALANIDÆ,

(OR SESSILE CIRRIPEDES);

THE VERRUCIDÆ,

ETC., ETC., ETC.

LONDON:
PRINTED FOR THE RAY SOCIETY.
MDCCCLIV.

Reprinted with the permission of the Ray Society

JOHNSON REPRINT CORPORATION
111 Fifth Avenue, New York, N. Y. 10003

JOHNSON REPRINT COMPANY LTD.
Berkeley Square House, London, W. 1

First reprinting, 1968, Johnson Reprint Corporation
Printed in the United States of America

DEDICATION.

TO

PROFESSOR H. MILNE EDWARDS,

DEAN OF THE FACULTY OF SCIENCES OF PARIS; PROFESSOR AT THE MUSEUM OF NATURAL HISTORY;
MEMBER OF THE INSTITUTE OF FRANCE;
FOREIGN MEMBER OF THE ROYAL SOCIETY OF LONDON, OF THE ACADEMIES OF BERLIN, STOCKHOLM, ST. PETERSBURG, VIENNA, KONIGSBERG, MOSCOW, BRUSSELS, HAARLEM, BOSTON, PHILADELPHIA, ETC.

THIS WORK IS DEDICATED,

WITH THE MOST SINCERE RESPECT,

AS THE ONLY, THOUGH VERY INADEQUATE ACKNOWLEDGMENT WHICH THE AUTHOR CAN MAKE OF HIS GREAT AND CONTINUED OBLIGATIONS TO THE
‘HISTOIRE NATURELLE DES CRUSTACÉS,’
AND TO THE OTHER MEMOIRS AND WORKS ON NATURAL HISTORY PUBLISHED BY
THIS ILLUSTRIOUS NATURALIST.

PREFACE.

Having so lately, in my volumes on the Recent and Fossil Lepadidæ, expressed as strongly as I could, and with the utmost sincerity, the obligations under which I lie to very many naturalists, I will not here repeat my thanks, and will only say that the assistance formerly rendered me from so many quarters has been most kindly continued without intermission. The references under the Habitats, in which I may remark the names of Mr. Cuming and of Mr. Stutchbury, and of the British Museum, so often recur, show my deep obligations to these gentlemen and to Dr. Gray, and indeed to most of the British and several Foreign[1] collectors of recent and fossil shells. At the period when the [Introduction] to this volume was printed, I stated that I did not know whether the Palæontographical Society would publish the few British fossil [Balanidæ]; the Council has now honoured me by determining on this publication, so that these species will hereafter be more fully illustrated than they could be in the present volume. I cannot conclude this short preface, without again tendering my most grateful thanks to the Council of the Ray Society for the publication of my two volumes, and for the very kind manner in which they have acceded to all my requests.

DOWN, KENT; July, 1854.

[1] I feel under special obligation to Mr. Dana for several very interesting communications connected with the present subject, and for information derived from his magnificent work on the Crustacea, collected during the United States Exploring Expedition. Also to M. Bosquet, of Maestricht, for the loan and gift of several interesting fossils, described and illustrated with the utmost fidelity, in his beautiful “Monographie des Crustacés fossiles du terrain Crétacé du D. de Limbourg.”

[MONOGRAPH]

ON

THE CIRRIPEDIA.

INTRODUCTION.

My former volume, published by the Ray Society, treated only of the Lepadidæ, one family of the Cirripedia: I was induced to print it from having the materials ready, though this partial publication has been in some respects inconvenient. The Palæontographical Society has done me the honour to publish, with ample illustrations, the fossil species of this same family of Lepadidæ. This present volume completes my work on the sub-class Cirripedia.[2] I had originally intended to have published a small volume on my anatomical observations; but the full abstract given in my former volume, which will be illustrated to a certain extent in the plates appended to this volume, together with the observations here given under the [Balanidæ], appear to me sufficient, and I am unwilling to spend more time on the subject. In the volume on the Lepadidæ, I gave the specific or diagnostic characters in English and Latin: I have here left out the latter, inasmuch as I have appended at the end of this volume a Latin Synopsis of all the species, recent and fossil, of the whole class. To each species is added a reference to the pages and plates of my three volumes, so that the Synopsis will serve as a systematic index to the three: an alphabetical index to the present volume is also given. In the Lepadidæ, I gave an additional specific character, derived from the softer parts of the animal’s body: in the [Balanidæ], these parts are more alike in the different species, and I have found it impossible to give a diagnostic character thus derived. In those cases in which a Family contains but one genus, or a Genus but one species, I have assigned my reasons for the institution of such groups, but have given, as heretofore, only a single description in full: it would have been easy to have separated, by analogy, this description into one for the species, another for the genus or for the family; but as I believe such separation and subordination of the characters would have been largely conjectural, I have thought it best to act as I have done, and give, thus saving useless repetitions, only a single description, and leave it for my successors, when more genera or species are known, to separate, with such certainty as is ever possible, the generic from the specific characters.

[2] The number of the British fossil species of the [Balanidæ] and [Verrucidæ] in a recognisable state is so small, that I do not know whether it will be considered worth while to publish in the Palæontographical series more detailed illustrations than are given in this volume.

In nomenclature, I have endeavoured rigorously to follow the rules of the British Association, and have never, at least intentionally, broken through the great law of priority. In accordance with the rules, I have rejected, that is, as compulsory, all names given before the introduction of the binomial system in 1758. I have given much fewer synonyms than is usual in conchological works; for it is impossible to recognise with any approach to certainty, several even of the common European forms, in the short descriptions given by most authors; this holds good in many cases in which figures, in appearance excellent, have been added. I assert this the more confidently, from having had the advantage of having gone over some of the Linnean synonyms with Mr. S. Hanley. I may further venture to express 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.

NOMENCLATURE OF THE SHELL OF A SESSILE CIRRIPEDE.

SHELL. Fig. 1.

Orifice of shell, surrounded by the sheath. Sheath formed by the alæ (a—a.) and by portions of the upper and inner surfaces of the parietes (p—p.)

N.B. In [Balanus], and many other genera, the Rostrum and Rostro-lateral compartments are confluent, and hence the Rostrum has the structure of Fig. 2.

COMPARTMENTS.

Fig. 2.

Fig. 3.

Fig. 4.

Fig. 2. Compartment with two radii, serving either as a Rostrum or Rostro-lateral compartment.
Fig. 3. serves as a Lateral and Carino-lateral Compartment.
Fig. 4. serves as a Carina or Rostrum.

OPERCULAR VALVES.

Fig. 5. SCUTUM (internal view of).

Fig. 6. TERGUM (external view).

Fig. 7. TERGUM (internal view).

Sessile Cirripedes, partly from being attached to surfaces having very different characters, partly from undergoing a varying amount of disintegration, and partly from unknown innate causes, are extremely variable. Under the head of Variation, [in the Family Balanidæ] and [under the Genus Balanus], I have enlarged on this subject, and have shown that there is scarcely a single external character which is not highly variable in most of the species. As whole groups of specimens often vary in exactly the same manner, it is not easy to exaggerate the difficulty of discriminating species and varieties. It is absolutely necessary in most cases, for mere identification, that the valves of at least one specimen in a group should be disarticulated and well cleaned (for which purpose caustic potash is most useful), so that the internal characters may be examined. Whoever attempts to make out from external characters alone, without disarticulating the valves, the species, (even those inhabiting one very confined region, for instance the shores of Great Britain,) will almost certainly fall into many errors: hence it is, and can thus only be accounted for, that I have not seen one collection of British specimens with all the species, though so few in number, rightly discriminated; and in the large majority of cases, either two or three species, certainly distinct, were confounded together, or two or three varieties, as certainly not distinct, were separated from each other.

On the Names Given to the Different Parts of Cirripedes.

In my former volume I have stated that I found it indispensable, in part owing to the extreme confusion of the nomenclature previously used, to attach new names to several of the external parts of Cirripedes. Almost all these names are applicable to the [Balanidæ], or sessile Cirripedes, and to the [Verrucidæ]; but a few additional names are requisite, which, together with the old names, will, I hope, be rendered clear by the accompanying woodcuts. In sessile Cirripedes, the whole of that which is externally visible, may for convenience sake be divided into the operculum or opercular valves (valvæ operculares), and the shell (testa), though these parts homologically present no real difference. The operculum is seated generally some little way down within the orifice of the shell; but in very young specimens and in [Verruca], the operculum is attached to the summit of the shell, and the shell, without the operculum being removed, can hardly be said to have any orifice; though, of course, the opercular valves themselves have an aperture for the protrusion of the cirri.

The shell consists of the basis (called the support by some authors), which is membranous or shelly, and flat or cup-formed, and of the compartments (testæ valvæ), which vary from eight to four in number, and occasionally are all calcified together.

The compartment, at that end of the shell where the cirri are exserted through the aperture or lips of the operculum, is called the carina (fig. [1]); the compartment opposite to it is the rostrum,—these two lying at the ends of the longitudinal axis of the shell. Those on the sides are the lateral compartments; that nearest the carina, being the carino-lateral (testæ valva carino-lateralis), that nearest the rostrum, the rostro-lateral, and the middle one, simply the lateral compartment; but these three compartments are rarely present together. The rostro-lateral compartment, which always resembles fig. [2], and may be always known by having radii on both sides, is often absent; and not rarely the lateral and carino-lateral compartments are confounded together, or one is absent; in such cases the compartment that is left is simply called the lateral one. The compartments are separated from each other by sutures, which are often so fine and close as to be distinguished with difficulty. The edge of a compartment, which can only be seen when disarticulated from its neighbour, I have called the sutural edge (acies suturalis).

Each separate compartment consists of a wall (paries), or parietal portion (pp in woodcuts), which always grows downwards, and forms the basal margin; and is furnished on the two sides either with alæ (fig. [4]), or with radii (fig. [2]), or with an ala on one side and a radius (fig. [3]) on the other.

The radius[3] (adopting the name used by Bruguière, Lamarck, and others) differs remarkably in appearance (though not in essence) from the walls or parietal portion, owing to the direction of the lines of growth and the state of its usually depressed surface. In the upper part the radii overlie the alæ of the adjoining compartments: in outline (r, fig. [1], [2], [3]), they are wedge-formed, with their points downwards; their summits (and this is often a useful specific character) are either parallel to the basis or as in fig. [1] and [2], oblique. The radii are sometimes not developed.

[3] The radii have been called by Ranzani and De Blainville “areæ depressæ” (the parietal portions of the compartments being the “areæ prominentes”); by Poli, “areæ interjectæ;” by Gray, “sutures;” by Coldstream, “compartments of the second order,” (the parietal portions being those of the first order); by some authors as “intersticia.” I may here add that the scuta are the “ventral valves” of Gray, the “anterior” of Ranzani, and the “inferior opercular” of De Blainville: the terga are the “posterior valves” of Gray and Ranzani, but the “superior opercular” of De Blainville: the rostrum, on the other hand, is the “anterior valve” of Ferussac and the “ventral” of De Blainville; the carina being the “dorsal valve” of the latter author.

The alæ (so called by Dr. Gray) are overlapped by the radii and by part of the walls; they usually extend only about half way down the compartment (a fig. [3], [4], [1]); their summits are either parallel to the basis or oblique. The alæ of the several compartments, together with the internal, upper, thickened surfaces of the walls, against a shoulder of which the sutural edges of the alæ abut, have been called (by Dr. Gray) the sheath (vagina). The upper and greater portion of the sheath is marked by transverse lines, caused by the exuviation of the opercular membrane, as that membrane may be called, which unites the operculum all round to the sheath, or upper internal surface of the shell.

The carina has always two alæ, as in fig. [4].

The carino-lateral and lateral compartments have always an ala on one (the rostral) side, and a radius on the other (the carinal) side, as in fig. [3].

The rostro-lateral compartment (when present) has always radii on both sides, as in fig. [2].

The rostrum has normally alæ on both sides, as in fig. [4], but very often from fusion with the rostro-lateral compartments on both sides, it has radii on both sides, as in fig. [2].

The walls of the shell, the basis, and the radii, are in very many cases composed of an outer and inner lamina, united together by longitudinal septa; a set of tubes or pores being thus formed. The points of the longitudinal septa generally project beyond the laminæ, and are denticulated on both sides (see woodcut, further on;) the septa are sometimes branched, several irregular rows of pores between the two laminæ being then formed (see Pl. [7], fig. [3 b], and Pl. [10], fig. [1 g], [1 h]).

Operculum, or opercular valves.—These consist of a pair of scuta and a pair of terga. They are joined to the sheath of the shell by the opercular membrane.

Scutum (woodcut [5]): this valve is generally sub-triangular, and its three margins are the basal, the tergal, so called from being articulated with the tergum, and the occludent, so called from opening and shutting against the opposed valve. The angles are called from the adjoining margins, as basi-tergal, &c.; the upper angle being the apex. The scutum is ordinarily articulated to the tergum by an articular ridge (crista articularis), running up to the apex of the valve, and by an articular furrow, which latter receives the scutal margin of the tergum. The articular ridge, instead of projecting straight up from the valve, when laid flat on its external surface, often bends over to the tergal side, and is then said to be reflexed. On the internal surface of the valve, there is almost always an adductor pit or cavity (fossa adductoris), for the attachment of the adductor scutorum muscle: this pit is often bounded on its tergal and basal sides, by a ridge, called the adductor ridge (crista adductoris), which, in its upper part, is often confluent with the articular ridge. Beneath the adductor ridge, in the basi-tergal corner of the valve, there is often a lateral-depressor pit (fossa musculi lateralis depressoris), for the attachment of the so-called muscle; and this pit is sometimes furnished with crests.

Tergum, (woodcut [6] and [7]):—this valve, also, has three margins, the scutal, basal, and carinal; its upper end, or apex, is sometimes beaked; on the basal margin a spur (calcar) depends; the outer surface of the valve is depressed or longitudinally furrowed (sulcus longitudinalis) in the line of the spur. The part called the spur is often so broad, that the name becomes not very appropriate. The angles are denominated, from the adjoining margins, as basi-carinal, or basi-scutal angle, &c. On the under side, in the upper part, there is an articular ridge, and on its scutal side, an articular furrow, receiving the articular ridge of the scutum. In the basi-carinal corner of the valve there are often crests for the attachment of the tergal depressor muscle.

Sack, Body, Cirri, Mouth.—A slit-like orifice between the opercular valves leads into the sack, in which the body is lodged. The body consists of the six (perhaps the seven) posterior thoracic segments of the archetype Crustacean; the first of these six segments (or first two, if there be seven segments) is developed on its dorsal aspect into a part, which I have called the prosoma[4] (see fig. [1, c], Pl. [25]). There is no abdomen. The thoracic segments support six pairs of cirri. Each cirrus consists of a two-jointed pedicel, carrying two multiarticulated rami. Rarely there are articulated caudal appendages (appendices caudales) on each side of the anus. The prominent mouth consists of a labrum, palpi, mandibles, maxillæ, and outer maxillæ, the latter resembling a lower lip: these organs may be conveniently spoken of, after Milne Edwards, as gnathites. Within the sack, attached to its carino-lateral end, a folded membrane forms the branchiæ. The sheets of ova lying within the sack are called the ovigerous lamellæ.

[4] A discussion on the homologies of the different parts is given under the head of the [Metamorphoses of the Balanidæ].

I have often found it convenient to designate the membrane investing the body, lining the sack, &c., by its proper chemical name of chitine, instead of by horny, or other such equivalents; but when covering parts of the shell, for brevity’s sake I have often spoken of it as an epidermis, but I do not believe that such is its nature. When this membrane sends into the body of the animal 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.

Relative position of parts.—The centre of the generally flat basis, which is cemented to the supporting surface, is properly the anterior end, and the tips of the terga, often hidden within the shell, are properly the posterior end of the external covering; but I have found it more convenient to speak of the upper and basal surfaces and aspects, which hardly admit of any mistake. A line drawn from the centre of the basis, along the middle of the rostrum to the tips of the scuta, shows the strictly medio-ventral surface of the shell; and another line drawn from the centre of the basis, along the carina, to the tips of the terga, shows the strictly medio-dorsal line; but from the crooked course of these lines, I have found it far more convenient to speak of the rostral and carinal end or aspect of the different parts of the shell; this is the more necessary with respect to the internal parts of the animal, owing to their remarkable changes of position during the metamorphosis, whence it comes that the dorsal surface of the thorax faces partly dorsally, partly anteriorly or downwards, and partly even ventrally; and the ventral surface of the whole posterior part of the thorax faces upwards or posteriorly; but when we refer these parts to the rostral, carinal, basal, and upper ends of the shell, there can be no mistake. There has moreover been great confusion in these relative terms, as applied by different authors.

