The Project Gutenberg eBook of A Monograph on the Sub-class Cirripedia (Volume 2 of 2), by Charles Darwin
This eBook is for the use of anyone anywhere in the United States and
most other parts of the world at no cost and with almost no restrictions
whatsoever. You may copy it, give it away or re-use it under the terms
of the Project Gutenberg License included with this eBook or online at
www.gutenberg.org. If you are not located in the United States, you
will have to check the laws of the country where you are located before
using this eBook.
Title: A Monograph on the Sub-class Cirripedia (Volume 2 of 2)
The Balanidæ, (or Sessile Cirripedes); the Verrucidæ, etc., etc.
Author: Charles Darwin
Release Date: July 25, 2014 [eBook #46408]
[Most recently updated: December 26, 2021]
Language: English
Produced by: Bryan Ness, Emmanuel Ackerman and the Online Distributed Proofreading Team
*** START OF THE PROJECT GUTENBERG EBOOK A MONOGRAPH ON CIRRIPEDIA, VOL 2 ***
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.
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.
Transcriber Added
Table of Contents.
Page
Dedication v
Preface vii
_Monograph on the Cirripedia_ 1
Introduction 1
On the Names given to the different parts of Cirripedes 3
Class Crustacea, Sub-Class Cirripedia 9
On the Sexual Relation of Cirripedes 23
Order I.--Thoracica 30
Family Balanidæ 33
Table of Contents 33
Structure of Shell 34
Structure of the Individual Compartments 43
Structure of the Radii 45
Structure of the Alæ 47
Structure of the Sheath 48
Structure of the Basis 49
Structure of the Opercular Valves (Scuta and Terga) 51
Growth of the Whole Shell, and Its Microscopical Structure
54
Muscles of Sack 61
Branchiæ 63
Thorax and Body 65
Muscular System 68
Movements and Muscles of the Cirri 71
Mouth 74
Cirri 81
Caudal Appendages 85
Alimentary Canal 85
Circulatory System 87
Nervous System 88
Eyes and Vision 93
Acoustic Organs 95
Olfactory Sacks 97
Male Organs of Generation 97
Female Organs of Generation 100
Metamorphoses and Homologies 102
Larva, First Stage 103
Larva, Second Stage 109
Larva in the Last or Pupal Stage 110
Act of Metamorphosis 126
On the Homologies of the Carapace and Shelly Valves 131
Cementing Apparatus 133
Affinities, Classification, Variation 152
Rate of Growth, Exuviation, Powers of Repairing Injuries
156
Geographical Range and Habits 159
Geological History 172
Sub-Family Balaninæ 175
1. Genus Balanus 177
Sections of the Genus 193
Section A 194
1. Balanus tintinnabulum 194
Var. communis, vesiculosus, validus, zebra, crispatus,
spinosus, coccopoma, concinnus, intermedius, occator,
d'Orbignii
Varieties 201
2. Balanus tulipiformis 204
3. Balanus psittacus 206
4. Balanus Capensis 209
5. Balanus Nigrescens 210
6. Balanus decorus 212
7. Balanus vinaceus 213
8. Balanus Ajax 214
Section B 216
9. Balanus stultus 216
10. Balanus calceolus 218
11. Balanus galeatus 220
12. Balanus cymbiformis 221
13. Balanus navicula 221
Section C 223
14. Balanus trigonus 223
15. Balanus spongicola 225
16. Balanus lævis 227
Var. nitidus, Coquimbensis
17. Balanus perforatus 231
Var. angustus, Cranchii, fistulosus, mirabilis
18. Balanus concavus 235
19. Balanus amphitrite 240
Var. communis, venustus, pallidus, niveus, modestus,
Stutsburi, obscurus, variegatus, cirratus
Varieties 245
20. Balanus pœcilus 245
21. Balanus eburneus 248
22. Balanus improvisus 250
Var. assimilis
23. Balanus nubilus 253
24. Balanus corrugatus 254
Section D 256
25. Balanus porcatus 256
26. Balanus patellaris 259
27. Balanus crenatus 261
28. Balanus glandula 265
Section E 267
29. Balanus balanoides 267
Varieties 270
30. Balanus cariosus 273
31. Balanus declivis 275
Section F 277
32. Balanus Hameri 277
33. Balanus amaryllis 279
34. Balanus allium 281
35. Balanus cepa 283
36. Balanus quadrivittatus 284
37. Balanus terebratus 285
38. Balanus vestitus 286
39. Balanus Imperator 288
40. Balanus flosculus 290
Var. sordidus
41. Balanus bisulcatus 293
Var. plicatus
42. Balanus dolosus 295
43. Balanus unguiformis 296
Var. erisma
44. Balanus varians 298
45. Balanus inclusus 299
2. Sub-Genus Acasta 302
1. Acasta spongites 308
2. Acasta sulcata 310
3. Acasta cyathus 312
4. Acasta undulata 313
5. Acasta glans 314
6. Acasta lævigata 315
7. Acasta fenestrata 316
8. Acasta purpurata 318
9. Acasta sporillus 319
3. Genus Tetraclita 321
1. Tetraclita porosa 329
Var. communis, nigrescens, viridis, rubescens, elegans,
communis (young), patellaris
2. Tetraclita serrata 334
3. Tetraclita rosea 335
4. Tetraclita purpurascens 337
5. Tetraclita costata 339
6. Tetraclita vitiata 340
7. Tetraclita cœrulescens 342
8. Tetraclita radiata 343
4. Genus Elminius 345
1. Elminius Kingii 348
2. Elminius modestus 350
3. Elminius plicatus 351
4. Elminius simplex 353
5. Genus Pyrgoma 355
1. Pyrgoma anglicum 360
2. Pyrgoma Stokesii 361
3. Pyrgoma cancellatum 362
4. Pyrgoma conjugatum 364
5. Pyrgoma grande 365
6. Pyrgoma milleporæ 367
7. Pyrgoma dentatum 369
8. Pyrgoma crenatum 370
9. Pyrgoma monticulariæ 372
Species dubiæ 374
6. Sub-Genus Creusia 375
1. Creusia spinulosa 376
Varieties with the Scuta and Terga calcified together 380
7. Genus Chelonobia 382
1. Chelonobia testudinaria 392
2. Chelonobia caretta 394
3. Chelonobia patula 396
Second Section of the Sub-Family of Balaninæ 397
8. Genus Coronula 397
1. Coronula balænaris 415
2. Coronula diadema 417
3. Coronula reginæ 419
4. Coronula barbara 421
Species Dubiæ 423
9. Genus Platylepas 424
1. Platylepas bissexlobata 428
2. Platylepas decorata 429
Species Dubiæ 430
10. Genus Tubicinella 430
1. Tubicinella trachealis 431
11. Genus Xenobalanus 438
1. Xenobalanus globicipitis 440
Sub-Family Chthamalinæ 446
12. Genus Chthamalus 447
1. Chthamalus stellatus 455
Var. communis, fragilis, depressus
2. Chthamalus antennatus 460
3. Chthamalus cirratus 461
4. Chthamalus fissus 462
5. Chthamalus dentatus 463
6. Chthamalus Hembeli 465
7. Chthamalus intertextus 467
8. Chthamalus scabrosus 468
13. Nov. Genus Chamæsipho 470
1. Chamæsipho columna 470
2. Chamæsipho scutelliformis 472
14. Nov. Genus Pachylasma 475
1. Pachylasma giganteum 477
2. Pachylasma aurantiacum 480
15. Genus Octomeris 482
1. Octomeris angulosa 483
2. Octomeris brunnea 484
16. Genus Catophragmus 485
1. Catophragmus polymerus 487
2. Catophragmus imbricatus 490
Remarks on Bronn's List of Fossil Balaninæ and Chthamalinæ 492
Family Verrucidæ 495
Genus Verruca 496
Powers of Excavation 512
1. Verruca Strömia 518
2. Verruca lævigata 520
3. Verruca Spengleri 521
4. Verruca nexa 522
5. Verruca prisca 525
Family Lepadidæ 526
Genus Alcippe 529
Alcippe lampas 530
Female 530
Male 555
Order II.--Abdominalia 563
Cryptophialus minutus 566
Female 566
Male 584
Order III.--Apoda 587
Proteolepas bivincta 589
Synopsis et Index Systematicus 606
Synopsis et Index Systematicus Specierum, et recentium, et fossilum
611
Description of plates 641
Plate 1. Balanus tintinnabulum 641
Plate 2. Genus Balanus 641
Plate 3. Genus Balanus 642
Plate 4. Genus Balanus 642
Plate 5. Genus Balanus 642
Plate 6. Genus Balanus 643
Plate 7. Genus Balanus 643
Plate 8. Genus Balanus 644
Plate 9. Sub-Genus Acasta 644
Plate 10. Genus Tetraclita 645
Plate 11. Genera Tetraclita and Elminius 645
Plate 12. Genera Elminius and Pyrgoma 646
Plate 13. Genera Pyrgoma and Creusia 646
Plate 14. Genera Creusia and Chelonobia 647
Plate 15. Genera Chelonobia and Coronula 648
Plate 16. Genus Coronula 649
Plate 17. Genera Platylepas, Tubicinella, and Xenobalanus 651
Plate 18. Genus Chthamalus 652
Plate 19. Genera Chthamalus, Chamæsipho, and Pachylasma 652
Plate 20. Genera Pachylasma, Octomeris, and Catophragmus 653
Plate 21. Genus Verruca 654
Plate 22. Alcippe lampas 655
Plate 23. Genera Alcippe and Cryptophialus 658
Plate 24. Genera Cryptophialus and Proteolepas 660
Plate 25. Genera Proteolepas and Balanus 662
Plate 26. Structure of the Mouth and Thorax 664
Plate 27. Nervous System and Senses 666
Plate 28. Cementing Apparatus 667
Plate 29. Cirri and Larvæ, first stages 669
Plate 30. Larvæ of Lepas: second and last stages of development
671
Errata 674
Index 675
Plates 685
THE
RAY SOCIETY.
INSTITUTED MDCCCXLIV.
[Illustration]
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
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.
[Illustration: 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.
[Illustration: Fig. 2.
Fig. 2. Compartment with two radii, serving either as a
Rostrum or Rostro-lateral compartment.]
[Illustration: Fig. 3.
Fig. 3. serves as a Lateral and Carino-lateral
Compartment.]
[Illustration: Fig. 4.
Fig. 4. serves as a Carina or Rostrum.]
OPERCULAR VALVES.
[Illustration: Fig. 5. SCUTUM (internal view of).]
[Illustration: Fig. 6. TERGUM (external view).]
[Illustration: 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.
Page
Structure of shell 34
" of the individual compartments 43
" of the radii 45
" of the alæ 47
" of the sheath 48
" of the basis 49
" of the opercular valves (scuta and terga) 51
Growth of whole shell and microscopical structure 54
Muscles of sack 61
Branchiæ 63
Thorax and body 65
Muscular system 68
Movements and muscles of the cirri 71
Mouth 74
Cirri 81
Caudal appendages 85
Alimentary canal 85
Circulatory system 87
Nervous system 88
Eyes and vision 93
Acoustic organs 95
Olfactory sacks 97
Male organs of generation 97
Female organs of generation 100
Metamorphoses and homologies 102
Larva, first stage 103
Larva, second stage 109
Larva, last or pupal stage 110
Act of metamorphosis 126
On the homologies of the carapace 131
Cementing apparatus 133
Affinities, classification, variation 152
Rate of growth, exuviation, &c. 156
Geographical range and habitats 159
Geological history 172
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.
[Illustration: 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.
[Illustration: 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.
[Illustration: 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.
[Illustration: 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.
[Illustration: 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.
When a Cirripede is alive, the action of the cirri is really beautiful:
from the position of the thoracic segments, the posterior cirri (three
pairs in the Balaninæ and four pairs in the Chthamalinæ) form a sort of
semicircle facing the mouth: the anterior cirri stand further apart,
and are opposed in pairs to each other, with the first pair pointing
beyond the mouth. Together the cirri form a hollow cone, not circular
but elongated, with the mouth situated at the lower anterior end. The
posterior cirri are protruded, by the movement of the whole thorax,
curled up, close along the carinal end of the orifice; as they are
protruded, they diverge, both by the movement of their pedicels, and,
as I believe, by the separation of the thoracic segments. As the two
rami of each separate cirrus are uncurled, they also diverge a little;
as do the double rows of spines on the segments in each ramus, by their
elasticity. By the movement of the thorax, the cirri are then swept
towards the rostrum; and, lastly, they are brought perpendicularly down
towards the mouth with a rapid movement, which would be beautifully
adapted to catch any object floating or swimming in the water; hence
I have called the action of the cirri, captorial. When the shell of
a Balanus is broken open, the second and third pairs of cirri are
repeatedly clasped over the mouth with a convulsive movement, in a
manner indicating, I think, that their chief function is to seize
and carry to the mouth any object entangled by the sweeping movement
of the three posterior pairs. The first pair is also well adapted
to aid in this seizing action; but I suspect that the long anterior
ramus likewise acts as an organ of touch, warning the animal of
danger. The mouth being itself moveable as a whole,--the outer maxillæ
being capable of a backward and forward sweeping action, and being
furnished with orifices apparently olfactory,--the inner maxillæ having
more diversified movements,--the toothed mandibles overhanging the
œsophagus,--and the œsophagus itself possessing a powerful swallowing
movement, are all admirably adapted to secure any prey, when once
entangled by the cirri.
_Mouth._
The mouth, in the sub-family Chthamalinæ, cannot be distinguished
from that of the Lepadidæ, which has been pretty fully described
in my former volume. In the Balaninæ, however, the labrum differs
considerably in not being swollen; that is, in its outer and inner
fold of membrane being close together, and in having a central notch:
the palpi are also larger, and the lower teeth on the mandibles, are
laterally (Pl. 26, fig. 5) double, as will be more particularly stated
under these two sub-families. I have given a drawing (Pl. 26, fig.
1) of the mouth, seen from above, of _Balanus perforatus_, with the
right-hand palpus (_d′_) and outer maxilla (_a′_) cut off, in order
that the labrum (_e_), mandibles (_c_), and inner maxilla (_b_) might
be better shown; the cut-off bases (_x_, _x_) of the first cirrus on
each side are also shown. In fig. 2 we have the deep supra-œsophageal
cavity in _Bal. improvisus_ torn open and laid flat, with the inner
surfaces of the labrum (_c_) and outer maxillæ (_a_) exhibited, the
palpi, mandibles, and inner maxillæ having been removed. Figs. 3 and
4 will presently be referred to; they are parts of the mouth, with
the muscles, &c. removed, of Coronula. The mouth differs extremely
little in the different genera and species of the Balanidæ, much less
than amongst the Lepadidæ. In the Balaninæ, the crest of the labrum
is sometimes hairy, instead of having, as is usual, from two to six
teeth on each side of the central notch: in _Balanus improvisus_ (Pl.