When a sessile Cirripede is held in the position in which they have generally been figured, namely with the basis downwards and the scuta towards the beholder, then the right and left sides of the Cirripede correspond with those of the holder.

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 confidence,—the specific determination being in each case made by myself.

CLASS—CRUSTACEA.
Sub-Class—CIRRIPEDIA.

Crustacea attached by the anterior end of the head, by cement proceeding from a modified portion of the ovaria; archetype composed of seventeen segments, with the three first of large size, and almost always developed into a carapace, not wholly exuviated, and capable of various movements; antennæ none; eyes rudimentary; mouth prominent, formed by the partial confluence of the labrum, palpi, mandibles, and two pairs of maxillæ; thorax attached to the internal sternal surface of the carapace, generally bearing six pairs of captorial, biramous, multiarticulated limbs; abdomen generally rudimentary; branchiæ, when present, attached to the under sides of the carapace; generally bisexual, when unisexual, males epizoic on the female; penis single, generally probosciformed, seated at the posterior end of the abdomen; oviducts none; metamorphoses complex.

Within the memory of many living naturalists, Cirripedes were universally looked on as belonging to the Molluscous kingdom; nor was this surprising, considering the fixed condition of their shells, and the degree of external resemblance between, on the one hand, Lepas and Teredo, and on the other hand, between [Balanus] and a Mollusc compounded of a patella and chiton. It is remarkable that this external false appearance overbore, even in the mind of Cuvier, his knowledge of their internal structure, namely, their lateral jaws, articulated appendages, and regular ganglionic nervous system, which now strike us as such conclusive evidence of their position in the great Articulate kingdom. Straus[5] was, I believe, the first who, in 1819, maintained that Cirripedes were most closely allied to Crustacea. But this view was disregarded, until J. Vaughan Thompson’s[6] capital discovery, in 1830, of their metamorphoses, since which time, Cirripedes have been almost universally admitted amongst the Crustaceans. It is well known, that it is hardly possible to give a definition of this great class, which shall include every member of it; nevertheless, even if the mature Cirripede alone be considered, the following characters, viz. the slight separation of the head and thorax, the latter generally bearing six pairs of appendages, and the being enclosed in a carapace—together with the periodical exuviation of the greater part of the external membranes, would, perhaps, suffice to show that it should be classed amongst Crustacea.

[5] Mémoires du Muséum d’Histoire Nat., tom. v, p. 381.

[6] Zoological Researches and Illustrations.

But it still remains undecided what rank in this class Cirripedes should hold. Before briefly discussing this point, it is indispensable to indicate their essential characters, which I will immediately attempt. For as long as it remained doubtful which was their anterior extremity, which the ventral or dorsal surface; as long as the peduncle was thought by one naturalist to be the legs, by another the abdomen, in a modified condition, it was hopeless to compare Cirripedes with ordinary Crustaceans, and assign to them their due rank.

In the larva in the first stage, an eye and two pairs of antennæ are in process of formation or are developed; here, then, according to the analogy of all Crustaceans, we have evidence of the existence of the first three cephalic segments. The mouth always consists of three pairs of gnathites, and hence again, from analogy, this part may be inferred to be formed of, and supported on, three other segments; making thus far six segments. In two Orders out of the three into which Cirripedes may be divided, namely, in the [Abdominalia] and [Apoda], eight quite distinct segments succeed the mouth; of these the first differs slightly from the seven succeeding segments, and may, I think, be safely considered as forming the seventh (cephalic) segment. The next seven segments resemble each other in all essential respects, and are no doubt the normal, seven thoracic segments. These, in both the above orders, are succeeded by three smaller segments, which differ in structure from the thoracic segments, and must be abdominal. Hence we here have, altogether, seventeen segments. It should, however, be observed that in the two orders just referred to, each includes only a single species; but I know of no good reason why, on this account, their structure should be valued the less. In the third order, the [Thoracica], which includes all common Cirripedes, two segments with their appendages are missing out of the eight that should succeed the mouth; from the open interval in the pupa, between the mouth and first pair of natatory legs, and from some other reasons, I believe that the two missing segments are the seventh and eighth, or last cephalic and first thoracic segments, and that they have coalesced close posteriorly to the mouth.[7] In the order [Thoracica], the abdomen is quite rudimentary, though often still bearing caudal appendages; in the pupa, however, of this order, as in the mature animal of the two other orders, it is formed of three segments. Hence I conclude that, notwithstanding the absence of the above two segments with their appendages in the [Thoracica], the archetype Cirripede may be safely said to be composed of seventeen segments.

[7] This question and the whole subject of the homologies of the several parts of a Cirripede, will be discussed under the head of the [Metamorphoses of the Balanidæ].

In the classification of Crustacea, the relation and number of the segments of the different parts of the body, are viewed both by Prof. Milne Edwards[8] and Mr. Dana,[9] as of the highest importance. I may premise that both these authors divide the Crustacea into Podophthalmia, Edriophthalmia, and Entomostraca; Milne Edwards making a fourth legion, the Branchiopoda, and another division, including Limulus, of equal value to the above four legions altogether; whereas Dana sinks Limulus and the Branchiopoda under his Entomostraca. As far as concerns our present discussion on Cirripedes, the first three divisions, as valued by Dana, will best serve as standards of comparison; but it is not unimportant to our present purpose, as showing how difficult it is to weigh the value of the higher divisions of a Class, to observe the wide difference in opinion of two naturalists, so eminent for their knowledge of the class in question and for their high abilities.

[8] Annales des Sciences Nat., tom. xviii, p. 120, 1852.

[9] Crustacea: ‘United States Exploring Expedition,’ p. 1395, 1852.

In the order [Thoracica], including all common Cirripedes, the cephalic and thoracic segments are as much confounded together (but with coalescence and abortion of two middle segments) as in most Podophthalmia; but in the two other orders, the cephalic and thoracic segments are as distinct as in the Edriophthalmia. The number of the segments, however, which strictly appertain to the anterior part of the head and mouth, being only six, is an Entomostracan character; on the other hand, the first pair of cirri in the [Thoracica], has some claim from their position, apparent functions, and separation from the succeeding pairs, to be said to belong to the mouth; on which view, the first nine segments would, in function, be cephalic, as in the highest Crustaceans. The fewness of the segments of the abdomen, and their not bearing in two of the orders appendages, is an Entomostracan character.

Cirripedes are permanently attached, even before their final metamorphosis, by a tissue or cement, first debouching through the second pair of antennæ, and, subsequently, in most cases, through special orifices, penetrating the anterior part of the head; this cement proceeds from glands, which certainly are modified portions of the ovarian system. This fact I consider of the highest classificatory importance, for it is absolutely the one single character common to all Cirripedes, besides such as show only that these animals belong to the articulated kingdom, and are Crustaceans. No structure of this kind has hitherto been observed in any other member of the class or kingdom. Even in the Suctorial Entomostracans, which become immoveably attached to the fish on which they prey, the males are free; and the means of attachment, as far as known, are quite different.

Both the first and second pairs of antennæ are absent in the mature animal; for the three terminal segments of the antennæ of the pupa, which may always be found cemented under the centre of the surface of attachment, are functionless, after maturity. The eyes are rudimentary, and are singular from being seated far from the anterior extremity of the head. In their rudimentary state, and in the absence of antennæ, we have characters common with certain Suctorial Entomostracans; and this similarity apparently arises from the fixed condition of the animals of both groups.

The carapace, which covers the dorsal surface of the larva in the first stage, in the last larval or pupal stage is developed so as to enclose, like a bivalve shell, the whole body. In the mature Cirripede, the whole external covering, whether shell and operculum, or capitulum and peduncle, can be conclusively shown to be the carapace of the pupa, modified. In thus enclosing the mouth and whole body, the modified carapace resembles that of several Entomostracans; but in being apparently formed (as I hope hereafter to show) by the development of the third segment of the head, and in consisting generally of distinct sclerodermic plates, arranged in an imbricated order, there is, I think, a closer resemblance to the same part in some of the Podophthalmia. The carapace, or portions of the carapace, being capable of other movements, besides merely opening and shutting, differs, I believe, from that of all other Crustaceans; as it likewise does[10] in the greater part not being periodically moulted.

[10] The carapace, however, of the Isaura, a Branchiopod, according to M. Joly (‘Annales des Sc. Nat.,’ 2 ser. vol. xvii, p. 293), is not moulted.

Moreover, in all Cirripedes there is another striking peculiarity connected with these parts, namely, the exclusive attachment of the whole thorax or included body to the internal ventral or sternal surface of the carapace and head. In the pupa, the thorax, as in all Crustaceans, opens into, and is continuously united with, the large anterior part of the head; but from the singular fact that the thorax of the young Cirripede is developed not within the thorax, but within the head of the pupa (Pl. [30], fig. [2]), with its longitudinal axis placed at right angles to that which it held in the pupal condition (the mouth and the whole exterior being developed conformably with that of the pupa), it comes to pass after the metamorphosis, that the Cirripede is, as it were, internally cut in twain (compare Pl. [25], fig. [1], and Pl. [30], figs. [2] and [3]). Thus it is, as will hereafter be more fully explained, that the sack originates, and thus the body becomes attached to the internal ventral surface of the carapace and front of head.

The thorax in two of the Orders bears no appendages, but in all common Cirripedes it is furnished with six pairs of biramous, multiarticulated cirri, which have a peculiar character, different from the limbs of other Crustaceans, not being natatory, ambulatory, or branchial, but “captorial” or fitted for sweeping the water, and thus catching prey.[11] The cirri, at least the anterior pairs, can, besides other movements, lengthen and shorten themselves; and this Milne Edwards[12] states is the case with the Podophthalmia, and is considered by him as an important character. The cirri of the first pair are attached on each side close to the bases of the mandibles, and, as already remarked, have some claim to be considered as maxillipeds or mouth organs. The three or the four posterior pairs of cirri in the [Balanidæ], form a series somewhat distinct from the two or three anterior pairs, thus recalling a characteristic feature in the Edriophthalmia.

[11] M. A. Hancock, in ‘Annals and Magazine of Natural History,’ 2d series, 1849, p. 312, speaks of the cirri acting like a prehensile net.

[12] ‘Annales des Sciences Nat.,’ tom. xviii, p. 121, 1852.

The mouth is prominent, and is formed by the partial confluence of the labrum, palpi, and lower segments of the mandibles, and of two pairs of maxillæ; it is capable of movement as a whole; in this respect we are reminded of the Suctorial Entomostracans; but I believe the above type of structure of the mouth is peculiar to Cirripedes.

The alimentary canal is simple, but can be distinctly divided into—(1st) an œsophagus, singular from the bell-shaped expansion of its lower end; (2d) the stomach, which is directed forwards and then doubled back; and (3d) the rectum. There is no distinct liver. The circulation is lacunal. In one family there are well-developed branchiæ, which differ entirely in their homologies and position from these organs in all other Crustaceans. In the nervous system, the sub-œsophageal ganglions vary in concentration from that degree observed in the lower Macroura, to that in the highest Brachyoura; but the supra-œsophageal ganglions are always much less concentrated, and are even embryonic in condition; presenting a difference not observed in other Crustaceans. On the under side of the sub-œsophageal ganglion, two nerves, apparently splanchnic, arise, and run almost parallel and under the collar surrounding the œsophagus; they are very remarkable from their great size, and from forming a plexus together with a large branch, arising on each side from the collar close behind the supra-œsophageal ganglion,—a structure unlike anything observed in other Crustaceans. The eyes, as already remarked, are rudimentary, and singular from being imbedded at a distance from the anterior end of the animal. In the basal confluent segments of the outer maxillæ there are two orifices, leading into little sacks, which I believe are olfactory organs: again there are two other orifices on each side of the thorax, beneath the first pair of cirri, leading into sacks, with a curious elastic vesicle suspended within them; and these I can hardly doubt are acoustic organs. Of these orifices and organs, there is no trace in the same relative positions in any known Crustacean.

Cirripedes are ordinarily bisexual, in which they differ from all Crustaceans: when the sexes are separate, the males are minute, rudimentary in structure, and permanently epizoic on the females; to these latter facts we have a partial analogy in some of the Suctorial Entomostracans; but a far closer analogy in certain Rotifers, which are considered by many naturalists as Crustaceans; but to the above subject I shall almost immediately have to recur.

The male excretory organ is probosciformed and capable of the most varied movements; it is single and medial; it is seated (in the one instance in which this point can be safely judged of) at the extremity of the abdomen, and therefore near the normal position of the anus; in all these respects there is a very great difference from other Crustaceans, in which the male organs are laterally double, and are not seated at the extremity of the abdomen. In regard to the female organs, the ovarian tubes and cæca inosculate together: there are no oviducts; the ova, connected together by membrane, and so forming the “ovigerous lamellæ,” become exposed by the exuviation of the lining tunic of the carapace or sack, and by the formation of a new tunic on the under side of these lamellæ; a process, I believe, unknown in other Crustaceans.

The metamorphoses are highly complex. The larva in its first stage bears a very close general resemblance, in having three pairs of natatory appendages, the first being uniramous and the two others biramous, and in having a single eye on its broad anterior front, to the larvæ of most Entomostracans; but I cannot avoid the belief, that this resemblance is only apparent, and not essential; and of false resemblances, how many instances occur in the animal kingdom! In the larva, when first freed from the egg, both pairs of antennæ are in process of formation within envelopes: the mouth is probosciformed and capable of movement, but is destitute of gnathites; it occupies a position between the three pairs of natatory limbs; and these limbs I must believe, for reasons hereafter to be assigned, answer (improbable as I am well aware it must at first appear) to the second, third, and fourth thoracic legs of the archetype Crustacean: the two hinder pairs of limbs apparently soon become captorial, or fitted to secure prey. Now, I cannot find in the published accounts of the larvæ of Entomostracans, any that answer to this description.

The larva in the last stage might be included in the vast class of Entomostracans: the attachment of the eyes to the singular apodemes produced inwards from the basal segment of the great prehensile antennæ, and the development of only the posterior six pairs of thoracic limbs, are its chief peculiarities: but its rudimentary mouth, owing to its transitional or pupal condition, renders the assignment of its proper rank difficult.

Having now given this short comparative sketch of the structure of a Cirripede, I may venture to express strongly my opinion, that the group is formed on a distinct type; as different from the other three or four main Crustacean groups, namely, the Podophthalmia, Edriophthalmia, Branchiopoda, and Entomostraca, as these differ from each other; the differences, moreover, being of the kind considered by the highest authorities on this subject, as the most important. It should be observed that there is no special blending at either end of the Cirripedial series, towards any one of the other main groups of Crustacea; it is hardly possible to take some one Cirripede, and say that it leads, more plainly than some other Cirripede, into ordinary Crustaceans. Moreover, a great range of structure, as we shall soon briefly show, is included within the group: I can adduce three or four undoubted Cirripedes, very considerably more different from each other, than any two members within the sub-class Podophthalmia, or within the Edriophthalmia, or the Branchiopoda, and quite as different as within the Entomostraca.

The opinion here expressed, that Cirripedes form a sub-class of equal value with the other main Crustacean groups, I am well pleased to find, accords with Mr. Dana’s[13] view, who remarks that this sub-class “has so many peculiarities of structure, that it should be regarded as a distinct type, rather than a subordinate division of the third (or Entomostracan) type.” M. Milne Edwards,[14] after dividing all Crustacea into two groups, divides one of them into four legions; and of one of these, the Entomostraca, he makes the Cirripedes a sub-group. I feel so entire a deference for any opinion on affinities or classification expressed by Milne Edwards, that I differ from him with the greatest hesitation. He does not give his reasons for assigning so subordinate a rank to Cirripedes, but I imagine it is from the nature of their metamorphoses: but if this be the case, I cannot understand why he should assign to his Branchiopods a rank equal to his Entomostracans. Moreover, I must repeat, that I do not believe that the larvæ do resemble the larvæ of Entomostracans and Branchiopods nearly so closely as at first appears to be the case. I may add, that Burmeister[15] has assigned to the Cirripedes a place amongst the Crustacea, almost equally subordinate to that given to them by Milne Edwards.

[13] ‘Crustacea: United States Exploring Expedition,’ p. 1407, 1852.

[14] ‘Annales des Sciences Nat.,’ tom. xviii, p. 120, 1852.

[15] ‘Beiträge zur Naturgesehichte der Rankenfüsser,’ 1834.

That Cirripedes have some special affinity to the Entomostraca, may be inferred from the fewness of the cephalic appendages, the biramous legs, the state of the abdomen, and the form of the carapace. Perhaps in the peculiar state of confluence of the lower segments of the gnathites, in the aborted antennæ, the rudimentary eyes, and in the minute parasitic males (when such exist), there is a more direct relation to the Suctorial division of the Entomostraca; but some of these resemblances are probably only analogical, resulting from the fixed condition of both groups. It should not be overlooked, that out of the three orders into which Cirripedes may be divided, in the two latter, the mature animal presents hardly any resemblance to an Entomostracan. From the distinct presence in either pupa or mature animal of the fourteen segments of the cephalo-thorax; from the apparent composition of the carapace, as will be subsequently explained; and from the concentrated condition of the nervous system, one is led to glance at the higher Crustacea; and here we shall find amongst the Podophthalmia, one aberrant group of low organisation, namely, that including Phyllosoma, Amphion, &c., in which more points of resemblance to Cirripedes may be detected, than, as I believe, in any other group whatever; for we here see that remarkable elongation of the head in front of the mouth, so eminently characteristic of Cirripedes; we have a carapace overlapping the thorax, which is sometimes free beneath; we have the abdomen sometimes almost obsolete; we have biramous legs: and especially we have the posterior cephalic and the first thoracic appendages more or less rudimentary and obsolete; and this, I infer from Mr. Dana, is a very rare phenomenon, though characteristic of all ordinary Cirripedes, in which the seventh and eighth segments with their appendages have disappeared. In the order including Phyllosoma, &c., namely, in the Macroura, the ganglions which give nerves to the five posterior thoracic limbs, are distinct from the great sub-œsophageal ganglion which supplies the several anterior appendages; this is the case with those Cirripedes in which all the infra-œsophageal ganglions are not concentrated into one. In the Macroura and Brachyoura, the first pair of legs almost always differs in structure from the others, so does the homologous or second cirrus in Cirripedes differ from the four succeeding pairs; in some few Macroura, the second leg is antenniformed, so in some few cases is the homologous (or third) cirrus; J. Vaughan Thompson was even struck by the resemblance in the curious, doubly pectinated spines on the anterior limbs of Mysis (allied to Phyllosoma[16]), and on those of many Cirripedes: these several latter resemblances may be small, but certainly I do not believe that they are accidental. Now the little group of Crustaceans, which includes Phyllosoma, &c., has lately been placed, by Milne Edwards, as a satellite amongst the Macrourous Podophthalmia; it leads into the Stomopoda, and likewise, as has been noticed by many authors, into the sub-class Branchiopoda, which latter sub-class is considered by Mr. Dana as only a part of the Entomostraca; this group, therefore, exhibits affinities radiating in several directions, and amongst these lines of relationship, one more must, I believe, be added, plainly directed towards the Cirripedia.

[16] M. Martin St. Ange (‘Mémoire sur l’Organ. des Cirripèdes,’ 1835, extrait des ‘Savans Etrangers,’ tom. vi) has compared the mouth of Lepas with that of Phyllosoma, and has given comparative figures; but the resemblance is founded, I believe, on quite false homologies.

One naturally wishes to ascertain how far Cirripedia are highly or lowly organised and developed; but in all cases this, as it seems to me, is a very obscure enquiry. Mr. Dana considers that, in Crustacea, the greater or less centralisation of all the appendages round the mouth is the main sign of high development; on this view, the anterior part of a Cirripede, from being so much elongated, must be considered as very low in the scale; the whole posterior part of the body, on the other hand, is, in ordinary Cirripedes, brought close to the mouth; but this is effected by the abortion of the seventh and eighth segments of the cephalo-thorax and of the whole abdomen, and so, I presume, would not, in Mr. Dana’s estimation, raise the class much in the scale. Von Baer[17] considers that the perfection of the type of any animal is in relation to the amount of “morphological differentiation” which it has undergone; on this view, Cirripedes ought to stand high in the scale, for they differ much morphologically from the type of the class to which they belong; as indeed is shown by the long time that elapsed before their true position, namely amongst the Crustacea, was even suspected; but something more must, I think, be added to Von Baer’s definition; for, to take as an example the eyes of a Cirripede,—as seen in the first larval stage, there is only one eye, and that most simple; in the pupa there are two, both compound, and furnished with complicated muscles; lastly, in the mature animal there are still two, but of very minute size, often almost confluent, and of the simplest structure; hence, then, there has been much morphological differentiation, but it is almost a contradiction in terms to speak, in relation to such a case, of perfection of type; and what has happened to one organ, might happen to other organs, and so to the whole animal. Lastly, under a physiological point of view, and taking the [Balanidæ] as the most perfect type of the class, the sub-œsophageal portion of the nervous system is highly concentrated; the organs of sense, excepting the eyes, seem more largely developed than in ordinary Crustaceans; the circulating system is of the simplest kind, being only lacunal; special Branchiæ, however, are developed by the metamorphosis of, as I believe, a special organ, occurring only in the Lepadidæ; the digestive organs are very simple, from not having any distinct liver; the generative system is very low, for both sexes are generally united in the same individual; and the testes and ovaria closely resemble each other. On the other hand, the thoracic limbs are, to a considerable extent, specialised in their structure and functions; only the three posterior pairs strictly resembling each other. Lastly, the dermal and muscular systems are complicated, and not, to use Professor Owen’s term, by mere vegetative repetition, as will be obvious to any one who will study the beautifully constructed and modified carapace—that is the operculum, shell and basis—of a [Balanus]. On the whole, I look at a Cirripede as a being of a low type, which has undergone much morphological differentiation, and which has, in some few lines of structure, arrived at considerable perfection,—meaning, by the terms perfection and lowness, some vague resemblance to animals universally considered of a higher rank.

[17] English Translation, in ‘Scientific Memoirs,’ 1853, vol. i, p. 228.

It has been seen that I divide the Cirripedia into three orders,—the [Thoracica], [Abdominalia], and [Apoda]; between which the fundamental difference consists in the limbs or cirri being thoracic in the first, abdominal in the second; and entirely absent in the third. For the sake of showing the range of character in Cirripedes, to which allusion has been made, I will briefly indicate the leading differences in each order. In the [Thoracica], three families are included,—the [Balanidæ], or sessile Cirripedes, the [Verrucidæ], remarkable from their quite asymmetrical shell, and the Lepadidæ, or pedunculated Cirripedes. The great difference in external appearance between these three families is known to all naturalists. Even within the one family of Lepadidæ there are great differences in external appearance, as will be admitted on comparison of Lepas, Pollicipes, Conchoderma, &c.; but we have also important internal differences, as in the case of Anelasma, in which the cirri are barely articulated, and are not capable of seizing prey, whilst the mouth is almost probosciformed, with the outer maxillæ and palpi rudimentary: still more important are the differences in [Alcippe], in which the cirri of the first pair act as brushes; the second, third, and fourth pairs being quite aborted; and the fifth and sixth pairs consist only of four segments, with one of the two normal rami converted into a crenated, button-like projection, for the sake apparently of triturating food; [Alcippe], also, is very remarkable in being destitute of a rectum and anus. In this same genus [Alcippe], in Ibla and Scalpellum, there are either separate males or Complemental males, some of which are so utterly abnormal in their characters, that by no definition which I could frame, could they be included even in their proper Order, much less in their proper Family.

In the second order of [Abdominalia] (Pl. [23] and [24]) the seventh or last cephalic segment is quite distinct, and bears rudimentary organs, answering to the first pair of maxillipeds of ordinary Crustaceans, of which organs, and of the segment supporting them, there is no trace in the [Thoracica]: the seven succeeding thoracic segments are destitute of any appendages; but the three segments of the abdomen bear three pairs of cirri. The mouth is peculiar in the labrum being developed into very large, moveable, lancet-formed organ; and the lower end of the œsophagus is armed with beautiful discs of teeth, and brushes of hairs,—a structure confined to this order. The male resembles the male of [Alcippe]; and the latter genus seems to be the connecting link between the [Thoracica] and [Abdominalia]. But the most important character of this latter order, in which it differs from [Alcippe], and all other known Cirripedes, is in its metamorphoses; all the first changes are merely indicated by changes in form in an egg-like larva, without the development of distinct organs; and the last, or pupal condition, which is attained within the sack of the parent, is very peculiar, by the entire absence of natatory limbs.

The third order of [Apoda] is the most peculiar of all; it contains, like the last, only one known species: the most acute naturalist, I am convinced, if he had not made the class his special study, would never even have suspected that this animal was a Cirripede. We see much magnified in Pl. [25], fig. [7] a naked, plainly-articulated animal, resembling the larva or maggot of a fly, attached by two threads; and these threads, on analysis, can be clearly shown to be the last rudiment of the carapace, specially modified. The last cephalic, the seven thoracic, and the three abdominal segments, are all equally destitute of appendages. The mouth is suctorial, and constructed on a plan unlike, I believe, anything known in the articulate kingdom; for the mandibles and maxillæ have rotated on their axes, and stand back to back; they can act only by tearing open a slit, and this action is performed in a hood, formed by the confluence of the broad palpi and labrum. Although the œsophagus is distinct, there is no stomach or anus. Lastly, owing to there being no carapace, the ova are developed, differently from in all other Cirripedes, within the thorax.

I will close this preliminary discussion on the confines and type of the sub-class, by recalling attention, now that a sketch has been given of the three Orders, to the remark before made, that a wide range of structure is included within it, and by reurging that the Cirripedia should be ranked, not as one of the subordinate groups, but as one of the main divisions of the Crustacea.

On the Sexual Relation of Cirripedes.

Cirripedes are commonly bisexual or hermaphrodite, but in Ibla, Scalpellum, and [Alcippe], members of the Lepadidæ in the order [Thoracica], and in [Cryptophialus] in the order [Abdominalia], the sexes are separate. As two of these genera were described in my former volume, and two others ([Alcippe] and [Cryptophialus]) are described in this volume, I may as well here give a brief summary of the facts as yet known on this very curious subject. The Males, in the above four genera, present a wonderful range of structure; they are attached in the usual way by cement proceeding from the not-moulted antennæ of the pupa, to different parts, in the different species, of the female. These males are minute, often exceedingly minute, and consequently generally more than one is attached to a single female; and I have seen as many as fourteen adhering on one female! In several species the males are short-lived, for they cannot feed, being destitute of a mouth and stomach. As the females are longer lived, successive crops of males, at each period of propagation, become attached to her. It is the females in the above genera which retain the characters of the genus, family, and order to which they belong; the males often departing widely from the normal type. Some of the males are rudimentary to a degree, which I believe can hardly be equalled in the whole animal kingdom; they may, in fact, be said to exist as mere bags of spermatozoa. So widely do some of them depart in every character from their class, that twice it has happened to me to examine specimens with a little care, and not even to suspect, until a long period afterwards, that these males were Cirripedes.[18]

[18] In my volume on the Lepadidæ (p. 200) in searching for analogies for the permanently epizoic and rudimentary condition of the male Cirripedes, I quoted two cases, which I believe are now known not to be analogous; namely, the Syngamus trachealis of Von Siebold, and the worm-like Hectocotyle, which latter was quite lately supposed to be a male Cephalopod, but has now been ascertained to be only one of the arms of the male wonderfully adapted and organised as a sperm-receptacle. The Asplanchna, the mouthless male of a Rotifer, (p. 292) alone remains for me.

In Scalpellum Peronii, and villosum, the males are but little abnormal, for if classified independently of their sexual relations, they would be considered as immature specimens of a new genus, standing next to Scalpellum; in Scalpellum rostratum, the male would form another and rather more distinct genus. The males, in the latter, are attached to the other sex, between the basal edge of the labrum and the adductor scutorum muscle; but in S. Peronii and villosum they are attached lower down, in the furrow between the two scuta, and are thus protected: in these three species, the internal parts of the male present nothing particular. In Ibla, the males are attached low down within the sack of the female; they may be said to consist of a mouth surmounted on a long peduncle, for there is no capitulum or general covering, and the whole thorax is in a rudimentary condition, the cirri being reduced to two distorted pairs. As these males certainly moult several times and grow a little, they must feed; and as they have no cirri fit for action, they must seize their food by the contortions of their peduncle, which we know homologically consists of the three first segments of the head. The movements of the peduncle must, also, supply those of the probosciformed penis, almost invariably present with other Cirripedes, but here absent. If compelled to class these males without regard to the female, great difficulty would be experienced; we could hardly place in the family of the Lepadidæ, a Cirripede without a capitulum, and without cirri, those very organs which give their name to the class, and with a thorax reduced to the dimensions of a lower lip; yet, if the presence of a peduncle did determine the classifier to place these males amongst the Lepadidæ, then undoubtedly the character of the mouth, &c. would fix their position next to Ibla.

The males of Scalpellum vulgare, ornatum, and rutilum, resemble each other in all essential points, and differ wonderfully in appearance and structure from all ordinary Cirripedes. They consist of a minute flattened bag with a small orifice at the summit, and at the lower end attached by the cemented pupal antennæ. On each side of the orifice, there is a pair of calcareous beads, representing the two scuta and two terga of ordinary Cirripedes; and between the scuta a minute black eye is generally conspicuous. In S. ornatum the beads, I may remark, on the two sides are not equal; those either on the right or on the left side, being larger than those on the opposite side, so that the animal externally is asymmetrical. Inside, within a tubular sack, the thorax is lodged, supporting four (instead of six) pairs of limbs; and these, instead of forming biramous, multiarticulated, captorial cirri, are reduced to almost a rudiment, supporting a few long sharp spines, which apparently act only as defensive organs. At the end of the thorax there is seated a large abdominal lobe, which does not occur in the other sex. Hence the thorax, though rudimental, has been specially modified. Of the mouth and stomach there is not a vestige. Constructed as these males are, assuredly they have no claim to be ranked amongst the Lepadidæ or pedunculated Cirripedes; nor is it possible to class them in any group whatever of ordinary Cirripedes. In S. vulgare the males are attached, often several together, to the extreme edges of the two scuta, and therefore immediately over the orifice leading into the sack; in S. rutilum and ornatum, they are attached in concavities on the under side of both scuta, just above the depression for the adductor scutorum muscle. In the former of these species, the pit for the reception of the male is formed by shelly matter not having been deposited over a certain space on the under side of the valve; and the pit is converted by a covering of membrane into a pouch. As there are two scuta so there are two pouches, in each of which a male is lodged; hence, according to the Linnean nomenclature, Scalpellum ornatum may be said to belong to Diandria monogynia. As these males, from being mouthless, soon die, they are succeeded by successive pairs; the pupa being led by a wonderful instinct to crawl into the pouch, and there undergo its metamorphosis.

Lastly, the males of [Alcippe] and [Cryptophialus] (Pl. [23], fig. [19], and Pl. [24], fig. [19]) are remarkable for their similarity to each other, considering the essential dissimilarity of the two females. The females live in cavities which they excavate in the shells of Molluscs, and within which they are attached by a horny disc; this disc is the only part of the outer integument which is not frequently moulted, and, apparently in consequence, the males are attached to its edges. It results from this position, that the males are protected by being enclosed within the cavity excavated by the female; and it further results, that the males are attached at a considerable distance from the orifice of the sack of the female, into which the spermatozoa have to be conveyed; and to effect this, the probosciformed penis is wonderfully developed, so that in [Cryptophialus], when fully extended, it must equal between eight and nine times the entire length of the animal! These males, like those last mentioned of Scalpellum, consist of a mere bag, lined by a few muscles, enclosing an eye, and attached at the lower end by the pupal antennæ; it has an orifice at its upper end, and within it there lies coiled up, like a great worm, the probosciformed penis, and beneath it a single testis, with a single vesicula seminalis. These organs complete the whole organisation of the male; for there is no mouth, no stomach, no thorax, no abdomen, and no appendages or limbs of any kind. Yet all these parts are present in the female. I know of no other instance in the animal kingdom of such an amount of abortion. The whole exterior of these males evidently is composed, as in all ordinary Cirripedes, of the three first cephalic segments; of the fourteen succeeding segments of the archetype Cirripede we have not a vestige, excepting the probosciformed penis, which, from analogy, should arise from the ventral apex of the seventeenth segment, the first three segments of the head being counted in the seventeen. Here, then, fourteen out of seventeen segments have aborted, the tip of the seventeenth having coalesced with the third cephalic segment! I am tempted just to notice the case of [Proteolepas], in the order [Apoda], as showing, within the limits of the same sub-class, a wonderful amount and diversity in abortion; for in [Proteolepas], the three anterior cephalic segments are reduced to the merest rudiment, encasing the cement-ducts, the fourteen succeeding segments being unusually well developed; whereas in the above described males, we have just seen the three anterior segments fully developed, whilst the fourteen succeeding segments are lost or have coalesced with the others; so that within the same sub-class all seventeen segments of the archetype have almost disappeared.