26, fig. 2) and _eburneus_, and in _Chelonobia_, the crest on each
side of the central notch (_e′_) is furnished with a row of finely
graduated teeth. A sub-triangular portion of the inner fold of membrane
of the labrum, which overhangs the œsophagus, is always thickened and
yellowish; it is also often punctured in patterns (Pl. 26, fig. 2,
_f_), which, I believe, give attachment to little muscles that serve
to open the upper end of the œsophagus. Opposite to this thickened,
sub-triangular portion of membrane, the thin membrane forming the
supra-œsophageal cavity (or the cavity surrounded by the gnathites)
is strengthened by a pair of curved ribs (_h_, fig. 2) of thickened
yellowish membrane, running down from the inner bases (_a′′_) of the
bilobed outer maxillæ to the opening of the œsophagus (_g_): a broad
branch from each of these ribs supports the sides of the orifice of
the œsophagus; and this branch almost joins on to a slightly thickened
rim or bar (_f′_), which branches off from the upper part of the
sub-triangular (_f_) inner fold of the labrum. This structure, in _Bal.
improvisus_, is represented in Pl. 26, fig. 2, as well as it could be,
considering that the deep supra-œsophageal cavity has to be torn open;
and then laid flat.
The _Palpi_ (Pl. 26, fig. 6) differ little, except in size, in the
different genera, being squarish, more or less elongated, or even
approaching to club-shaped: in most of the Balaninæ they are larger
even than the mandibles, of which they normally form a part. Their
upper margins, especially towards their free extremities, are always
thickly clothed with spines; and there is generally a single row,
either short (_r_) or long, of spines of greater length, which arise
from a little above, and stand almost in a parallel line to, the basal
margin. On the internal surface, there is sometimes a row (_t_) of
very short little spines, which overhang the crest of the labrum. The
_Mandibles_ (Pl. 26, fig. 5) have from three to five teeth; the lower
point or angle is generally pectinated. In Coronula and its close
allies, there are some small teeth intermediate between the four or
five main teeth; and in these genera, though members of the sub-family
Balaninæ, the lower teeth exhibit only rudiments of being laterally
double.[33] The _Maxillæ_ sometimes have a notch under the upper large
pair of spines, and in _Octomeris brunnea_ there is a double notch: in
many species of Balanus, the inferior corner stands up like a step (Pl.
26, fig. 7, _a_): in many other genera and species, the whole edge is
straight. In all, or almost all cases, the row of spines on the middle
portion is double. The _Outer Maxillæ_ are always bilobed on their
inner faces, and are clothed with bristles. On all the gnathites, the
bristles are often doubly serrated.
[33] M. Martin St. Ange describes, in his 'Mémoire sur
l'Organisation des Cirripèdes,' pp. 15 and 32, "_une petite
langue_" in the mouth of Lepas; but I may venture to assert that
such does not exist; it is merely the point of union between the
outer maxillæ. M. St. Ange, in his comparison of the mouth of
Lepas with that of Phyllosoma, compares the mandible of the latter
with the palpus of Lepas; the first maxilla of Phyllosoma with the
mandible of Lepas; and so on with the other gnathites.
_Muscles and functions of the Gnathites, and their confluence._--The
outer maxillæ appear at first like a deeply-lobed lower lip, for they
reach over almost to the labrum (Pl. 26, fig. 1), and thus partially
cover the other organs; they are separately capable of a strong and
rapid, to and fro movement, by which no doubt they sweep any prey,
entangled by the cirri, towards the other gnathites. Each outer maxilla
is furnished with a pair of muscles, apparently a flexor and extensor;
there is also a little muscle between the two maxillæ, I presume for
the purpose of bringing them together. The outer and inner maxillæ
generally stand close together, and in several genera a little way
apart, from the mandibles; but there is no trace of any labrum or true
lower lip, bounding the mandibles and orifice of the œsophagus. The
outer and inner maxillæ and mandibles are not opposed in pairs to each
other, but against the thickened inner fold of the labrum; almost in
the same manner as the posterior pairs of cirri are not opposed one to
the other, but to the mouth.
I have described pretty accurately the muscles of the mandibles in
my former volume, and there given a drawing (Pl. 10, fig. 1) of them.
There are four muscles: first, the depressor muscle, which is the
largest, and is attached, at its upper end, to ligamentous apodemes
under the free toothed portion of the jaw; and at its lower end,
spreading considerably out, is attached to a concavity close above the
basal margin of the labrum; to understand the action of these muscles,
it should be borne in mind that the mandible almost faces the labrum.
In some genera, as in Coronula,[34] the swelling near the basal margin
of the labrum (Pl. 26, fig. 3, _k_), caused by the internal concavity
for the above muscle, is conspicuous. The depressor muscle is opposed
by a small elevator, attached to the mandible close by the depressor;
thence it runs upwards, and is united at its upper end to the base of
the palpus, at the point where the latter adheres to the labrum: I have
ventured to call this muscle the elevator, from being apparently so
well fitted for this purpose; but I feel some little doubt, from having
observed an apparent slight movement in the palpi of living Balani;
and this is the only muscle entering those organs. The free part of
the mandible is articulated on a square, thickened piece of membrane,
forming part of the side of the mouth (Pl. 26, figs. 3, 4, _c^1_;
and Pl. 10, fig. 1, _a_, _b_, in my volume on the Lepadidæ); to this
square piece of membrane, two short muscles are attached, one above the
other, and which ought, in the Plate in my former volume, to have been
represented crossing the depressor muscle at nearly right angles; at
their further ends they are attached to about the middle of the labrum,
where, at least in Coronula (Pl. 26, fig. 3, _i_), a slight concavity
can be detected. The action of these two muscles must be to draw the
whole mandible against the labrum; and the depressor muscle might, at
the same time, draw the toothed edge downwards, and thus force any prey
into the œsophagus.
[34] This is figured by Burmeister in his 'Beiträge zur
Naturgeschichte der Rankenfüsser,' Tab. 2, fig. 6.
The inner maxillæ are likewise furnished with four muscles, very
nearly as figured in my former volume (Pl. 10, fig. 10); namely, two
muscles, one inside and the other outside the curious apodeme, which
in the Balanidæ (Pl. 26, fig. 7, _b′_) is as invariably present as in
the Lepadidæ: these two muscles are attached at their lower ends to
the outer membrane of the mouth, close to its basal articulation: the
outer one of these two muscles would, I presume, act as an elevator,
and the inner one as a depressor; the free part of the organ working on
the top of the apodeme, like an axe, on a hinge, on the top of a pole.
But there is also a larger depressor muscle, in an analogous position
with that (_i. e._ the first-mentioned muscle) of the mandibles; and a
fourth muscle, crossing the latter depressor at nearly right angles,
and attached (as far as I could make out) on the side of the orifice
of the œsophagus, close under the mandibles: the action of this latter
muscle would be to draw the whole organ towards the labrum.
I must not conclude my description of the mouth, without drawing
particular attention to its peculiar compounded nature. It is
prominent, and is capable, as a whole, of movement; it is separated
from the body by a fold or articulation, which can be traced all
round. It is, as we have seen, composed of a broad labrum and three
pairs of gnathites, which latter have only their terminal segments
free; and these surround a conical hollow, at the bottom of which
lies the opening of the œsophagus. The prominence of the whole mouth
appears to result from the lateral fusion of the two basal segments of
the three pairs of gnathites. I have examined the mouth of ordinary
Crustaceans, and can see no trace of a structure like this. That there
has been some union of the parts is indisputable; for the palpi, which
in ordinary Crustaceans are quite free, are here firmly united to the
upper and outer corners of the labrum; and indeed, at first appear to
be more intimately connected with the labrum than with the mandibles.