It may be asked how I know that the several above described rudimentary epizoons are really the males of the Cirripedes to which they are attached. Even if the whole course of the metamorphoses had not been known in three of the cases, the mere fact of these epizoons being cemented by the three terminal segments of their peculiar, pupal antennæ, would have been sufficient to have shown that they belonged to the class of Cirripedes. In nearly every case, I was able to demonstrate, and not in one or two but in many specimens, that these epizoons were males; and as in several instances the spermatozoa were developed, and as, notwithstanding, in no instance was there a vestige of ova or ovaria, it may safely be concluded that they were not hermaphrodites, and therefore required females of some kind. If these epizoic Cirripedes had been independent animals, as they all belong to the same sub-class, and all have such peculiar habits, it might have been expected that they would have shown some special affinity towards each other; but this is not the case; the epizoon of Ibla is more nearly related to Ibla, and the epizoon of Scalpellum more nearly related to Scalpellum, than are these epizoons to each other. If the several epizoons were classed by themselves, they would be grouped in divisions, corresponding with those of the Cirripedes on which they are attached, which is just what might have been expected if these latter were their females. There are, also, many special relations between the male epizoons and the Cirripedes to which they are attached; thus, the mouth of the epizoon of Ibla, is so like the mouth of Ibla, which is peculiar in several respects, that I should easily have recognised it as belonging to a member of that genus. Scalpellum villosum is remarkable as one out of only two or three members of the whole Family, which is destitute of caudal appendages, so is its male epizoon; again, S. villosum is unusually spinose, so is its male epizoon; on the other hand, Scalpellum ornatum is remarkably smooth, so is its male epizoon; and I could give other similar instances. Will it be believed that these coincidences are accidental, and that the epizoons have no special or sexual relation to the Cirripedes to which they are attached?

One other instance of coincident structure is so important, that it must, even in this sketch, be noticed; the prehensile antennæ of the pupa are most important and complicated organs, and differ in the different genera of the same family; they are preserved in a functionless condition throughout life, and in two instances I was able accurately to compare these organs in the epizoon and in the Cirripede to which it was attached, and they were identical in every particular. The full force of the excessive improbability of this resemblance, and of the above coincidences in structure, on the supposition of the epizoon and its support not being sexually related, will hardly be perceived without referring to the facts given in detail in my former volume.

Lastly, in the case of [Cryptophialus] (and indirectly in that of [Alcippe]) the nature of the male epizoon is, I think, actually demonstrated; for I traced both it and the female or ordinary form of [Cryptophialus], through the same several larval stages, from the egg, enclosed within the sack of the female, to the pupa and mature animal. Moreover, if the male nature and sexual relation to the supporting Cirripede, be admitted in any one of these epizoons, then so close is the agreement in habits, and to a certain extent in structure, in all the foregoing epizoons, that probably no one admitting one instance would dispute the others, and further evidence would even be superfluous. Indeed, had it not been for the following facts, I should not have brought forward, either here in abstract, or in other places in detail, so many arguments and so much evidence.

I have as yet not entered in detail on the sex of the supporting Cirripede: in [Cryptophialus], [Alcippe], and in one species of Ibla, I was able to demonstrate in many specimens, that all the male organs, internal and external, were entirely absent; and consequently that these Cirripedes were exclusively female. In Scalpellum ornatum, also, there is no trace of external male organs (the state of the four dried specimens not allowing the internal organs to be examined), and there cannot be any reasonable doubt that this species likewise is exclusively female. It should be borne in mind that the male organs, external and internal, are most easily discovered, and that in the above cases I had an abundant supply of excellent specimens. On the other hand, in Ibla Cumingii, and in four species of Scalpellum, I was able to demonstrate in the supporting Cirripede the presence of all the male organs, as well as of the female; and in the vesiculæ seminales of several specimens, both in the Ibla and in Scalpellum vulgare, spermatozoa were contained; the male organs, however, not being very amply developed. These species, consequently, are not exclusively female, but are hermaphrodite, though having male epizoons attached to them. This statement, I am well aware, is enough, at first, to cast a doubt on all that I have said; but let any one reflect on the evidence, of which I have here given a summary, and which has been elsewhere given in full, and I think he must admit that at least those epizoons which are exclusively male, and which are attached to Cirripedes exclusively female, are sexually related and form one species; but if he admit this, he cannot possibly escape from the conclusion that some of the other epizoons, for instance that of Ibla quadrivalvis, are the males of the hermaphrodites to which they are attached,—these epizoons not exclusively impregnating the ova of a female, but aiding the self-impregnation of an hermaphrodite. Hence I have called these males Complemental Males, to show that they do not pair with a female, but with a bisexual individual. Nothing strictly analogous is known in the animal kingdom, but amongst plants, in the Linnean class, Polygamia, closely similar instances abound.

In the series of facts now given, we have one curious illustration more to the many already known, how gradually nature changes from one condition to the other,—in this case from bisexuality to unisexuality. Finally, in the four genera so often named, we meet the following several cases, some of them even the most diverse, occurring in closely allied species. (1st), a female, with a single male (rarely with two) permanently attached to her, protected by her, and capable of seizing, by the movements of its pedunculated body, any minute animals or substances found within her sack; (2d), a female with successive pairs of short-lived, mouthless males, inhabiting pouches on each side under her scutal valves; (3d), a female with many, in one instance fourteen, short-lived males, destitute of mouth, thorax, and appendages, but furnished with a stupendously long male organ, attached to a thickened portion of her outer integuments, but lying within the cavity which she has excavated; (4th), an hermaphrodite with a male attached within the sack, capable of feeding itself, as in the first case; (5th), an hermaphrodite with from one to three males, organised like ordinary Cirripedes, and apparently capable of seizing prey in the common way; and attached between the scuta, and thus protected; (6th and lastly), an hermaphrodite with from one or two up to five or six, short-lived, mouthless males, like those in the second case, attached in one particular spot, on each side of the orifice leading into the sack.

ORDER I.—THORACICA.

Cirripedia having a carapace, consisting either of a capitulum on a peduncle, or of an operculated shell with a basis. Body formed of six thoracic segments, generally furnished with six pairs of cirri; abdomen rudimentary, but often bearing caudal appendages; mouth with the labrum not capable of independent movements; larva firstly uniocular, with three pairs of legs, lastly, binocular, with six pairs of thoracic legs.

In the sketch of the three Orders given in the [Introduction], it will have been seen that the differences in their structure are so great, that it would have been hardly possible to have given a single blended account of the whole Class. But as all common Cirripedes are included in the present Order, here would have been the natural place for a full description of their external and internal structure. Having, however, been necessarily, yet perhaps unfortunately, led to give, in my former volume, a description of this kind of the Lepadidæ; and as it is necessary to give a similar account of the other great family of the Order, namely, the [Balanidæ], I have found it more convenient to make this latter account comparative and supplemental to the former one on the Lepadidæ, and so serve for the Order, rather than attempt to give a separate description in full of it. For this latter plan would have involved much useless repetition, as, on account of the many exceptions and limitations necessary to almost every statement, there is little choice between a description of great length and a mere diagnostic character of the Order, such as I have given above.

The [Thoracica] may be divided into three very natural Families, of nearly equal value; firstly, the [Balanidæ], or sessile Cirripedes, which may be subdivided into two sub-families, also very natural, the [Balaninæ] and [Chthamalinæ]; secondly, the [Verrucidæ], containing only one genus; and thirdly, the Lepadidæ, or pedunculated Cirripedes. These three families differ from each other, besides in mere external appearance, almost exclusively in the relation of the different portions of their external covering or carapace, and of the muscles moving such portions. In the [Balanidæ], the four opercular valves surrounding the orifice leading into the sack, are capable of other movements, besides being opened and shut; whereas all the other valves are immoveably united together. In the Lepadidæ, the valves answering to the opercular valves, are furnished with a muscle only for shutting them; whereas the peduncle answering to the basis in the [Balanidæ] is capable of various movements. In the [Verrucidæ] the shell is singularly asymmetrical; only half of the operculum (either the right or the left side, this varying even in the same species) being moveable; the other half being immoveably united to the remaining valves; and the whole shell has only one muscle serving to shut the moveable half of the operculum. All the internal parts and organs are very similar in the above three Families. If, however, the internal structure of one of the two sub-families, into which the [Balanidæ] may be divided, namely, of the [Balaninæ], be compared with that of the Lepadidæ, several important differences may be detected;—on the one hand, in the [Balaninæ], the presence of branchiæ, the extremely complicated cementing apparatus, the difference in structure between the third and succeeding pairs of cirri, the large palpi, the notched labrum, and the laterally double teeth of the mandibles;—and on the other hand, in the Lepadidæ, the presence of ovigerous fræna, caudal appendages, bullate labrum, and often prominent olfactory orifices. But if the Lepadidæ be compared in these several respects with the other sub-family, or [Chthamalinæ], which cannot possibly be removed out of the family of [Balanidæ], many of these differences break down and disappear, in some or all of the species.

The Lepadidæ include, as has previously been noticed, a much greater range of difference than the [Balanidæ]; and this is what might have been expected, for it is the most ancient family, and extinction has done its work, separating genera, which, in accordance to analogy, we may suppose were once more nearly connected by intermediate forms. The Lepadidæ, in one sense, may be taken as the type of their order; for they have undergone less “morphological differentiation;” that is, they differ the least from the last larval stage, and seem to give the most general idea of a Thoracic Cirripede. On the other hand, if we mean, as some authors do, by the word type, that form which, in the group in question, has been most modified, and illustrates every peculiarity of its class in the strongest manner, then we must look to the [Balaninæ], and to its typical genus, [Balanus], for the most Cirripedial form. In this genus the different portions of the carapace differ most, and subserve to a certain extent different ends, and in minute structure are most complicated; here the cementing apparatus, which offers the main characteristic of the whole sub-class, is most complex; here the several pairs of cirri differ most from each other in structure and action; here the peculiar branchiæ (organs apparently derived from the modification of another organ, itself confined to Cirripedes, viz., the ovigerous fræna) are best developed; here the nervous system is most highly concentrated; and, lastly, here we meet with the largest and most massive species of the whole group.

1. Family BALANIDÆ, (or Sessile Cirripedes).

Cirripedia without a peduncle; scuta and terga furnished with depressor muscles; other valves united immoveably together.

TABLE OF CONTENTS.

Almost every one who has walked over a rocky shore knows that a barnacle or acorn-shell is an irregular cone, formed generally of six compartments, with an orifice at the top, closed by a neatly-fitted, moveable lid, or operculum.[19] Within this shell the animal’s body is lodged; and through a slit in the lid, it has the power of protruding six pairs of articulated cirri or legs, and of securing by their means any prey brought by the waters within their reach. The basis is firmly cemented to the surface of attachment. The whole shell, basis, and operculum consists, as we have already seen, of the first three segments of the head, modified into a singularly constructed carapace, which encloses the mouth and rest of the body. The anterior extremity of the shell is situated in the centre of the basis, where indeed, by due care, the antennæ of the pupa may be always detected; and the posterior extremity is directed vertically upwards.

[19] The best published description of the structure of the shell of a sessile Cirripede, is given by Dr. Coldstream, in the ‘Encyclopædia of Anatomy and Physiology,’ article ‘Cirrhopoda.’

Structure of Shell.

When the shell of a sessile Cirripede or barnacle, for instance, of a [Balanus], is first examined, the structure appears extremely complicated; but this can hardly be considered as really the case. The structure will, I think, be best understood by recalling to mind that of Pollicipes,—the oldest known genus, from which, in one sense, all ordinary Cirripedes, both sessile and pedunculated, seem to radiate. I must premise, and the fact in itself deserves early notice, that the homologies of the several parts in the pedunculated and sessile Cirripedes admits of no doubt,—that is, if amongst the pedunculated, the genus Pollicipes, or certain species of Scalpellum, be taken as a standard of comparison.[20] The peduncle corresponds with the basis, as may be clearly seen, if a Pollicipes with a short peduncle, and a [Balanus], with a deep cup-formed or cylindrical basis be compared, for the contained parts are similar, and both grow at their upper edges upwards and outwards. Secondly, the valves round the lower part of the capitulum of a Pollicipes, though generally much more numerous, and forming more than one whorl or circle, and not so closely packed together, answer to the compartments forming the shell of a sessile Cirripede; this is shown by their lateral and downward growth, by their upper ends generally projecting freely above the cavity in which the animal’s body is lodged; and in the case Pollicipes mitella, by an actual resemblance in outline, some being triangular, some broad at the upper end, and some sub-rhomboidal, and, lastly, in the manner in which they slightly overlap and indent each other: moreover further resemblances in the relative position and even in the size of the several valves, will hereafter be pointed out between certain sessile genera amongst the [Chthamalinæ] and certain genera of the Lepadidæ. Thirdly, the scuta and terga in Pollicipes, so strikingly resemble in manner of growth in position relatively to the animal’s body—in shape—and even in being articulated together, the valves which form the operculum or lid of sessile Cirripedes, that their identity is at once obvious.

[20] Dr. J. E. Gray long ago observed these homologies. If Lepas be taken, the comparison is not quite so simple, owing to the growth of all the valves in that genus being upwards; but in several species of Scalpellum we may see the intermediate steps between the normal downward growth of the valves in Pollicipes, and the abnormal upward growth in Lepas.

It may be well here further to premise, that apparently none of the sutures in the shells of Cirripedes correspond with the articulations between the three archetype cephalic segments, of which the whole shell is formed; or with the eight elemental pieces, of which each separate segment in the archetype crustacean is known to consist. But, as I believe, the several valves in the shell of a Cirripede are homologous, or at least analogous, with the sclerodermic plates,[21] of which the carapace of the Podophthalmia is formed; with this difference, that in the latter they become, after their first formation, united together into a single piece, and are thus moulted as a whole; whereas in Cirripedes, the valves or sclerodermic plates are not moulted, but continue to be added to throughout life.

[21] Milne Edwards, ‘Annales des Sciences Naturelles,’ tom. xviii, (1852), p. 236.

In Pollicipes, there is no difficulty in understanding the growth of the lower valves of the capitulum, especially if a species be taken in which these valves stand a little way apart: at each period of growth, they are added to at their basal edges and a little way up both sides; at the same time, a new membrane connecting them together is formed, the old membrane disintegrating, or being left hanging in tatters to the last zone of growth. Now if we look at the shell of a sessile Cirripede, there is no essential difference in the growth of the compartments or valves; all grow downwards and laterally; but they overlap each other much more laterally than in Pollicipes, and the connecting membrane is in most parts reduced to a mere film jammed in between the valves; but, in the case of the opercular membrane, it still remains wide, and is periodically moulted.

In the annexed woodcut (fig. [1]), of the rostrum of [Balanus Hameri], the downward growth and the lateral growth on both sides is plain. The modified sides (rr) for convenience sake, have been called the radii; they invariably overlap the adjoining compartments. The middle part (p), has been called the wall, or parietal portion: in the specimen figured, the walls and radii are distinctly separated, but in some cases, especially amongst the [Chthamalinæ], the lines of growth are absolutely continuous from one to the other. In fig. [2] of a Lateral compartment of the same Balanus, we have the same essential structure; but the left side (a) is more protuberant, and is hollowed out in its lower half; it is, also, more distinctly separated from the parietal portion: this side has for convenience been called the ala; it is invariably overlapped by the adjoining compartment: in some few cases, as in [Pachylasma], the ala is not hollowed out in its lower part, and from being added to in a straight line along its whole edge, with the lines of growth continuous with those on the wall, it differs hardly at all in appearance from a radius. Lastly, in fig. [3] of the carina, or compartment facing the rostrum, we have alæ (aa) on both sides; these being, as in all cases, overlapped by the adjoining compartments.

Fig. 1.

Fig. 2.

Fig. 3.

p, p, Parietes; r, r, Radii; a, a, Alæ.

Fig. 1, Rostrum with two radii, serving in the [Chthamalinæ] for rostro-lateral compartments.

Fig. 2, always serving for lateral and carino-lateral compartments.

Fig. 3, Carina, serving in the [Chthamalinæ], also, as a rostrum.

Now, the compartments in the shell of every sessile Cirripede, are without exception constructed on the above three simple patterns. In number, they are 8, 6 or 4, or all confluent together.

Considering this simplicity in growth and form of the separated compartments, it seems at first surprising that the construction and enlargement of the whole shell in [Balanus], should long have been viewed as a difficulty. But the radii, from growing against rectangular indentations, or rather furrows, in the opposed compartments, come to be set a little inwards; and their external surfaces assume a very different appearance from the wall-portions of the compartments, which grow against the surface of attachment. In different species, the summits of the radii (and of the alæ) grow either very much more obliquely than in the species figured, or more squarely—that is, they extend from tip to tip of the adjoining compartments, parallel to the basis. In this latter case, and when the surfaces of the radii differ considerably in appearance from the walls, as in [Balanus tintinnabulum] (Plate [1]), I am not at all surprised that the radii should have been described as separate elements, and called “areæ interjectæ,” or “compartments of the second order:” for the shell of this Balanus seems to be composed of six wedges with their points upwards, namely, the parietal portions of the compartments, and of six other narrower wedges, the radii, with their points downwards; and the fact that these latter wedges consist simply of the sides of the parietal portions, modified by growing against the adjoining compartments, is completely masked. I should add, that sometimes the radii are not developed, which simply means that the overlapping lateral edges of the compartments have not been added to during growth.

The alæ are originally developed at the period of the metamorphosis, as slight lateral protuberances in the upper part of the compartments; from being overlapped, and therefore not exposed to external influences, and from growing (as in the case of the radii) against rectangular indentations or furrows in the adjoining compartments, they generally assume an extremely different appearance from the parietes, and might naturally be thought to have a very different nature. But the alæ in all cases (as is obvious in [Pachylasma]) are nothing but the protuberant lateral edges of the compartments, rendered thin and modified during growth. In order that the margins of the alæ should be received in an indentation, the upper internal surfaces of the walls of the recipient compartments are thickened all round, excepting where they receive the alæ. This thickened, upper, internal portion of the walls or shell, together with the alæ themselves, form the part called the sheath. The sheath sometimes blends insensibly into the lower parts of the compartments, and then perhaps it would not be thought to be a distinct element; but often it is abruptly separated by an overhanging edge (see Pl. [9], fig. [5 b], [9 b]; Pl. [20], fig. [1 b]; Pl. [25], fig. [1], K′) from the lower part, and then the sheath greatly complicates the internal appearance, but not the essential structure of the shell. The sheath acts beautifully, like an internal hoop, in strengthening the shell round the orifice, where it is naturally weaker than at the lower or basal end, where it adheres to the surface of attachment: in the upper part of the shell, moreover, the sutures between the compartments do not go straight through, but owing to the alæ projecting and being overlapped, are extremely oblique; or the joints, in the language of carpenters, may be said to be broken.

There is one other point of structure in the shells of the [Balanidæ], more especially of species like [Balanus tintinnabulum], which adds to their apparent complexity, namely, that the rim or orifice of the shell formed by the upper ends of the compartments, projects considerably above the opercular valves. In a young [Balanus], immediately after the metamorphosis, the operculum is attached by the opercular membrane all round to the summits of the compartments, and there cannot be said to be any orifice to the shell itself, but only an orifice or slit between the opercular valves; but during growth, as the compartments are added to at their basal edges, their upper ends are deserted, and cease to enclose the sack, within which lies the animal’s body. Hence the upper ends come to project freely, either quite separately as in some species of Pollicipes, where they cannot be said to form an orifice; or more or less united into a ring so as to form an orifice, as in the different species of [Balanidæ]. It follows, that to understand the real shape of a [Balanus], or rather of the cavity enclosing the animal’s body, all that part of the shell which projects above the opercular membrane, may, in imagination, be removed as something extraneous, like so many spines; not that I mean to say that these points of shell are dead; on the contrary, they are often porose and penetrated by numerous threads of corium. This upper part of the shell, thus produced so as to form an orifice, no doubt serves to protect the less strong and moveable operculum.

Fig. 4.
Octomeris.

Fig. 5.
Chthamalus.

Fig. 6.
Chamæsipho.

Fig. 7.
Balanus.

Fig. 8.
Tetraclita.

a, Rostrum; b, Rostro-lateral, c, Lateral, d, Carino-lateral compartment; e, Carina.

Horizontal sections through the Shells of the principal genera of [Balanidæ], showing the arrangement of the Compartments. Genera 4, 5, and 6 belong to the [Chthamalinæ]; 7 and 8 to the [Balaninæ].

Number and Arrangement of the Compartments.—I have already stated that the shell, in every one of the [Balanidæ], consists of eight, six, or four compartments, or of all fused together into a single piece; and that the compartments themselves are all constructed on the three simple patterns of which woodcuts (figs. [1], [2], [3]) have been given. They are arranged in a certain definite order. The type arrangement is found amongst the [Chthamalinæ], as might have been expected, inasmuch as this sub-family is so closely related to the ancient genera Pollicipes and Scalpellum, whence all the Thoracic Cirripedia may be said to radiate. In [Octomeris] (fig. [4]) the type-arrangement of the compartments, eight in number, is well shown; the rostrum and carina resemble each other, and have alæ on both sides, and therefore are overlapped on both sides: the rostro-lateral compartments have radii on both sides, and therefore overlap the adjoining compartments on both sides; the lateral and carino-lateral compartments have radii on their carinal, and alæ on their rostral sides; and therefore overlap on one side, and are overlapped on the other side. Now the shell of every other sessile Cirripede differs, I believe, from that of [Octomeris], only in the fusion together or abortion of some of the eight normal compartments: in one genus, however, [Catophragmus], outer whorls of small compartments, arranged like the lower valves in the capitulum of Pollicipes, are superadded. The genus [Chthamalus] (fig. [5]) differs from [Octomeris] only in the carino-lateral compartments being aborted, (as will presently be discussed), and hence has six compartments. [Chamæsipho] (fig. [6]) differs from [Chthamalus] only in the rostro-lateral and lateral compartments being fused together; and hence has only four compartments. In [Balanus] (fig. [7]) and the whole sub-family of the [Balaninæ], the rostrum is compounded of the true rostrum, as seen in the type [Octomeris], and of the two rostro-lateral compartments; hence this compounded rostrum has radii instead of alæ on both sides, and there are only six compartments. [Tetraclita] (fig. [8]) and [Elminius] differ from [Balanus] only in having the carino-lateral compartments absent, and probably aborted; hence there are only four compartments. Lastly, in [Pyrgoma], all the compartments are blended together into a single piece.

In Pollicipes, the old type-form of the whole order, and in Scalpellum, we have four valves, (answering to the operculum), surrounding the aperture leading into the sack, and the valves below are arranged in successive whorls, with a strong tendency to alternation. For, the rostrum alternates with, that is faces the interval between, the two scuta; the carina alternates with the two terga; and the upper lateral valves alternate with the scutum and tergum on each side. These four valves, namely, the carina and rostrum, which resemble each other in structure, and the pair of upper latera, which are larger than the other lateral valves, together form the uppermost whorl, or that beneath the scuta and terga. In the next whorl we have the rostro- and carino-lateral valves, alternating with those above them; and beneath them there are generally other valves, which decrease in size and display the same tendency to alternation. The valves here just specified, namely, the rostrum, carina, and three pairs of lateral valves, in the Lepadidæ, are so much larger, and are so much more commonly present, than the other valves of the capitulum, that to them alone I affixed special names. Now if amongst sessile Cirripedes we look to that genus, viz., [Catophragmus], which comes in its whole structure the nearest to Pollicipes, one of the Lepadidæ, we find (as in fig. [4]), firstly, a rostrum and carina resembling each other, and a pair of lateral compartments, larger than the other lateral pairs; these four valves alternating with the opercular valves: and, secondly, we find, but forming part of the same whorl, a pair of rostro-lateral and a pair of carino-lateral compartments, which, just as in Pollicipes, are larger than the exterior and lower valves. These lower little valves, I may remark, decrease in size in the successive whorls, and tend to alternate in position, just as in Pollicipes. Observing these several striking points of correspondence in the valves, (and indeed in the whole structure), of [Catophragmus] and Pollicipes, one is strongly inclined to suspect that in [Catophragmus], and therefore in [Octomeris] and other sessile Cirripedes, although the rostro- and carino-lateral compartments appear to lie in the same whorl with the rostrum, carina, and large lateral compartments, yet that they really belong, as in Pollicipes and Scalpellum, to a lower whorl. Now if a very young shell of [Balanus], immediately after the metamorphosis, be examined, the carino-lateral compartments will be found not to have been developed; they first appear after two or three zones of growth have been added to the other compartments; bearing in mind that in Pollicipes and in [Catophragmus] the lower whorls are added successively during growth, we find in this fact strong confirmation of the view that the carino-lateral compartments normally belong to a whorl beneath that including the rostrum, carina, and lateral compartments. Whether the rostro-lateral, like the carino-lateral compartments, are developed subsequently to the others, I have had no opportunity of ascertaining, and therefore cannot confirm the above analogical conclusion, namely, that they, also, belong to a lower whorl.

In the sub-family [Balaninæ], which includes [Balanus] (woodcut [7]), and [Tetraclita] (woodcut [8]), the shell is characterised by not having rostro-lateral compartments, and by the rostrum having radii: now in [Pachylasma giganteum], which undoubtedly belongs to the sub-family [Chthamalinæ], at a very early age the rostro-lateral compartments become blended with the true rostrum, making a compound rostrum, exactly like the rostrum in the [Balaninæ]; distinct evidence of a similar fusion is retained throughout life (Pl. [15], fig. [1]) in all three species of [Chelonobia], which is undoubtedly a member of the [Balaninæ]. Hence, I think, we may conclude that in all the genera of the [Balaninæ] the rostro-lateral compartments are probably not aborted, but are blended with a normal rostrum (resembling that in woodcuts [4], [5], [6]), making together a compound rostrum furnished with radii: it must, however, be observed that I could not detect any actual evidence of this fusion in [Balanus], even immediately after the metamorphosis. In [Chamæsipho] (woodcut [6]), either the rostro-lateral compartments attain a most unusual breadth, or, as is more probable, they have become confluent with the lateral compartments, which in the Lepadidæ seem to be the most persistent of all the lateral valves. In such genera as [Tetraclita] and [Chthamalus], in which the carino-lateral compartments are absent, they may be fused with the lateral compartments or with the carina; but seeing that they are normally developed later than the other valves, it appears to be the simplest theory to assume, until the contrary be proved, that they are aborted. Finally, the somewhat unexpected conclusion that the shell (not including the operculum) of sessile Cirripedes normally consists of eight valves, four belonging to an upper whorl, and four to a lower whorl, all forced into a single ring, and often more or less fused together, though not strictly proved, is rendered highly probable. I will only further add, that the Basis perhaps represents several whorls of the small valves or scales on the peduncle of Pollicipes, fused together; the comparison of the basis with the calcareous cup, forming the lowest portion of the peduncle in Lithotrya, which has been made by some authors, I do not think is very accurate, as the cup in Lithotrya seems to have a special relation to the boring habits of that genus.

Fig. 9.

Basal edge of wall of compartment in [Balanus tintinnabulum]; a, a, outer lamina; b, b, inner lamina; c, c, longitudinal septa uniting the inner and outer laminæ with their ends denticulated.

Structure of the Individual Compartments.

If the basal margin of a compartment, for instance, of [Balanus tintinnabulum], be examined, it appears sufficiently complicated, being composed of an outer and inner lamina, separated by longitudinal septa, which are denticulated at their bases; and the tubes formed by these longitudinal septa are crossed by transverse septa. On the other hand, in some cases, as in the genera [Chthamalus] and [Elminius], each compartment consists of a simple shelly layer. These two extreme states graduate into each other: we have, firstly, on the internal surface, quite irregular points and ridges; these become regular, causing the internal surface to be longitudinally ribbed; then these ribs themselves become finely furrowed on their sides and at their lower ends, producing sharp, minute ridges, the ends of which I have called the denticuli; and, lastly, some of the denticuli on the adjoining longitudinal septa become united into a solid layer, forming the internal lamina of the wall. These denticuli do not generally cover the whole surface of the longitudinal ribs, but leave a portion near the outer lamina of the compartment smooth. The denticulated ends of the longitudinal septa project beyond the basal edge of both the outer and inner laminæ, and enter the mouths of the tubes (where such occur) in the basis, and thus strengthen the shell. The whole of the internal lamina generally is more or less striated longitudinally, thus displaying its origin from the union of the inner edges of the longitudinal septa. I need only further remark that on the internal surface of the outer lamina, between the main longitudinal septa, there are generally (as in the woodcut) smaller longitudinal ridges, which do not reach the inner lamina, and on this account alone are not called septa.

Tubes are formed by the longitudinal septa, between the outer and inner laminæ. These tubes are almost square, and are occupied by threads of corium, which enter at pores left open between the edge of the compartment and that of the basis on which it rests. The tubes extend high up the compartments; but in the uppermost part they are generally cut off by thin, transverse, calcareous septa, deposited by the ends of the threads of corium; a cancellated structure being thus produced. Or the uppermost part of each tube becomes filled up solidly with compact shelly layers, which are always first thrown down on the side of the tube facing the outside, and thus greatly strengthen the shell: in several instances, as in [Balanus perforatus] and [Tetraclita porosa], in which the disintegration of the upper part of the shell is a necessary element in its growth for the enlargement of the orifice, these filled up tubes become exposed. In [Coronula] and [Tubicinella], the tubes in their upper parts are, I believe, crossed only by transverse membranous septa.

Anomalies and exceptions.—In [Tetraclita] (Pl. [10], fig. [1 g], [1 h]) from the branching of the longitudinal septa, several irregular rows of tubes are formed. In certain varieties of [Balanus balanoides] (Pl. [7], fig. [2 b]), and in [B. cariosus] (Pl. [7], fig. [3 b]), slight branching ridges on the internal surface of the walls, seem to answer to the longitudinal septa, and produce, during the downward growth of the shell, extremely irregular cells, and short tubes. In [Balanus vinaceus] (Pl. [2], fig. [7 d]), the internal lamina, instead of being solid, as in every other species, is left cancellated, and thus betrays, much more plainly than usual, its origin in the united denticuli of the adjoining longitudinal septa. In [Balanus porcatus], between the main longitudinal septa, there are (Pl. [6], fig. [4 e]) what may be called rudimentary and disconnected longitudinal septa. In [Coronula] and its allies (Pl. [16], fig. [6], and Pl. [17], fig. [4 c]) it is the outer lamina of the compartment which is anomalous; for in the two or three lower zones of growth, it forms only a ledge on each side of the longitudinal septa; which ledges, higher up, become confluent, and so form an ordinary outer lamina. In [Coronula], also, the wall of each compartment (see transverse section, Pl. [16], figs. [5], [7]) is very remarkable from being deeply folded, the folds being on their internal faces firmly calcified together, and on their external faces closely pressed together (often with a neatly serrated suture), so that the whole nature of the shell might be, as has happened, easily quite misunderstood; and the walls be considered as very thick, instead of being, as is really the case, very thin. In [Chelonobia] (Pl. [15], fig. [1]), however, the walls are truly of such great thickness, that the nature of the relative parts might likewise be misunderstood; in this genus the ovarian tubes enter the walls, extending up between the longitudinal septa, or, as they may here be more naturally called, the radiating septa. I will specify a few more peculiarities worthy of remark:—in some species of the sub-genus [Acasta], clefts are left, covered only by membrane, on the lines of suture (Pl. [9], figs. [7 a], [8 a]), between the compartments, just above the basis; and in other species the basis is perforated by numerous membrane-covered, minute orifices. In [Platylepas], each compartment has one deep inward fold (Pl. [17], [fig. 1]), somewhat analogous to the three folds in [Coronula]; this fold is produced into an internal midrib, supporting and rendering convex the membranous basis; in this genus, also, the rostrum, owing to its midrib, is generally thrust a little on one side, and the shell thus rendered asymmetrical. In [Chamæsipho scutelliformis] the shell is symmetrically perforated (Pl. [19], fig. [4 a]) by four apertures. Lastly, in [Chthamalus Hembeli] and [intertextus], after a certain age, the basal edges of the walls become inflected, and continue to grow inwards till they entirely take the place of the true membranous basis.

Structure of the Radii.

Radius.—This term, as we have seen, is applied to that side of the compartment, the growth of which is modified, by abutting against and overlapping the adjoining compartment. Hence the structure of the radius is essentially the same with that of the parietal portion of the compartment. When best developed, as in [Balanus tintinnabulum], the radius consists of an outer and inner lamina, separated by denticulated septa, extending in horizontal lines parallel to the basis, and is consequently perforated by minute tubes or pores. The tubes become filled up solidly much more commonly than do the parietal tubes; and the inner lamina, in such cases, is hardly distinct from the outer lamina. The denticuli often fail, or are present only on the lower sides of the septa; and very frequently the edge of the radius can only be said to be crenated. Notwithstanding these frequent anomalies, if a series of species and genera be taken, it is certain that there is, as might have been expected, a close relationship in internal structure, between the radii and the parietes. The edge of the radius is received in a slight furrow (generally marked like a seal, with the impression of the denticulated septa) in the opposed compartment: sometimes the outer edge or lid of the recipient furrow, is so broad as to give the false appearance of a radius having been developed, at least in the lower part of the shell, on both sides of the suture. A crest of corium runs into each suture between the edge of the radius and the furrow in the opposed compartment; and when the radius is permeated by pores (as in woodcut [10]), threads of corium branch off this crest, and enter the pores. In the lower part of the shell, these crests of corium project from the corium forming and surrounding the sack; but in the upper part of the shell, above the opercular membrane, and therefore above the sack, the corium is produced up each line of suture as a separate ribbon. In proportion as these ribbons extend more or less near to the summit of the shell, so do the edges of the radii continue to be added to, to a greater or less height from the basis; and consequently their summits become less or more oblique.