The palpus on its upper and exterior surface, is in direct continuity
with the square thickened piece of membrane, on which the mandible is
articulated, and likewise with that side of the upper or free portion
of the mandible which faces the labrum. This face of the mandible,
beneath the toothed edge, is hollowed out or arched (Pl. 26, fig. 5,
_p_), owing to the above-mentioned continuity of its membrane with
that of the palpus. On the lower surface, the palpus is firmly united
to the lateral corners of the labrum; or indeed the corners of the
labrum may be almost said to be formed by the soft, swollen bases of
the palpi: the point of union, when viewed from the outside, is seen
to form a knob on the shoulder of the labrum, beneath the level of
its crest, and at this knob (Pl. 26, fig. 3, close to _d′_) several
thickened bands in the surrounding membrane unite. The free portion of
the palpus stands out transversely behind (_i. e._ anteriorly to, in a
homological sense) the labrum. I suspect that the palpus possibly may
consist of two segments, of which the terminal one is free, and the
lower one confluent with the labrum.
Before proceeding any further, I should observe that figs. 3 and 4, in
Pl. 26, represent the membranes of the mouth of _Coronula diadema_,
perfectly cleaned. In fig. 3, all the front part of the mouth has been
removed, the mandible on one side, the labrum with the two palpi, and
the œsophagus being alone left, and these are viewed from the inner
side; the front part, however, of the supra-œsophageal cavity has been
cut away. In fig. 4, the labrum, with the œsophagus, has been removed,
whilst the two outer maxillæ, the right-hand inner maxilla and mandible
(with the exterior and basal portions, _d_, _d′′_, of one palpus) are
seen from the outside; but in order that these parts should all be
shown, the whole of the right-hand side of the mouth has been spread
out, for the teeth of the mandible should have stood in a vertical
line between the two outer maxillæ. In the mandibles, the free upper
part is separated, by a distinct articulation, from the square piece
of thickened membrane (fig. 3, _c^1_) on which it is supported; and
this latter is separated by a second articulation from a portion of
thickened membrane (_c^2_), the basal edge of which forms the third and
lowest articulation, separating the mouth from the body. This basal,
thickened portion of membrane curls round and inwards, towards the
outer maxillæ or front of the mouth, and its terminal points sometimes
even penetrate a little way within the muscles, like apodemes: it is
not distinctly separated by any line or suture from the membrane, which
forms the whole broad labrum; so that I at first concluded that the
labrum dipped under the mandibles, and thus afforded a support on which
they were articulated; but this appears so opposed to all analogy,
that it is more probable that the above basal thickened portion of
membrane is truly the basal segment of the mandibles, completely
confluent with the labrum; and it is, I think, not very improbable
that even a large portion of what in appearance belongs to the labrum,
namely, those concavities to which the muscles of the mandibles are
attached, may, also, be part of the basal segment of the mandibles.
Whether or no there really are two segments beneath the upper free
portion of the mandibles, which have become laterally confluent with
other parts, I must think that the square thickened piece of membrane
(_c^1_) represents at least one segment. I may here observe, that Prof.
Milne Edwards seems to consider the mandible of the higher Crustaceans
as answering homologically to the haunch of the leg; but, according to
M. Brullé,[35] there ought to be two basal segments (sous-maxillaire
and maxillaire) bearing the proper mandible, and giving rise, on the
outer side, to the palpus,--a structure which perfectly corresponds
with my view of the mandible and palpus in Cirripedes.
[35] 'Annales des Scienc. Nat.,' 3d series, Zoolog., tom. ii, p.
271.
_Maxillæ_: the point whence the long apodeme (_b′_, Pl. 26, fig.
4 and fig. 7) arises, according to Audouin's views, must mark an
articulation, and this would separate the upper free segment from the
lower segments, which I believe to be laterally confluent with the
organs on each side. The thickened membrane, of which the upper free
part is formed, extends a little distance beyond the insertion of the
apodeme; and this small portion beneath the point of insertion may
possibly answer to the square, thickened piece of membrane, or second
segment, supporting the mandibles. Beneath it, a rather wide expanse of
thin, flexible membrane reaches down to the basal fold surrounding the
mouth, and may thus form the third segment.
_Outer Maxillæ_: the upper free segment has a spinose lobe (_a′′_,
Pl. 26, figs. 2 and 4), on its inner face, which may indicate a lower
and second, almost free segment. Passing over this, we have, on the
outside of the mouth, beneath the free, upper segments, an expanse of
membrane, which, on the side, close to the inner maxillæ, is perforated
(Pl. 26, fig. 4, _n_) by orifices which I believe are olfactory. In
some species, as in _Bal. eburneus_ and _improvisus_, there is a
longitudinal medial suture in this expanse of membrane, which I suppose
indicates the lateral confluence of the middle segments of the two
outer maxillæ. A short, transverse articulation or fold separates this
middle segment (fig. 4, _a^1_) of each maxilla from the third or basal
segment; and this latter (_a^2_) is separated from the body by a very
distinct fold, which (at least amongst the Lepadidæ) sends inwards a
short, medial, tongue-formed apodeme. Here, then, we apparently have,
as in the mandibles, two segments under the upper free segment of
each outer maxilla, laterally confluent with the adjoining organs.
But I must state that, in old specimens, and only in old specimens of
_Coronula diadema_, I have found under the outer maxilla an additional
transverse ridge and fold, which plainly shows how easily a mere
thickening of the membrane might be mistaken for an articulation. I
can, however, hardly persuade myself that the articulated membrane,
under the free part of the mandibles, which has now been figured and
described, has no homological signification; and the fusion of the
palpus and labrum seems too plain to be mistaken. Hence I must conclude
that the mouth, in the Cirripedia, does truly exhibit a compounded
structure of a very peculiar nature.
_Cirri._
There are always six pairs; each biramous and multiarticulated,
supported on a pedicel formed of two segments. A shield-like swelling
at the exterior bases of these pedicels often appears like another
segment; but such, I believe, is not its nature. The five posterior
pairs answer to the five pairs of ambulatory legs in the higher
Crustaceans; and as in the case of the latter, the three, or the four
hindermost pairs almost invariably resemble each other. The first
pair, which is homologous with the outer maxillipeds of ordinary
Crustaceans, is separated by an interval from the second pair;--though
this is not the case with the legs of the pupa, from which the cirri
are metamorphosed. These anterior cirri are attached to the lateral
edges of the mouth, namely to the thickened rim of membrane, forming
the supposed basal segment of the mandibles. They are capable of more
diversified movements than the other cirri: the anterior ramus is
always elongated, with the terminal segments more or less tapering,
and is directed beyond (or anteriorly to) the mouth: the shorter
ramus closely resembles in structure the rami of the second pair.