Fig. 10.

Edge of the radius of [Balanus tintinnabulum]. a, outer lamina; b, inner lamina; c, denticulated septa, uniting the two laminæ.

Peculiarities in the Structure of the Radii.—In some of the species of [Tetraclita], in which genus the walls consist of several rows of tubes, the radii are likewise perforated by several rows; and in some of the other species (Pl. [10], fig. [1 h]), the edge, or disarticulated surface of the radius, is marked by irregularly branching ridges; and these evidently correspond with the branching septa or ridges of the wall. In [Chelonobia], the outer lamina of the radius, as well as of its recipient furrow, is of extraordinary thickness; and this lamina, in [C. testudinaria] (Pl. [14], fig. [1 a], [5], b, and Pl. [15], fig. [1], f), is modelled into sharp transverse ridges and valleys. In the [Chthamalinæ], the radii, like the parietes, are simply solid; and apparently in consequence, for the sake of strengthening the sutures, the edges of the radii, and of the recipient furrow in [Octomeris] (Pl. [20], fig. [3 a]) and in [Chthamalus dentatus] and [Hembeli] (Pl. [18], fig. [3 b], [5 a]), are neatly dentated. In some other species of [Chthamalus] (Pl. [19], fig. [1 a]), the radii present a slight modification of this structure, the sutures being formed by oblique interfolding laminæ. In the radii of [Coronula] and [Tubicinella], there is a peculiarity, in apparent connection with the fact, that in these genera the parietal tubes are not crossed by transverse calcareous septa, namely, that the pores by which the radii are permeated keep unclosed throughout their length, and open into a special longitudinal tube (Pl. [16], fig. [7], d′), which runs along that margin of the wall, whence the radius arises. In [Coronula] the wall is of extreme thinness, and in conformity so is the true radius, but that the shell might not thus be rendered very weak, complementary or pseudo-radii are developed on their inner sides (Pl. [16], fig. [7], adjoining the true radii A d, C d, and shaded by distant convex lines). Even in the allied genus [Xenobalanus], in which the whole shell tends to become rudimentary, traces of these pseudo-radii (Pl. [17], fig. [4 b], d) can be detected. In [Coronula], though the radii (Pl. [16], fig. [7], A d, C d) are, by the above special means, rendered thick, and though the alæ also are thick (C a′, D a′), yet together they do not equal in thickness the folded walls; and consequently, there is left between the radii and alæ square chambers (v), occupied by the branching ovarian tubes.

Structure of the Alæ.

These project, generally abruptly, from the sides of the upper part of the compartments; they appear from the first growth of the shell; they are overlapped by the radius and by part of the wall of the adjoining compartment; they are thinner, and have, owing apparently to being overlapped, a very different aspect from the parietal portion; but they do not differ from it in essential nature. They are solid, that is, they are never permeated by pores; but their edges are generally crenated, and there is, in some cases, as in [Chelonobia], sufficient evidence that these crenations answer to the horizontal septa on the edges of the radii (also often reduced to mere crenations), and consequently, likewise, to the longitudinal septa of the parietes. In [Coronula] the edge of each ala consists of a medial ridge, sending off denticulated septa on both sides, and is therefore anomalous as compared with the alæ in other genera, but corresponds in structure with the similarly anomalous radius of [Coronula]. In order to allow of the growth of the edge of the ala, a fine thread of corium runs up the narrow furrow in which the edge is lodged, proceeding from the corium of the sack. In proportion as this thread runs up higher or lower, so are the summits of the alæ rendered, during growth, less or more oblique.

Structure of the Sheath.

As the compartments overlap each other, the edges of the alæ would have projected, and the inner surface of the orifice of the shell would not have been smooth and rounded, had not that part of each wall, which does not overlie an ala, been thickened so as to allow of the formation of a shoulder or indentation, against which the edge of the ala fits and abuts. The thickened portions, and the alæ themselves, together form the sheath, of which the use seems to be to strengthen, like a broad internal hoop, the upper part of the shell round the orifice, where naturally it is weak. The sheath is composed of successive, fine, shelly layers, which extend, as the shell is added to at the basal margin, lower and lower down on the inner surface of the walls. The lower edge of the sheath either simply projects a little inwards, or more commonly is formed into a sharp depending ridge, as represented in fig. [1], K′, Pl. [25]. In some species of [Pyrgoma] (Pl. [13], fig. [2 b]), the sheath reaches nearly to the bases of the compartments; and in [Chelonobia] (Pl. [14], fig. [4] e c e), the inner layer of shell surrounding the sack, which seems to correspond more nearly to the sheath than to the inner lamina of the walls, actually rests on the basal membrane. The opercular membrane is generally, but not invariably, attached only a little way above the lower edge of the sheath: at each exuviation, a new opercular membrane is formed, and is attached to the next lower zone of the sheath; the old membrane being cast off, but a circular slip of it is left, investing the last zone. Hence the whole upper part of the sheath above the opercular membrane, comes to be thus invested; and is marked by circular lines, one above the other, caused by the successive exuviations. This investing membrane often supports rows of minute bristles, directed upwards. Generally, a film of shell is deposited, at the period of the formation of each new opercular membrane, on that part of the sheath which lies immediately beneath. This innermost film or thin layer of shell, on the lines of suture between the compartments, breaks joint, at least in some cases, with the underlying shelly layers,—that is, the suture in this last-formed film does not lie exactly over the suture in the subjacent layers of the sheath. In [Tubicinella], the sheath extends down close to the basis; and what is unique in this one genus, the opercular membrane, gradually thinning out downwards, closely adheres to the whole inner surface of the shell. In [Tubicinella] and in [Xenobalanus] (Pl. [17], fig. [4 b]), the sheath separates easily into separate successive rims of shell; and this structure evidently is for the sake of facilitating the breakage of the upper end of the shell, which, as we shall presently see, is necessary to allow of the increase in size of its orifice.

Structure of the Basis.

This, in several genera and species, is composed of simple membrane, and consists of successive, concentric, circular slips, added round the outside, at each period of growth. In some species of [Tetraclita] and [Balanus] the basis is calcareous, but diaphanous, very thin, smooth, or somewhat granulated. In other cases it consists of a single calcareous lamina, either smooth, or with ridges radiating from its centre; it is formed of two laminæ, (as is most usual in [Balanus],) separated by radiating septa. These septa, as well as the radiating ridges in the case of the single lamina, are homologous with the longitudinal septa of the parietes. The denticulated ends of the latter enter the mouths of the tubes formed by the radiating septa of the basis: threads of corium pass between the denticuli of the parietal septa, and thus enter the basal tubes. The ends of these threads of corium generally deposit transverse calcareous septa, exactly as within the parietal tubes. When the basis is thick the septa themselves (ccc) between the proper basal tubes, become porose, (or rather cancellated,) and they sometimes expand into a very thick, cancellated layer, separating the outer lamina (a) of the basis from the proper basal tubes, which always lie close under the inner lamina (b). This structure differs only slightly from that seen in the parietes of [Tetraclita], in which the branching of the longitudinal parietal septa, produces thick walls, formed of several rows of tubes or pores. With respect to peculiarities in structure of the basis, [Balanus lævis] offers the most remarkable case; for here, in specimens which have grown crowded together, the whole interior appears sometimes to have become too much elongated or too deep for the animal’s body, and consequently the lower part of the deeply-concave basis has been filled up (Pl. [4], fig. [2 a]) by thin, irregular, calcareous diaphragms. In elongated specimens, also, of [Balanus balanoides], the shell sometimes appears to have grown too long for the animal’s body; but in this case the membranous basis becomes extremely convex inwards; it still reaches the basal edges of the parietes all round, but in the middle it is raised high above the surface of attachment; yet sometimes threads of the cementing tissue depend from the middle part to the surface of attachment. In [Balanus terebratus] (Pl. [8], fig. [2 a], [2 b]), and in some species of [Acasta], the basis is riddled, as previously stated, by numerous, minute, membrane-covered orifices. In [B. declivis] the membranous basis is always extremely oblique, owing to the rostral end of the shell being twice as high as the carinal and opposite end.

Fig. 11.

Portion of edge of basis of [Balanus tintinnabulum], a, a, outer lamina; b, b, inner or upper lamina; c, c, c, porose or cancellated radiating septa.

Regarding the very remarkable means by which the basis of sessile Cirripedes is cemented to the surface of attachment, it will be convenient to defer for a little the description, on account of its necessary length.

Structure of the Opercular Valves (Scuta and Terga).

These are situated on each side of the slit or orifice leading into the sack; from their shape, their powers of movement, their separation by flexible membrane from the shell, to which they serve as a lid, they appear at first as if they constituted an element very distinct from the shell itself, but this is not the case. They are, together with the opercular membrane, as essentially as the whole of what is externally visible, a part of the modified carapace, of which they occupy the upper or posterior extremity: from tracing the metamorphoses, or even by comparison of a [Balanus] with Pollicipes, there can be no doubt of the truth of this conclusion. The opercular valves are four in number,—a pair of scuta and a pair of terga; but the latter in [Coronula diadema] and [reginæ], are either aborted or represented by a mere rudiment; and in [Xenobalanus] both scuta and terga are quite absent. In several cases, more especially in the genus [Pyrgoma] (Pl. [13], fig. [1 b]), the scutum and tergum on each side are calcified together, so that sometimes not even a trace of the line of junction can be discovered. In most cases the scutum is firmly united, being articulated in a manner presently to be described, to the tergum; but in [Coronula], [Tubicinella] (Pl. [17], fig. [3 c]), and [Platylepas], the ends of these valves are simply approximated.

Scuta.—These valves are important, inasmuch as the animal’s body is attached to them; in Pl. [25], fig. [1], the broken line, surrounding a, b, shows where the body has been cut, in removing the scutum on the near side, the other scutum, S, being left articulated to the tergum, T. In shape the scuta are generally sub-triangular; but in some species of [Pyrgoma] and in [Chelonobia], &c. they are much elongated. The lines of growth are usually prominent; and along the occludent margins the alternate, or sometimes every third or fourth line, is developed into a knob, which produces a serrated edge, serving to lock the two opposed valves together; there is, however, no trace of this structure in [Coronula] and [Tubicinella]. In some species of [Pyrgoma], a ledge of considerable breadth (Pl. [13], fig. [3 e], &c.) is developed along the occludent margins of the two scuta, as well as of the two terga, giving them an anomalous structure. The Terga differ considerably in outline in the different genera and species: their shape approaches more nearly to a triangle than to any other regular form; but there is generally a projection or spur on the basal margin, on the side towards the scutum. In some species of [Pyrgoma], the tergum is of so irregular a shape as to defy description. In most cases, a longitudinal depression or furrow runs down the valve, from the apex to the extremity of the spur; and it not rarely happens that the sides of this furrow become folded inwards and almost closed. The spur probably answers to the basal point of the usually sub-rhomboidal tergum in Pollicipes and Scalpellum.[22] The tips of the terga in some species of [Balanus], &c., are specially modified into sharp points or beaks (Pl. [2], fig. [3 b], [3 d]), bowed a little inwards, and projecting considerably above the tips of the scuta; this is effected by the medial, uppermost part of the valve being internally thickened and hardened, and then, by the disintegration of the two margins and the external surface, the internal modified portion becomes exposed. The whole valve, also, at least in such cases as in [Balanus psittacus], appears to be forced slowly upwards in the articular furrow of the scutum. I am assured, by a competent observer, that the beaks of the terga in [B. porcatus] can give an object placed within the orifice of the shell a sharp tap.

[22] In comparing the Tergum of one of the [Balanidæ] with that of a typical member of the Lepadidæ, for instance, that of [Balanus] with that of Pollicipes, apex corresponds with apex: the extremity of the spur in [Balanus] corresponds with the basal point of the whole valve in Pollicipes: the scutal margin, (which in [Balanus] homologically extends down to the extremity of the spur), corresponds with the scutal margin of Pollicipes: the carinal margin in [Balanus] corresponds with the upper carinal margin in Pollicipes: the basal margin of [Balanus] on the carinal side of the spur, corresponds with the lower carinal margin in Pollicipes: lastly, (and this is the chief difference), in [Balanus] there is no appreciable occludent margin, the apex of the valve being brought close to the upper angle of the scutal margin; in [Chthamalus], however, there is yet left some remnant of an occludent margin,—which margin in Pollicipes is conspicuous.

The scutum and tergum, with the few exceptions above stated, are articulated together at a large or open angle. The articulation (see Pl. [11], fig. [5 b], [c], [d], and fig. [6 b], [c]) is effected by the margin of the tergum being a little inflected, and lodged in a furrow in the margin of the scutum. This furrow in the scutum has its further border generally prominent and often reflexed or curved over; I have called it the articular ridge; it, also, is lodged in a furrow in the upper part of the tergum, which again is bordered by a ridge, viz., the articular tergal ridge. So that in both scutum and tergum there is an articular furrow, bordered in each case, on one side by the margin of the valve, and on the other side by the so-called articular ridge. In [Chelonobia] (Pl. [14], fig. [1 b]) the articular ridge of the scutum is horny. When, as often happens, the scuta and terga have been much worn, the manner of their articulation (Pl. [18], fig. [1 a]) is pretty well shown even from the outside; in this case their external appearance is very different from what it is in those individuals (fig. [1 c]) of the same species, which have not suffered disintegration. This articulation of the scuta and terga is prefigured amongst the Lepadidæ, in Pollicipes mitella, and in Lithotrya.

The scuta are brought together by a short, strong, straight, adductor muscle (Pl. [25], fig. [1, a]); its attachment leaves (with very few exceptions, as in [Tubicinella]) a rounded impression, or even pit, on the under side of the valve in its upper part. This pit is frequently bounded, on its lower side, by a sharp ridge, which, though not in actual connexion with the adductor muscle, I have, for convenience sake, called the adductor ridge; it serves apparently to give support to the animal’s body; in some few cases (as in [B. psittacus], Pl. [2], fig. [3 c]) it is confluent at its upper end with the articular ridge, and converts the whole basi-tergal corner of the valve into a deep cavity. In some of the species of [Pyrgoma] (Pl. [12], fig. [5 c], [7 b]), and in some varieties of [Creusia], this adductor ridge is enormously developed, so as to depend far beneath the true basal margin, or that to which the opercular membrane is attached. At the basi-tergal corner of the valve, there is generally a small pit or impression, and sometimes distinct crests, for the attachment of the lateral depressor muscle. At the rostral end there is, also, a small cavity formed by the overfolding of the occludent margin (rarely furnished with crests) for the attachment of the rostral depressor muscle. In the Terga, at the basi-carinal corner, there are usually crests, though sometimes feebly developed, for the attachment of the tergal depressor muscle. But in [Chelonobia], [Coronula], [Tubicinella], [Platylepas], and in some other cases, there are no crests. The crests, when well developed, are furnished with rectangular sub-crests or denticuli on both sides; in fact they resemble, and are probably homologous with, the denticulated ribs or septa in the parietes, radii, and basis. Altogether the scuta and terga are attached, as far as muscles are concerned, to the shell and sack, by three longitudinal pairs.

Growth of the Whole Shell, and Its Microscopical Structure.

The opercular valves are added to along their basal margins alone;[23] the animal’s body, together with the several muscles, becoming attached at each period of growth lower and lower down to the valves; this no doubt is effected by the absorption of the upper surfaces of the muscles, and the formation of new fasciæ on their lower surfaces. The opercular membrane, which, though thin and flexible, forms part of the general outer surface of the animal as much as does any portion of the rigid shell, with which indeed it is strictly homologous, is periodically moulted, together with the integuments of the whole included animal. The new opercular membrane is of course each time formed a little larger than the old one. In [Coronula] and [Tubicinella], however, several successive opercular membranes are preserved one over the other, and the outside membrane gradually disintegrates; in these cases the undermost opercular membrane is formed wrinkled and considerably too large, so as to allow of being stretched, before it is finally cast off. In [Tubicinella], the opercular membrane runs down, adhering to the inner surface of the shell, to nearly the basis, and hence during the diametric growth of the shell, it is longitudinally split, and is repaired by slips of new membrane, which resemble the radii in form and in direction of the lines of growth.

[23] In some species of [Pyrgoma], the ledge (limbus occludens) which is added along the occludent margin of both scuta and terga, and in some species of [Balanus] a narrow rim, or slight protuberance which is added along the carinal margin of the terga, offer unimportant exceptions to the rule, that the opercular valves grow only at their basal margins.