In the Chthamalinæ the second pair, and in the Balaninæ the second
and third pairs (as will be more particularly described under these
sub-families) differ in structure from the posterior pairs, from which
they are separated by a slight interval. The number of segments on
the posterior cirri is often great, amounting in Chelonobia even to
fifty. Each segment normally is furnished on its inner face, which
is usually somewhat protuberant, with from two to rarely eight or
ten pairs of long spines or bristles, placed in a double row; the
two spines in the lower pairs stand nearer to each other, and are
shorter than the spines in the upper pairs. Between each pair of
spines there is either a single, very thin bristle, or often a tuft of
such. The pairs are directed somewhat upwards, and they diverge when
the cirri are uncurled; their function is obviously to entangle the
prey. On the dorsal or exterior surface of each segment, close to its
upper margin, there is a tuft of spines, often composed of thicker
and thinner spines; these, I believe, serve to prevent any creature
intruding within the sack. On both sides of the upper margin of each
segment, there is generally a row of short, blunt, excessively minute
spines, which only deserve notice, inasmuch as it is by their increase
in number and size, and by the spreading out of the dorsal tufts,
and, lastly, by the increase of the little tuft intermediate between
the pairs of spines situated in front, that the segments on the two
or three anterior pairs of cirri become covered, like brushes, with
bristles. The bristles or spines on the second and third cirri are
often, especially in Tetraclita, doubly and coarsely pectinated. The
bristles on the pedicels follow the same arrangement as on the rami;
namely, being in regular pairs on the posterior cirri, and crowded
thickly, like a brush, on the anterior cirri. The segments in the
shorter ramus of the first cirrus, and in both rami of the second,
and often of the third cirrus, are broader than the segments of the
posterior cirri; they are, also, especially in the genus Balanus,
frequently produced in their upper, ante-lateral corners, into
remarkable prolongations (see Pl. 29, fig. 4, of the third cirrus of
_Bal. perforatus_), clothed on their inner surfaces, and at their
extremities, by numerous bristles. The number of the segments in each
cirrus is in some degree variable, and increases with age; this is
likewise the case, to a certain extent, with the number of the spines
borne on each segment.
As compared with ordinary Crustaceans, I presume the two rami answer
to the "_tige_" and "_palpe_" of Milne Edwards; and the pedicel (as I
have called it) to the two basal segments of the leg.[36] The "_fouet_"
or flabellum does not appear to be developed in any Cirripede; for
though the filamentary appendages in certain genera of Lepadidæ, might
at first be thought to be of this nature, yet their usual position
_beneath_ the basal articulation of the first pair of cirri, and the
occasional presence of more than one, proves, I think, that such is not
the case.
[36] According to this author's new nomenclature, the pedicel would
consist of the coxopodite and basipodite; the tige would be the
ischiopodite and following segments; and the palpe would be the
exopodite; the epipodite or flabellum being absent. ('Annales des
Sciences Naturelles,' tom. xviii, 1852.)
Though the structure of the cirri is very uniform, yet we meet with
some peculiarities. In Chelonobia, the segments of the posterior cirri
bear only two pairs of main spines; whereas in some varieties of
_Balanus balanoides_, they carry as many as ten pairs in a longitudinal
row; but in this latter species, the number of these spines varies,
in a singular manner, from six to ten pairs. In Tubicinella, the
pairs of spines on the segments of the posterior cirri are arranged
so closely one under the other, that they appear almost like a single
transverse row. Considering the whole family, the third pair of cirri
differs most in structure in the different genera. Thus, in _Chthamalus
antennatus_, the anterior (or outer) ramus (Pl. 29, fig. 3) is thicker
and much longer than the posterior (or inner) ramus; the number of the
segments in one instance being, in the two rami, 53 and 18; in the
longer ramus, the spines are arranged abnormally, tending to form a
little circle round each segment; and the whole ramus may be said to be
antenniformed, and I believe acts as an organ of touch: the relative
number of the segments, I may add, in the two rami and the arrangement
of their spines varies greatly in this species. In two other species of
the same genus Chthamalus, we have _occasionally_ the anterior ramus in
some degree antenniformed, so that this whole structure is variable.
In the allied _Chamæsipho columna_, it is the posterior or inner
ramus which is antenniformed, but this peculiar development is more
plainly marked in the case of the second pair of cirri than in that
of the third pair. In _Tetraclita porosa_ it is, also, the posterior
ramus of the third pair which is antenniformed; in this third pair,
and indeed in the other cirri, the relative numbers of the segments
vary extremely. A similar structure in the third pair, but in a lesser
and variable degree, may be observed in some of the other species of
Tetraclita. In _Balanus vestitus_, also, we have, in the third pair,
an analogous structure. It is scarcely possible to believe that the
circumstance of the second pair of legs, which answer to the third pair
of cirri, being antenniformed in certain decapod Crustaceans, is an
accidental coincidence; it must be owing to some special affinity in
the two groups.
In Chelonobia, the third pair of cirri is of unusual length compared
with the second pair, but does not otherwise differ from the type
of its sub-family: in Coronula and its allies, on the other hand,
the third pair is very short and broad, as may be seen (Pl. 29, fig.
5) in Xenobalanus: in this latter genus, the front surfaces of the
segments of the pedicels (fig. 6) of the posterior cirri, are extremely
protuberant, almost as in _Scalpellum vulgare_.
The last peculiarity in the cirri at all worth mentioning, is in
the sub-genus Acasta, in which, differently from in all other known
Cirripedes, the anterior ramus of the fourth pair does not absolutely
resemble the rami of the fifth and sixth pairs; in most of the species,
the spines on this anterior ramus are more crowded together, are
larger, and are mingled with some short thick points; and the spines
in the dorsal tufts are also longer than in the two posterior pairs of
cirri; but in _A. sulcata_ (Pl. 29, fig. 2), and in a lesser degree
in _A. cyathus_ and _A. purpurata_, the front margins of the lower
segments of this anterior ramus, and of the upper segment of the
pedicel, are developed into strong, downwardly curved teeth: it is very
remarkable that so beautiful a structure should be extremely variable,
as it certainly is in _Acasta sulcata_.
_Caudal Appendages._
With extremely few exceptions, these are present in all the Lepadidæ
and Verrucidæ; whereas amongst the Balanidæ they occur only in the
two species of Pachylasma, and in one species of Catophragmus;
these being the genera most closely allied to the Lepadidæ, and
where, consequently, their presence might have been anticipated.
These appendages are seated close together over the anus; they are
multiarticulate, each segment being sub-cylindrical, with a few small
bristles round its upper edge.
_Alimentary Canal._
I have not much on this head to add to what I have said under the
Lepadidæ. As in that family, the strong internal membrane of the
œsophagus terminates in a remarkable, bell-shaped expansion (Pl. 26,
fig. 3, _g′_), which, as observed by M. St. Ange, serves to keep
the upper broad end of the stomach expanded. The œsophagus is well
furnished with constrictor and radiating muscles for closing and
opening it; and it is thus capable of a strong swallowing movement.
The stomach runs down to the lower end of the prosoma, and then
doubling back on itself extends to the anus. As the prosoma is much
elongated in Tubicinella and Xenobalanus, so is the stomach of
unusual length in these genera. In several species of Balanus, the
upper edge of the stomach is surrounded by from six to eight cæca;
these cæca I ascertained, in _Balanus perforatus_, are branched, and
penetrate a considerable way into the body; and some of them at least
expand a little at their extremities. Each cæcum, from the manner in
which it retained fluid, must, I think, be furnished, at the point
where it enters the stomach, with a sphincter muscle. In Tetraclita,
Chthamalus, Tubicinella, Coronula, and Xenobalanus, there are no cæca;
but in Xenobalanus and _Coronula balænaris_, there are longitudinal,
approximate folds in the upper, broad end of the stomach, which would
serve to expose the food to a greater extent of digesting surface.[37]
[37] The presence and absence of these cæca in genera so closely
allied as Balanus and Tetraclita, shows, I think, that these
cavities are not of high importance; and I must doubt whether Von
Siebold's view ('Anatomie Comparée,' tom. i, p. 445), that these
cæca form a passage to a true or isolated liver, such as exists in
the higher Crustacea, can be admitted. Cæca are said by Von Siebold
to occur in some of the Entomostraca, as Daphnia, Argulus, &c.