The basis is added to only round the circumference, and hence increases in diameter, and, when concave, in depth. The compartments grow at their basal margins, where they are in contact with the basis; hence the shell is added to in height, and, owing to the outward inclination of the compartments, also, in basal diameter; but the compartments likewise, in most cases, grow along both lateral margins, that is, on the edges of the radii and alæ; and hence the upper part of the shell, also, increases in diameter. The orifice of the shell, moreover, thus becomes enlarged. In some cases the shell is destitute of radii, only sutures being present, that is, the compartments do not grow laterally; and sometimes, as in the whole genus [Pyrgoma], there are not even sutures, the compartments having been fused together: in both these cases, the shell can increase in diameter only at the base; and the orifice, it might have been thought, would necessarily have remained, to the destruction of the animal, of the same minute size, as when first formed after the metamorphosis: this certainly would have been the case had not the upper ends of the compartments, surrounding and forming the orifice, been nicely adapted always to yield, in a certain limited degree, to the disintegrating influences to which every shell is exposed, but which most Cirripedes can resist; and the disintegration of the narrow end of a conical tube, of course increases the diameter of its orifice. In [Tubicinella], in which the shell is furnished with narrow radii, and does increase in diameter from top to bottom, the increase is not sufficient in proportion to the continued elongation of the shell; to compensate for this, the orifice is enlarged at short intervals by the breakage of the upper end of the shell, for which purpose (as explained under the genus) it is evidently constructed. Hence we see that, in certain Cirripedes, decay or disintegration, and breakage, are necessary elements in their growth! It is a remarkable fact, which I cannot explain, that in some species in which the orifice of the shell is usually increased by disintegration, if individuals are so situated that they are not exposed to sufficiently energetic disintegrating influences, as may be inferred from the well-preserved condition of the whole surface of the shell, then the radii become developed, and the orifice is increased in size by the diametric growth of the upper part of the shell: I have seen instances of this in [Tetraclita porosa], and [purpurascens], and in [Balanus perforatus]: it appeared, but of course erroneously, as if the lateral growth of the compartments had been subjected to the will of the animal.

Considering the strength of the shell of sessile Cirripedes, the separation of their compartments one from another and from the basis, during growth, has justly been thought a surprising circumstance. In most [Chthamalinæ] and in some species of [Balanus], however, if the shell be boiled in caustic potash, the compartments fall apart with a touch; this shows that their union is due to animal and probably to organised matter, and the growth of such matter between the opposed edges of the compartments, and their consequent gradual separation, offers no particular difficulty. But in many Balani, boiling in potash for hours does not seem even to weaken, in the least degree, the sutures, which are wonderfully strong—the shell often breaking rather than yield on these lines; if, however, the shell be dissolved in acid, the animalised tissue which is left easily separates on the lines of suture, and if this tissue be boiled in potash, the remnants of the compartments fall quite separate. These facts seem to me to show, that the compartments in such cases are joined along the lines of suture by tissue, which must be in a calcified state, but which, nevertheless, continues to grow by intersusception; in other words, I believe that the tips of the complicated ridges and points interlocking on the lines of suture, are not separated from each other by films of corium or simple animal matter, but are actually united by corium in a calcified, yet still growing condition.

In ordinary Crustaceans, the growth is periodical and sudden; a new and larger carapace, for instance, is formed under the old one, and after the exuviation of the latter, the new one soon hardens, and does not subsequently increase in size; so it is in the case of Cirripedes, with the membranes of the body, and even with certain parts, as the opercular membrane, of the external covering. But a Cirripede cannot, like a crab, crawl into some crevice and remain protected till its shell becomes hardened; hence, probably, it is that the shell is never[24] wholly moulted. Even if the margins of the opposed compartments and of the basis were to grow rapidly, the shell would necessarily be much weakened on the lines of suture, and unable to withstand the heavy breakers, to which so many species of sessile Cirripedes are exposed. On the other hand, although the margins are thus compelled to grow slowly, they do not grow continuously, as may be seen in the zones of increment on all the valves, corresponding, I believe, with the periods of exuviation of the membranes of the body. A layer of shell, often very thin, seems to be generally deposited over the whole internal surface of the several valves, at the same time that the marginal zones are added; so that the only essential difference in the growth of the external covering, in Cirripedes as compared with ordinary Crustaceans, is that the old shell is not cast off, but adheres to the outside of the new shell, and that the margins are added to (in certain definite directions) slowly yet not continuously, instead of the whole being formed at a single period.

[24] In the genus [Alcippe], and in [Cryptophialus], the whole of the external membranes are moulted, excepting the surface of attachment; but then these Cirripedes live in cavities which they form for themselves, and are thus protected. In Lithotrya the membrane of the peduncle, with its little valves or scales, is moulted, but here, again, this very part is protected by the tubular cavity, which the animal forms and inhabits. Neither of these three genera belong to the [Balanidæ], or sessile Cirripedes, which we are now more especially describing.

If, now, a section of one of the shelly zones of growth be carefully examined, it can in some cases be distinctly seen to be formed of successive, excessively fine laminæ; but the animalised tissue (which differs much in amount in different Cirripedes) left after the shell has been dissolved in acid, exhibits, in most cases, neither laminæ nor any other structure whatever. The shell seems to be the actual pulpy corium, or true skin, in a calcified condition, but generally with its cellular structure modified and much reduced: I have taken a bit of recently-formed shell of [Tetraclita] and of [Coronula], with the corium still adherent to its under surface, and after dissolution in acid, I could not distinguish the part, which had just before existed as shell, from the corium itself. In the case of [Coronula], immediately prior to the period of moulting and growth, I found the unaltered corium so charged, as to effervesce, with carbonate of lime, either in a state of dissolution, or in granules too minute to be visible under the highest powers.

The sutures between the several compartments and the basis are covered by thin membrane, which is continually splitting during the growth of the opposed edges of the underlying shell; but previously to each splitting, a new slip of membrane is, I believe, already formed under the old one; so that the corium is not even momentarily exposed. Owing to this manner of growth, the slips of membrane consist of successive rims united together; in most cases, these soon become abraded from the older parts of the shell, but are sometimes preserved. The last-formed slip of membrane over a suture is homologous with the opercular membrane; and both are strictly analogous with the ring of flexible membrane, forming the joint of the leg of a crab. In the latter case, the flexible membrane and hardened crust are both moulted together: in the opercular membrane, there is a double line of splitting, one close round the opercular valves, and the other at the basal edge of the sheath, and the intermediate portion is moulted, but with a zone of membrane left adherent to the non-moulted valves and sheath: lastly, in the slips of membrane covering the sutures, there is only a single line of splitting, and no portion, I believe, is moulted; the rims of membrane on each side remaining adherent on the compartments and basis, until worn away.

The opercular membrane, when closely examined, exhibits no structure, except that it can sometimes be plainly seen to be composed of successive, numerous, excessively thin laminæ. Occasionally, however, it presents the false appearance of being permeated by parallel and anastomosing vessels: this appearance is clue to one or more of the component laminæ having been wrinkled before a succeeding lamina was thrown down and attached to its under side. If a small piece of an opercular valve of [Tubicinella], with the opercular membrane adhering to it, and with the corium under both, be dissolved in acid, it may be clearly made out that the corium under the valve has gone on being converted into shell, whereas under the opercular membrane it has been converted and condensed into fine constituent laminæ of chitine. Inasmuch as the successive layers of shell, during each period of growth, go on encroaching on those of the membrane, the line of junction between the shell and chitine becomes oblique or bevelled. The membrane on this bevelled line of junction assumes a slightly different aspect to what it has elsewhere; it becomes yellowish or brown, thicker and very much tougher. In many genera it is also furnished with a row of small bristles. At the period of exuviation the opercular membrane separates just outside this modified portion, leaving the latter adherent, as a rim or slip, on the valves. If, however, the opercular membrane be rudely torn off before its proper period of exuviation, it carries with it the as yet continuous, but already modified, slip. A slightly indented line may sometimes be traced before the period of exuviation, showing where the separation will take place: what produces this line I know not. The coloured, thickened, and modified slips of opercular membrane, which are thus retained adhering to the valves, and which together form an investing membrane, have been considered by most authors as the epidermis; but they have no more right to be thus called than has any other part of the opercular membrane. Exactly similar slips of membrane are left investing the sheath. So, again, the membrane which, when well preserved, invests the walls of the shell, is made up, as already stated, of successively adherent slips, which originally covered the lines of suture.[25]

[25] In the case of [Coronula] there is a peculiarity, described in the last section of this [Introduction], (under the head of Cementing Apparatus), namely, that the two or three last-formed, exterior zones of the Basal membrane continue for a period to increase in width; being, as I believe, dragged one from over the other, with fresh laminæ of membrane continually thrown down. In this same genus, and in [Tubicinella], the walls of the shell are invested by membrane, which is doubled inwards under their basal edges; and as the latter grow, the investing parietal membrane splits and separates from the basal membrane, and is pulled outwards and downwards. This inflected, often broad border of membrane, seems to me more strictly comparable with the opercular membrane, than with those narrow, thickened rims of yellowish membrane which in other Cirripedes cover the suture between the basal edges of the walls and the basis.

The little bristles above alluded to, which arise from the slips of membrane left adherent on the opercular valves, sheath, and walls, stand in rows; a row corresponding to each period of exuviation of the opercular membrane. The bristles are generally largest on the opercular valves and sheath; in [Balanus tintinnabulum], they are from 1 to 2/1000ths of an inch in length, but they are longer in some other species. I may here mention, as showing the connexion of these bristles with the opercular membrane, that similar bristles occur in [B. perforatus], scattered over the surface of that membrane, and are necessarily moulted with it. In the imbedded genera [Coronula] and [Tubicinella], none of these bristles exist. When a portion of valve or shell, furnished with bristles, is dissolved in acid, tough, sinuous, and apparently hollow, threads are seen to run from their bulb-like bases, into and up the corresponding layer, which, before dissolution, existed as shell; and they terminate internally in very fine points, which I believe are united to the underlying corium. These threads, or tubuli,[26] as I have called them in my volume on the Lepadidæ, are, in [Tetraclita porosa], about 1/5000ths of an inch in diameter, but only half that size in [B. tintinnabulum]. On parts of the shell where there are no bristles, similar tubuli penetrate the shelly layers, and come to the surface. The tubuli running to the lowest and last-formed row of bristles, just after a period of exuviation, are so delicate as hardly, or not at all, to be distinguished; in the row above, they are plain and longer, and for the next two or three upper rows they are, in some cases, as in [Tetraclita porosa], longer and longer, having been added to during each successive thickening of the valve. These tubuli consist of chitine, and no doubt first existed as threads of corium; they are so tough that they must serve to strengthen the successive layers of shell, but I imagine their chief function is to keep up the vitality of the newly-formed layers of shell. May we not, also, venture to suppose that by their means, some degree of sensibility is given to the bristles? I need only further remark, that in some species of [Balanus] and of [Chthamalus], the under side of the shell is penetrated by irregular pores, large enough to be visible to the naked eye, into which threads of corium penetrate; but these can hardly be said to appertain to the microscopical structure; and are more nearly related to those pores and furrows, formed by the greater or less development of the longitudinal septa, and in which the threads of corium deposit, or rather become changed into, transverse septa, or solid shelly matter, as previously described.

[26] I regret that I have used this term “tubuli”; for the threads thus designated, I believe, are not the same with the tubuli of Dr. Carpenter, which are not left after dissolution in acid. I have seen tubuli, as called by me, in the shell from the leg of a crab, after having been placed in acid.

Muscles of Sack.

In the pupa, the thorax, as we shall hereafter more fully see, is continuous with, and opens into the large anterior end or front part of the head; but during the metamorphosis (Pl. [30], fig. [2]), the thorax of the Cirripede becomes, owing to the almost transverse position occupied by the young animal within the pupa, to a great extent internally separated from the anterior end,—which anterior end forms, as we know, either the peduncle or the basis. Hence it comes to pass that the body or Thorax (Pl. [25], fig. [1]) is lodged within a sack (f) within the shell. The chitine membrane lining this sack is excessively thin and transparent, but less so in [Xenobalanus] and [Tubicinella]; it is obviously continuous with that investing the body of the animal; it is also essentially continuous with the opercular valves and membrane, and consequently with the whole shell. It is periodically moulted. It is lined by corium, as is likewise the surrounding shell; hence the corium is double round the sack, as indeed might have been expected from the shell and opercular valves (at least their upper parts) being formed by the prolongation, as is obvious in the pupa, of the posterior edges of the carapace. Between the two folds of corium, which are united together by transverse ligamentous fibres, branching out at both extremities, like the roots and branches of a tree, we have the longitudinal muscles, which go to the opercular valves; and likewise a layer-like mass of branching ovarian tubes (Pl. [25], fig. [1, g]): the ovarian tubes, however, are often confined to the base of the sack. In [Xenobalanus], the two folds of corium are united by longitudinal membranous septa, making a series of quite peculiar, square tubes.

The above-mentioned muscles are attached at their upper ends to the opercular valves, and at their lower ends to the basis. There are, in fact, three pairs, but the pair attached to the basi-carinal angles of the two terga (Pl. [25], fig. [1, i]), are almost invariably confluent, forming one great bundle; the second pair is attached to the lateral or basi-tergal corners of the two scuta, and are hidden in the figure; the third pair (h) is attached also to the scuta, to their rostral angles. These muscles can only act as depressores; they are often extremely powerful; they belong to the voluntary class, for they are transversely striped. By their action, the opercular valves are capable of varied slight movements, within the limit allowed by the width of the flexible opercular membrane. By the action of the lateral scutal depressores, the orifice leading into the sack is opened, the movement being generally aided by the protrusion of the cirri. By the sudden contraction of the rostral scutal depressores, the blows which are sometimes given by the beaked terga at the opposite end of the operculum, are probably effected. By the contraction of all three pairs of muscles, the opercular valves are held down with quite surprising force. The valves can be raised only by the action of the animal’s body against the basis.

In [Coronula] these muscles are more spread out, and do not extend down to the basis; their lower portions, as is likewise the case in [Tubicinella], do not exhibit transverse striæ, and hence tend to pass either into the involuntary class, or into ligament. This condition of the muscles, in the above two genera, accords with the little-developed state of their opercular valves. In [Xenobalanus], there is no longer any evidence of the muscles being collected into five or six bundles, for they are thinly and almost uniformly spread out, and show in no part transverse striæ. I may add that in much elongated specimens of [Balanus balanoides], these muscles become in their lower part ligamentous, and destitute of striæ.

Branchiæ.

In the [Balaninæ], a pair of Branchiæ is always present: they lie on each side, in a somewhat curved position, in the angle between the sides of the shell and the basis. In Pl. [25], fig. [1], they are exactly covered, on the further side, by the body of the animal. They are attached near each other at the carinal end of the sack in a vertical line, and likewise on each side in a transverse line, extending from close beneath the spur of the tergum towards the point of attachment of the body to the scutum. In [Balanus], as in the figure (Pl. [25], fig. [3]) of [B. tintinnabulum], each branchia consists of a medial fold of skin, a little curved conformably with the sack, and slightly tapering towards its rostral and free extremity; but this fold is almost hidden by the vertical sub-folds or membranous ridges, themselves plicated and sub-plicated, which project on both sides: these vertical folds are free at their tips: at their lower attached ends, they are thickest. On the side nearest the wall of the shell, the whole branchia has a bilobed appearance, owing to a very deep indentation caused by the projection of the scutal lateral depressor muscle; the sub-folds on this side are also more plicated. The branchia essentially is an inward plicated fold of the membranes of the sack; for its outer, very thin tunic is continuous with and moulted with that lining the sack; and within it we have two layers of delicate, pulpy, transparent corium, united together (as is best seen in [Coronula]) by ligamentous fibres, branched at their two ends, all exactly as in the corium surrounding the sack. There are here no distinct vessels, any more than in other parts of the body, but a fluid could easily circulate in the interspaces of the corium. From the large size of this organ, and its simplicity of internal structure, being adapted exclusively to expose a great surface of skin to the water, I do not doubt that it has been correctly considered as a respiratory organ. By the voluntary movements of the opercular valves (i. e. part of the carapace) the water is constantly being pumped in and out of the sack; the movement, indeed, may be almost compared to the heaving of a man’s chest. Moreover, the branchiæ on each side are attached so closely to the spur of the tergum, that each time the latter is moved, the whole branchia must, I think, be agitated, and the folds opened, as by the action of a lever.

In our two commonest, tidal, sessile Cirripedes, viz. [Balanus balanoides] and [Chthamalus stellatus], I have observed that, when left uncovered by water, they kept the orifice of their operculums a little open, with a bubble of air within their sacks, so that the orifice was in fact closed by a thin septum of water, with air beneath; when disturbed, they closed their operculums with force, and expelled the bubble of air with a clicking noise, which has been noticed by Dr. Coldstream,[27] and has been thought to be made by the movement of the operculum itself. [Bal. crenatus], a deep-water species, when out of water, keeps its operculum closed.