As in the case of the Lepadidæ, a transparent, structureless,
epithelial tube, composed of chitine, containing more or less digested
food, is found, in specimens preserved in spirits, occupying the whole
length of the stomach, and where there are cæca, sending branched
prolongations into them. It does not extend into the œsophagus or into
the rectum. This epithelial tube or model of the stomach, filled with
excrement, is expelled by the rectum, whole, that is in a single piece,
as I observed in some living specimens of _Balanus balanoides_: in
some specimens, however, of _Chthamalus stellatus_, the excrement was
ejected, perhaps from the animal being confined, in fragments, and the
sack thus became befouled. Beneath the epithelial layer, the stomach is
lined by a delicate, pulpy and cellular mucus layer, which easily peels
off in flakes: this is surrounded by a muscular layer with the fibres
closely approximated and transverse; and this by a layer of stronger,
longitudinal muscles, but more distant from each other. Lastly,
outside this double muscular layer, there is a rather thick, somewhat
laminated, pulpy layer, abounding with cells, often nucleated, and
frequently containing much oily matter. This structure agrees closely
with Dr. C. H. Jones's[38] account of the external covering of the
stomach in Daphnia, and which he believes to be hepatic: as in Daphnia,
there does not seem to be any ducts. I may here observe, that within
the upper part of the prosoma, but not immediately connected with the
stomach, I have often observed much white pulpy substance, permeated by
lacunal passages, and exhibiting no structure except some excessively
minute cells.
[38] 'Philosophical Transactions,' 1849, p. 116. Karsten ('Nov.
Actorum Acad. Nat. Cur.,' 1845, tab. xx) has excellently figured
the testes, as the hepatic glands; and has indicated the ovaria
as salivary glands; it is singular that this anatomist overlooked
the ducts which lead from his supposed hepatic glands, into the
vesiculæ seminales, within which he observed spermatozoa.
The rectum, lined by membrane continuous with that investing the
thorax (and seen through it, in Pl. 26, fig. 8, _c_), extends inwards
to about opposite the bases of the third or fourth pairs of cirri.
It is longitudinally plaited; the ends of the folds forming a sort
of valve where joined on to the stomach. It is coated by circular,
transverse muscular fibres: judging from the movements, the anus
itself is surrounded by a strong sphincter muscle. The anus opens on
the dorsal surface of the thorax (fig. 8, _b_); but as in the genera,
in which caudal appendages occur, it opens under them, the orifice, I
believe, is homologically terminal, and owes its dorsal aspect to the
aborted state of the whole abdomen, and to the great development of the
probosciformed penis; for the anus may be said to be situated on the
dorsal base of this organ.
Altogether we see that the alimentary canal is of a very simple
structure. The food, judging from the contents of the stomach, seems
generally to be composed of infusoria and minute animals: but in the
case of Tetraclita, I have been surprised at the size and number of the
included amphipod, isopod, and entomostracan Crustaceans, in one case,
together with an annelid. I have, also, sometimes seen some confervoid
matter within the stomach.
_Circulatory System._
On this subject I can add nothing, except to express my conviction that
there is no heart, or true vessels; the circulation being strictly
lacunal. A passage has often been quoted from Poli, in which he states
that he saw a pulsating organ, close above the anus; but I have seen
this movement, which appeared to me to be a convulsive twitching of the
sphincter muscle of this orifice. The largest lacunal channel extends
down the middle of the rostral compartment of the shell: and this
answers to the rostral channel down the peduncle in the Lepadidæ. Large
nerves and the main pair of unbranched ovarian tubes (Pl. 25, fig. 1,
leading into _g_) extend along this channel. At the basis (at least in
Coronula) this channel joins on to a large circular one, running all
round the sack, and sending off branches into the mass of ovarian tubes
and cæca.
_Nervous System._
It has been shown in my former volume, that in Lepas and in some
other genera of the Lepadidæ, there are six main ganglions; one
supra-œsophageal, and five infra-œsophageal, or thoracic. In
Pollicipes, however, there are only four thoracic ganglions, the
last ganglion supplying the three posterior pairs of cirri with
nerves, whereas in the other genera, the last ganglion supplies only
the fifth and sixth pairs of cirri. In this genus, moreover, the
lateral fusion of the ganglions has been so complete, that there is
no evidence of their having been formed by the union of two. Amongst
sessile cirripedes, we discover evidence of much higher concentration
even than in Pollicipes. My chief examination has been confined to
_Coronula diadema_, and to _Balanus tintinnabulum_: and in these
genera we find (and the fact appears to me highly remarkable) as high
a degree of concentration in the infra-œsophageal ganglion as in any
decapod Crustacean, for instance, as in Maia, judging from the figure
given by Milne Edwards; for all the nerves, with the exception of
those connected with the supra-œsophageal ganglions, radiate from a
single great ganglion.[39] The nervous system is, moreover, otherwise
complicated.
[39] It must, however, be observed that, according to Mr. Dana,
there is in certain suctorial Entomostracans, as in Caligus,
only one infra-œsophageal ganglion. Mr. Dana speaks of this as
resulting from reduction. In Cirripedes, from the gradation which
may be observed from Lepas through Pollicipes into Balanus, the
ganglions are certainly not reduced but concentrated. In Van de
Hoeven's figure of the nervous system in Limulus, there is seen to
be no chain of thoracic ganglions; all the nerves rising from the
circa-œsophageal collar; but this, on the other hand, seems hardly
developed into a ganglion.
To begin with _Coronula diadema_ the great infra-œsophageal ganglion
(Pl. 27, fig. 1, A) is seated nearly opposite to the anterior margin of
the second pair of cirri, which are homologous with the first pair of
legs in the decapod Crustaceans. This ganglion shows no trace of any
longitudinal medial suture; its shape is hardly discoverable, for it
is formed by the union of eleven principal pairs of nerves, besides
several arising from its under surface; in outline, however, it may be
said to be divided into a posterior and anterior half; the latter being
somewhat heart-shaped, and the posterior half elongated. The nerves
going to the five posterior pairs of cirri rise from the posterior
margin of the ganglion, and run for some distance in a sheet, parallel
and close together; the pair, however, going to the second pair of
cirri soon branches off from the others. Each of these nerves enters at
the inner and posterior margin of the cirrus to which it belongs, and,
at least in the case of the first pair, divides into two branches as it
enters. The nerves (Pl. 27, fig. 1, _r^5_, _r^6_) going to the fifth
and sixth pairs of cirri are more closely united together than are the
others, and appear, till they branch off, like a single large nerve.
That which belongs to the sixth cirrus gives off, opposite to the
fifth cirrus, a branch (_s_) nearly as large as itself, which enters
the probosciformed penis. I may remark, that homologically this is the
only abdominal nerve in any cirripede of the Order. From the under side
of the nerves which run to the five posterior pairs of cirri, small
branches are given off, extending dorsally into the thorax.
The anterior end of the great infra-œsophageal ganglion is formed by
the union of a set of nerves, extending parallel in a bundle in a
directly opposite direction to those running to the five posterior
pairs of cirri. These nerves consist of an outer larger pair (_r^1_)
entering the first pair of cirri; and within these, and rather dorsally
to their roots, we have the circa-œsophageal chord (_c_, _c_), or
collar nerve; between the roots of the latter, and on the ventral
surface (or near side of the figure), there are three closely united,
small pairs, running to the gnathites, and, as I believe, to the
olfactory sacks. From the under (or dorsal) surface of the anterior
end of the ganglion, two nerves, larger even than the circa-œsophageal
chord, and which I shall call the splanchnic pair (_d_, _d_) arise;
and the singular course of these nerves will presently be described;
between this great pair, there is a single (_b_) medial nerve, which
runs down and branches into that large diverging muscle, which is
attached to the upper ventral surface of the stomach. Posteriorly to
these three nerves, we have two pairs of much smaller nerves (not
figured), running dorsally into the body, so that we have seven nerves
rising from the under surface of the infra-œsophageal ganglion. I
need only further add, that on each side of this ganglion, between
the nerves going to the first and second pairs of cirri, there is a
moderately sized nerve (_k_), which appeared to run into the muscles of
the thorax: a nerve in a similar position is figured by Milne Edwards
in Maia.
The circa-œsophageal chord (_c_, _c_) nearly equals in length the whole
distance from the centre of the main ganglion to the posterior end
of the thorax. This collar bows out on each side, where passing the
œsophagus (_œ_), which is seated at its anterior end. From the collar a
branch is given off on each side, which I traced as far as between the
mandibles and maxillæ; from analogy with other Crustaceans, it perhaps
runs to the mandibles. The collar has not a transverse commissure, such
as described by Milne Edwards in the Podophthalmia, and as figured by
Van de Hoeven in Limulus.
The supra-œsophageal ganglions (B) present a singular contrast with
the infra-œsophageal ganglion in their little development, size, or
degree of confluence. They lie directly under the basal edge of the
labrum. They are laterally quite distinct, and consist merely of a
slight enlargement of the circa-œsophageal chord. From the anterior
edge of each ganglion, a broad nerve (_f_) extends for some distance in
a straight line, and, on close examination, can be seen to be formed of
two nerves closely united, of which the inner and smaller one, after
a space, appears to cross over the larger nerve: both become at this
point tortuous, and, giving off branches (_m_, _m_), form a plexus. The
two nerves (_f_) then bend inwards, and almost touching each other, run
down, together with the two ovarian simple ducts, along the rostral
compartment of the shell. No doubt, if the smaller branches from these
nerves could be traced, they would be seen to form a network over the
whole sack; and would therefore enclose, as in a cage, the rest of the
nervous system. These nerves correspond, I believe, to the two pair
of antennular nerves of ordinary Crustaceans, and hence I will call
them by this name. Just in front, at the outside corners of the two
supra-œsophageal ganglions (B), a branch (_e′_) arises, which I traced
to the ends of the adductor scutorum muscle, and to those several
muscles which serve to retract the interspace of membrane between the
mouth and the adductor.
The pair of great splanchnic nerves above alluded to, which arise from
the anterior and dorsal surface of the infra-œsophageal ganglion, are
in Pl. 27, fig. 1, _d_ _d_, (and in fig. 2), laid flat; but in nature
they first bow outwards, and then, penetrating deeper into the body,
approach each other, and running nearly parallel, pass round the lower
end of the œsophagus: their course consequently is nearly similar
to that of the circa-œsophageal chord, with this difference, that
the outwardly bowed portion is situated near the infra-œsophageal,
instead of near the supra-œsophageal ganglion. The splanchnic nerve,
a little beyond the supra-œsophageal ganglion, joins a plexus (_d′_);
and into this plexus another large nerve (_e_) which I will call the
supra-splanchnic nerve, sends branches; this nerve takes an almost
semicircular bend over the ovarian glands. The supra-splanchnic nerves
(_e_, _e_), though appearing to spring from the supra-œsophageal
ganglions, do really arise, as may be seen by tracing the constituent
fibres, from the circa-œsophageal chord. The plexus (_d′_) lies close
to the coats of the upper end of the stomach: several branches,
proceeding from it, run further on, but I was able to trace only a few
of them: one went (at least in the case of _Balanus perforatus_), to
the adductor scutorum muscle: another branch spread out on the flanks
of the prosoma: I strongly suspect that one branch goes to the acoustic
sack: it appeared, also, as if some of the small branches entered the
second plexus (_m_), where the inner antennular nerve and ophthalmic
nerve cross over the outer antennular nerve.
I have called the nerves (_dd_, _ee_) splanchnic and supra-splanchnic,
from their course and apparent function in supplying the viscera. In
the descriptions of the nervous system of other Crustaceans I can find
nothing analogous to my great splanchnic nerve (_dd_); the so-called
supra-splanchnic nerves (_ee_), which arise from the circa-œsophageal
chord, seem to be the analogues of the ordinary splanchnic nerves,
though these latter are always described as uniting into a single
medial branch. The plexus (_d′_) is the cervical ganglion of M.
Martin St. Ange,[40] who has likewise indicated the course of my
splanchnic and supra-splanchnic nerves; but the plexus, when viewed
as a transparent object, hardly appears to me to be ganglionic in its
nature. In my former volume on the Lepadidæ, I quite misunderstood the
course of these splanchnic nerves.
[40] 'Mémoire sur l'Organisation des Cirripèdes,' p. 19.
From the commissure between the two supra-œsophageal ganglions, a
straight chord (Pl. 27, fig. 1, _g_) arises, which terminates in
a small ganglion (C), barely exhibiting traces of being formed of
two laterally confluent ganglions. This is the ophthalmic ganglion.
The chord connecting it with the two supra-œsophageal ganglions is
accompanied by a small nerve (_h_) which runs on to the muscles round
the adductor scutorum muscle; the chord is encased by much fibrous
tissue, and its dissection is thereby rendered difficult. From the
ophthalmic ganglion, on each side, a nerve (_i_) goes forth and crosses
the antennular nerve; these, if I could have traced them, would have
been found to run, as may be safely inferred from what is known in
_Balanus tintinnabulum_, to a pair (D, D) of eyes.
In _Balanus tintinnabulum_, the structure of the great infra-œsophageal
ganglion (Pl. 27, fig. 2, A) is essentially the same as described under
Coronula. The great pair of splanchnic nerves springing from its under
side, are here actually twice as large as the circa-œsophageal chord.
The plexus (_d′_) formed by the splanchnic nerve (_d_), on each side,
with the supra-splanchnic nerve (_e_), which arises close posteriorly
to the supra-œsophageal ganglion, is here much less complicated, but
is perfectly distinct; and there was no appearance of the cervical
ganglion of M. Martin St. Ange. The chord (_g_) running from between
the two supra-œsophageal ganglions to the ophthalmic ganglion, is
nearly as large as the double antennular nerve (_f_) on each side of
it. The ophthalmic chord (which is accompanied in its whole course by a
small branch running to the adductor scutorum muscle) terminates in a
small ophthalmic ganglion (C), which seems to be formed by the almost
complete fusion of two ganglions. This ganglion is hardly larger than
the chord which it terminates: it appeared to me to give rise to more
than one pair of nerves, and a single nerve (to my surprise) joined the
branch just mentioned, which goes to the adductor scutorum muscle.
From each supra-œsophageal ganglion, two closely united antennular
nerves (_f_) extend, of which the inner one crosses over the main
or exterior nerve, nearly opposite to the ophthalmic ganglion, and
here forms (_m_) a plexus. The structure of this plexus I was not
able, any more than in Coronula, to make out thoroughly, but I traced
quite distinctly a long nerve (_i_) running from it into what must be
considered as the eye. As in the case of Coronula, I traced a nerve
on each side from the ophthalmic ganglion into the plexus, where I
lost it; and as here in Balanus, I saw on each side of the ophthalmic
ganglion a cut off nerve, of about the size of that which runs from
the plexus on each side into the eye, I think we may safely conclude
that the latter or optic nerve does really arise from the ganglion here
called ophthalmic. I may add that the analogy of the nervous system in
the Lepadidæ most strongly confirms the view of this latter being the
ophthalmic ganglion.