[27] ‘Encyclopædia of Anatomy and Physiology;’ article Cirrhopoda.

In [Coronula], [Platylepas], [Tubicinella], and [Xenobalanus], each branchia[28] consists of two unequal folds, both plicated on both sides: in the two latter genera, they extend far down the deep and elongated sacks, and hence the area of surface altogether gained is extremely great. In most of the species of [Chthamalus], the branchia consist of a small fillet barely plicated: in the allied [Chamæsipho columna], they are rudimentary, forming a smooth little pouch only 1/100th of an inch in length: in [Chthamalus scabrosus] they are quite aborted, being perhaps represented by a slight hairy ridge; but in [Chthamalus dentatus], and therefore within the limits of the same genus, the branchiæ (and this seems to me a singular fact) are large, each being composed of two plicated folds, as in [Coronula]. Tapering filaments situated near the bases of the cirri, such as those occurring in several species of the Lepadidæ, are not found in any sessile Cirripede; but I have observed nearly similar filaments, projecting upwards and inwards at the base of the sack, in several species of [Balanus] and in [Coronula]; those which I examined were simply occupied by delicate corium, and no doubt must aid in exposing a greater surface of corium to the circumambient water.

[28] Burmeister has given a good figure (Tab. 2, fig. 10) of the branchiæ of [Coronula], (but the two folds are shaded too unequally), in his ‘Beiträge zur Naturgeschichte der Rankenfüsser.’

In my former volume on the Lepadidæ, I have described the ovigerous fræna occurring on the two sides of the sack, to which the ovigerous lamellæ are attached by a peculiar glandular secretion: in the [Balanidæ] there are no ovigerous fræna, but the branchiæ just described are identical with the fræna in essential structure and in position; differing only in being placed a little nearer to the carinal end of the sack, and in being generally (but not always) larger and more plicated: seeing this, and that in [Alcippe lampas], and in some species of Pollicipes,—the genus which comes nearest to the [Balanidæ],—the ovigerous fræna are large and are destitute of glands, and have therefore lost their normal function of supporting the ovigerous lamellæ, I can hardly doubt that the branchiæ in the [Balanidæ] are the ovigerous fræna of the Lepadidæ in a modified condition; a transformation of function not greater than that of the swimming bladder of a fish into the lungs of the higher Vertebrata.[29]

[29] There is, I conceive, no foundation for the belief of some authors that the branchiæ of the [Balanidæ] are in any way connected with the ovaria.

Thorax and Body.

Parts of the body included within the shell or carapace.—These parts (Pl. [25], fig. [1]) consist of the prominent mouth, and of the thorax (c′), with its largely developed portion, called the prosoma (c), and with its appendages. The abdomen is quite rudimentary, being represented merely by a small portion of membrane surrounding the anus, and sometimes inserted like a wedge between the inwardly inflected posterior thoracic segments; in only two genera ([Catophragmus] and [Pachylasma]), its nature is rendered somewhat plainer by supporting caudal appendages. The probosciformed penis lies folded under the thorax; and I believe (from what is seen in the anomalous genus [Proteolepas]), that it normally arises from the ventral surface of the terminal point of the rudimentary abdomen.[30] The thorax is laterally compressed, the ventral surface being very narrow, with the bases of the cirri placed closely together. It consists, in appearance, of two very different portions; one a soft, more or less rounded bag, which I have called the prosoma; and the other, which supports the five posterior pairs of cirri, is narrower, invested with stiffer membrane, and is more or less distinctly composed of five segments. These segments (Pl. [26], fig. [8]) on their dorsal and dorso-lateral surfaces, are generally driven like wedges one into the other, with their points directed anteriorly: on the ventral surface the articulations are transverse. The prolongation (e) of the thin membrane (a) surrounding the anus (b), that is, the rudiment of the abdomen, which sometimes carries caudal appendages, almost divides (in appearance, whether really I know not) the hindermost thoracic segment along the medio-dorsal line, into two parts. I have given the above drawing of these segments, but with the dorsal surface much flattened, in [Coronula diadema]; in most species of [Balanus], however, the wedges formed by one segment being driven into another, are much sharper; on the other hand, in [Xenobalanus] they are nearly straight and transverse. The three posterior segments are always the most distinct; the two next segments are also distinct laterally, but along the dorsal surface they become, excepting in [Xenobalanus] and some few other cases, completely confluent. The greater distinctness of the posterior segments is conformable to what takes place in the higher Crustacea. The articulations between the segments are folded inwards, and are formed of thin membrane, which in some cases, as in [Coronula diadema], forms a marked contrast with the much thicker, stiffer, and yellowish membrane of the segments themselves; in [Balanus tintinnabulum], however, the whole membrane of the five thoracic segments is very thin, excepting small wedge-shaped portions along the medio-dorsal line. The infolded articulations between the segments supporting the three anterior pairs of cirri (at least in the [Balaninæ]), are much wider than those between the three posterior segments; the former segments, with their cirri, being consequently capable of being moved further apart from each other. Could there have been any doubt as to the distinctness and reality of the five thoracic segments, it would have been set aside by the arrangement of the muscles attached to them, as will presently be described. I need only add, that in many genera there are shield-like swellings at the exterior bases of the pedicels of the posterior cirri, which I for some time thought were the epimeral elements of the thoracic segments; but I now believe them to be parts of the pedicels of the cirri. The basi-exterior margin, moreover, of the pedicel of the third pair of cirri, in many species of the [Balaninæ] (Pl. [25], fig. [1]), is produced as a plate, thickly fringed with fine hairs, half across the dorsal surface of the thorax; serving, apparently, as a brush to clean the sack, or to prevent the ingress of any intruding substance.

[30] Von Siebold and Stannius, in their ‘Anatomie Comparée,’ tom. i, p. 473, and p. 440, (foot-note), consider the articulated probosciformed penis as an elongated abdomen; a view which, at the commencement of my examination, I was tempted to admit; but the position of the caudal appendages on the dorsal basis of the penis, suffice, I think, to show that this view is not correct; for these caudal appendages evidently correspond with those borne on the very extremity of the abdomen in the pupa. Nor, indeed, does the position of the anus accord well with such a view.

The soft, rounded, bag-like portion of the body, which I have called the prosoma, is usually separated by a notch from the five posterior thoracic segments; at its upper end it may be said to carry the mouth and first pair of cirri. The prosoma includes the main part of the stomach and the broad ends of the vesiculæ seminales. It is always clothed by very thin membrane, which in [Chthamalus dentatus], is hairy. In [Tubicinella] and [Xenobalanus], the prosoma is much elongated, being produced far down the deep sack. That the prosoma is mainly formed by a great development of that segment (homologically the second thoracic segment) which carries the first pair of cirri, is certain, and I should not have hesitated to have said that it was exclusively so formed, had not the first thoracic segment in the anomalous genus [Cryptophialus] been developed as a distinct and free segment, not attached to the carapace; showing that possibly in other Cirripedes, the dorsal half of this first thoracic segment may be concerned in the formation of the free prosoma.

Muscular System.

Attachment of the Body to the Shell.—The prosoma which carries the posterior thoracic segments, and in appearance the mouth, is the only part of the body which is attached to the general covering (Pl. [25], fig. [1]), namely, to the opercular valves. Except through the continuity of the lining membranes of the sack, the body lies free within the walls of the shell. The area of attachment (shown by a sinuous broken line round a and b) extends from about the middle of the two scuta down to their basal margins. As these valves lie obliquely across the orifice of the shell, the animal’s body comes to be suspended almost in the middle of the sack. The two scuta, as we have seen, have the power of opening and shutting a little; and are brought together by the adductor scutorum muscle (a), which is generally very powerful. The body is attached to these valves, round and beneath the adductor, so as to hide it until one of the valves be removed. The attachment is chiefly effected by three pairs of widely expanded, superficial muscles, two pairs of which are spread over the flanks of the prosoma, and the third pair over its rounded (properly dorsal) surface, which lies close to the rostral compartment (A, fig. [1]) of the shell. I should have stated, that my chief examination of the attachment of the body to the scutal valves, has been made on [Coronula balænaris], and less closely on [Balanus tintinnabulum]. Within these three pairs of superficial muscles, there are (besides the adductor) no less than five other pairs; of these one long pair is attached at one end to the basal margin of the labrum (e), and at the other end, to the under side, near to the basal margin of the scuta: two other, shorter, parallel pairs of muscles are attached at one end to the interspace of membrane between the basal edge of the labrum and the adductor scutorum muscle, and at the other end, to the under side of the scuta, above the attachment of the first pair: the fourth and shortest pair curls close under the adductor, and is there attached at both ends beneath it. The action of these four pairs of muscles must be to draw back, from the orifice of the shell, the mouth, and that interspace of body between the basal margin of the labrum and the adductor muscle. This movement I saw in living specimens. The last and fifth pair of muscles is small, but of considerable length; it is a diverging pair, attached at the converging end, above and exteriorly to the adductor muscle; and at the diverging end, low down on the under side of the scuta; I am very doubtful regarding the function of this pair. Altogether we have seen that round and within the fleshy pedicel, by which the body is attached to the scuta, there are no less than eight pairs of muscles. The central space between these muscles is hollow, and here many lacunal channels seem to converge. These muscles receive nerves from the supra-œsophageal ganglions. The interspace above alluded to, between the basal edge of the labrum and the adductor scutorum muscle, occupies a very different position according as the animal’s body is protruded as far as it can be, or is retracted. It is homologically part of the third cephalic segment; and consequently the mouth ought to have stood posteriorly (i. e. above, in the position figured in Pl. [25], fig. [1]) to this interspace; yet, in fact, when the animal is retracted within its shell, the mouth usually lies almost directly beneath this interspace and the adductor scutorum muscle.

Besides these muscles of attachment, the prosoma is furnished with several other muscles. There are superficial muscles running up towards the basal margin of the sides of the mouth; and other deeper muscles, to which, I presume, the movements of the mouth, as a whole, are due. The muscles moving the gnathites do not, as far as I could make out, extend beneath the basal edge of the mouth. There are, also, powerful muscles giving movement to the basal segments of the pedicel of the first pair of cirri. Again, there are superficial muscles running to the next succeeding thoracic segment; the anterior ends of which are separated by a clear interspace from the lower ends of the above-described superficial muscles, by which the prosoma is attached to the scuta. On each flank, moreover, but more deeply imbedded, are the long flexor and extensor muscles, presently to be described, running to the five posterior thoracic segments. The last muscle which I need here mention, is a deep-seated diverging pair, attached near the upper end of the stomach, on its ventral surface, and diverging from this point to the sides of the prosoma high up beneath the mouth. The probable action of this pair, as well as of the three superficial pairs of muscles by which the body is attached to the scuta, is to draw up the whole prosoma towards or from the orifice; and likewise to contract it firmly, so as to serve as a fulcrum for the movements of the five posterior thoracic segments, together with the cirri, which they carry.

The muscles of these five thoracic segments are numerous and powerful; they are also complicated, chiefly owing to the segments on their dorsal and dorso-lateral surfaces being driven, like wedges, one into the other. As far as I could make out, there are on each side three, superficial, dorso-lateral and lateral muscles (generally, if not always, destitute of striæ), which do not cross the articulations, but extend merely from articulation to articulation; and of which the function can be only to contract each separate segment, and consequently to open out the intermediate infolded articulations; the effect of this would be to separate slightly the cirri from each other,—more especially those borne on the two or three anterior segments, between which the infolded articulations are deeper or broader. There are other more deeply imbedded, powerful, long, dorso-lateral extensor, and ventri-lateral flexor muscles, attached at one end within the flanks of the prosoma, and at the other end to the successive segments of the thorax. The action of the former is to straighten and stretch out the thorax; of the latter, or ventri-lateral muscles, to retract it. In tracing these muscles, a fascia could be seen to become attached to a segment, and then this same fascia would run on to the next succeeding segment: the effect of this must be, that each segment can be retracted and protracted either from the prosoma as a fulcrum, or from the antecedent segment as a fulcrum: we have, also, seen that each segment can, by the agency of the superficial, non-striated muscles, contract itself. Hence these thoracic segments are capable of diverse movements, as was very evident when the shell of a living specimen was opened. By one movement in common, the whole five posterior segments could be drawn back, so as to become even partly imbedded in the prosoma: lateral, twisting or wriggling movements were also quite distinct: the three posterior segments seemed to be capable of less independent movements than the anterior segments; and I observed that the more powerful flexor and extensor muscles did not run into these three posterior segments. The cirri, of course, partake of the movements of the thorax; and in watching, in an uninjured specimen, the alternate, protruding, gracefully sweeping and retracting movements of the posterior pairs of cirri, it was evident that the thorax was the chief agent in their movement. Besides the muscles now mentioned, there are some immediately to be noticed, which extend from within the thoracic segments to within the pedicels of the cirri.

Movements and Muscles of the Cirri.

Although the cirri have not been described, it will be most convenient here to treat shortly of their muscles. Each cirrus consists of a pedicel, having a long basal and a short upper segment, supporting two multiarticulate rami. The lower segment of the pedicel can be drawn forward by an adductor muscle, attached low down within this segment, and crossing at right angles (at least in the case of the anterior cirri) the corresponding muscle of the opposed cirrus, on the central, ventral surface of the thorax. This segment can also be drawn back by a muscle springing from the dorso-lateral surface of the thorax, and running only a little way within the segment: I am far from sure that the lower segment does not possess other muscles. The short upper segment of the pedicel can be moved backwards and forwards, as I saw in living specimens, independently of the lower segment; this movement being best seen in the anterior cirri, which are much more often moved independently of each other than are the posterior cirri. The rami are capable, I believe, of being moved backwards and forwards as a whole, by the movement of the few lower segments, which are generally more or less confluent. They can, also, be curled up and uncurled by the combined movement of each separate segment. The uncurling seems to separate the two rami a little laterally. Each ramus, at least in the two or three anterior pairs, can be moved to a certain extent, independently of the other ramus of the same cirrus; and the few terminal segments, either of both rami or of one ramus, are often a little moved and curled (and this is especially the case with the long anterior ramus of the first pair), without the lower segments or the pedicel being moved.

The flexor and extensor muscles, which, as I believe, move the upper segment of the pedicel (a and b, Pl. [29], fig. [1]), are attached at their upper ends to its basal margin, and are thus enabled to draw it a little way down within the lower segment, and so move it. The short flexor muscle (c), which is attached at its lower end within the upper segment of the pedicel, and the longer extensor (d), also, attached within this same lower segment, serve, I believe, to move the lower, partially confluent segments of each ramus as a whole. In the case of these muscles, and of those last mentioned, I am surprised that the extensors (b) and (d) are not attached nearer to the exterior and dorsal surface. Other muscles (e, f) attached at their lower ends within the upper segment of the pedicel, run up each of the two rami to their tips, with some of the fasciæ terminating within each segment: of these muscles, the outer one (f, f) appears to be the extensor, and the inner one (e, e) the flexor. But besides these, there are other short flexor muscles (g, g) which run on the anterior face,[31] from segment to segment, serving to pull the front edge of one segment within the edge of the next lower segment. These muscles differ much in plainness in the several genera: they are very distinct in [Coronula]. In some specimens of this genus, a few of the articulations between the basal segments of the rami having been obliterated, the short muscles (g, g) running from articulation to articulation were absent, and their presence and nature in the upper segments thus rendered the plainer. The muscular system in the several pairs of cirri seems to be the same, with the exception of the first pair, in which the muscle answering, as I suppose, to (a), namely, the flexor of the upper segment of the pedicel, is much spread out at its lower end, and is there attached to the exterior surface of the lower segment.

[31] For a considerable time I thought that there were muscles going to the spines, especially to those which arise from the upper dorsal edge of each segment; but I have since ascertained that these are the cases within which new spines, with their lower ends doubled like the fingers of a glove hastily pulled off, are in process of formation.

The backward and forward movements of the segments, both in the rami and in the pedicels of the cirri, are apparently effected, as already noticed, by the outer or inner (as the movement may be) basal edge of one segment being drawn a little way down within the next succeeding lower segment. If, at the same time, both the inner and outer margins of all the segments were drawn one within the other, the whole limb would necessarily be shortened; and I distinctly saw a shortening action, with very slight movement in any other direction, in the first and second pairs of cirri; and I think it almost certain that this movement might be performed by the other cirri. If I correctly understand a statement of Milne Edwards,[32] this is an important fact, as he asserts that only the higher Crustaceans possess the power of shortening their limbs.

[32] ‘Annales des Sciences Naturelles,’ tom. xviii, 1852, p. 121.