_Eyes and Vision._
The optic nerve (_i_), running from the plexus to the eye, is of
considerable size; it runs nearly parallel to the main antennular
nerve, diverging from it a little. It retains nearly the same diameter
throughout; and gives off only one single, small, inner branch. It can
be traced beyond the basal edges of the scuta, to just under the upper
edge of the transparent opercular membrane, which unites the scuta
to the sheath of the rostrum. The nerve itself, at a little distance
from its further end, was, in a full-sized specimen, 5/1000ths of an
inch in diameter; widening a little, it expands slightly, and abruptly
terminates in a circular disc, about 8/1000ths of an inch in diameter,
(see Pl. 27, fig. 5). The nerve just beneath this slight expansion,
is coated all round by pellets of dark purple pigment-cells, but not
actually united into a continuous layer. These pigment-cells are
the more conspicuous from the surrounding parts being colourless. I
could not make out distinctly any cornea; and I suppose the external
transparent membrane, to which the above slight circular expansion is
attached, acts as such. This description very closely agrees with that
given of these organs in _Bal. rugosus_ of Gould, (_B. crenatus_?) by
Dr. Leidy,[41] who first discovered the eye in the adult cirripede,
but he did not observe the ophthalmic ganglion. These eyes differ from
those in some of the genera of the Lepadidæ, only in the greater length
of the optic nerve, and by standing laterally further apart from each
other.
[41] 'Proceedings of the Acad. Nat. Sciences of Philadelphia,'
vol. iv, 1848, p. 1. I may add that I have, also, observed the
supra-œsophageal and ophthalmic ganglions in _Bal. perforatus_.
I may here mention that I tried a few simple experiments on the senses
of _Balanus balanoides_, _B. crenatus_, and _Chthamalus stellatus_.
I found these three species very sensitive to shadows, that is, to
an object like my hand passing even quickly, and at the distance of
about a foot, between them and the source of the light.[42] They were
indifferent to a gradual change from bright to obscure light; but
instantly perceived and drew in their cirri, when my hand was passed
between the basin in which they were kept and the window, even when
this was tried rather late on a dusky evening; and likewise when my
hand was passed between them and a single candle. I took, of course,
the precaution of passing my hand in other directions, but this
never produced any effect. These species are moderately sensible to
any vibration in the vessel in which they were kept, but they were
indifferent to noises made in the air, or in the water. I found it
impossible to touch, under water, an individual shell ever so lightly
with a needle, without all the immediately surrounding individuals,
when several adhered together, perceiving it, and retracting their
cirri: it made no difference whether the one touched had already
withdrawn its cirri and was motionless: from this fact, and from seeing
that a similar but slighter effect was produced by touching the rock
on which the specimens adhered, I infer that the perception by the
others of the one being touched, is communicated by vibration. When an
individual was touched under water, not by a needle, but by a pointed
camel-hair brush, it generally withdrew its cirri, but the neighbouring
specimens took no notice: when touched by a single hair of the brush,
no notice was taken, unless the skin of the orifice leading into
the sack was so touched. In these trials, it is of course necessary
carefully to avoid intercepting the light. I could not make out that
cirripedes perceived odours diffused in the water.
[42] I find that this fact was long ago observed by Von Siebold,
'Anatomie Comparée,' tom. i, p. 434.
_Acoustic Organs._
These are situated in the same position as in the Lepadidæ, namely,
in a slight swelling on the sides of the thorax (Pl. 25, fig. 1,
_d′_) just beneath the basal articulation of the first pair of cirri.
The orifice in Tubicinella and Xenobalanus is slightly produced, or
is tubular; the free part in the former genus projecting 5/100ths
of an inch. The structure of all the parts is essentially the same
as in the Lepadidæ, but I think all are proportionally larger. The
external membrane of the body is turned inwards at the orifice,
as a short flattened tube, which widens considerably (being, in a
middle-sized specimen of Coronula, 4/100ths of an inch in width) before
it abruptly terminates. The meatus, as I have called the sack-like
cavity which encloses the true acoustic sack or vesicle, is formed
of pulpy membrane, and is apparently continuous with the corium of
the whole body, but by dissection it can be separated entire. The
acoustic vesicle is of various shapes, as we shall immediately see; but
in all essential respects it is identical with the same part in the
Lepadidæ; it is formed of the same peculiar, soft, elastic, brownish,
transparent tissue, which seems to be composed of fine, transverse
pillars, becoming towards the outside fibrous, and at their inner ends
appearing when viewed vertically from above, like hyaline points. In
_Coronula diadema_, I observed on the outside of the acoustic vesicle,
some excessively minute bristles, only 1/3000ths of an inch in length,
seated on little eminences. I examined carefully the contents of the
vesicle in this species, in specimens well preserved in spirits, and
there was nothing within but a very little, thin, pulpy fluid, and a
few yellowish nucleated cells, here and there aggregated into small
groups. In Coronula, the flattened acoustic vesicle is elongated, with
a somewhat sinuous, but not very irregular margin (Pl. 27, fig. 4),
and is without any ridges on the surface; its neck or orifice projects
at right angles to the elongated portion, which stands obliquely to
the tubular orifice of the meatus. In a moderately-sized specimen of
_Coronula diadema_, the elongated portion of the acoustic vesicle was,
6/100ths of an inch in length. In Tubicinella, the acoustic vesicle
is heart-shaped, with the neck attached to its broader end; and the
surface is covered by zig-zag ridges. In _Balanus tintinnabulum_ (fig.
3), the acoustic vesicle is almost square at the lower end, with the
neck placed at one of the upper corners; on the external surface,
there is an oblique prominent ridge or fold, which sends off downwards
another ridge; its length, in a large individual, was 5/100ths of an
inch.
In all these cases, the acoustic vesicle is mainly attached by its
neck, to the upper end of the sack-like meatus; but there is likewise
a layer of soft, pulpy, cellular matter, slightly connecting that side
of the vesicle which is opposite to the neck, with the walls of the
meatus or outer sack. The mouth or orifice of the vesicle is closed
by a delicate lid or diaphragm, which can easily be separated; and
this diaphragm is formed by the expansion of a large nerve, which here
abruptly terminates. In a very large specimen of _Coronula diadema_ I
clearly made out the existence of this nerve, and traced its course
for some distance from the point where the summit of the meatus and
the neck of the vesicle are joined together; the nerve first runs
posteriorly, and then turns inwards and doubles back or anteriorly;
and I clearly followed it to the antero-lateral sides of the uppermost
end of the stomach, where it seemed to enter a ganglion, so that I
unfortunately cut it off, but found only a slight plexus, with the cut
off nerve apparently running onwards with nearly the same diameter.
The diameter is great, fully equalling, in its widest part, that of
the circa-œsophageal chord; but it is very much flattened, and so
has not nearly so much bulk as that nerve. Before it reached the
stomach, it gave off one branch, which ran towards the mouth. The only
nerves which, from their size, could, I think, be continuous with
this from the acoustic sack, are the main branches proceeding from
that plexus (_d′_) formed by the interbranching of the splanchnic and
supra-splanchnic nerves.[43]
[43] I have always feared that anatomists would reject my view of
these organs being acoustic, owing to the absence of otolithes;
but I observe that so high an authority as Von Siebold ('Anatomie
Comparée,' tom. i, p. 433) does not believe that otolithes occur
in the acoustic organs even of the highest Crustacea. He considers
an "ampoule volumineuse, a parois mince, remplie d'un liquide
transparent," and a "membrane tympanique," though having a fissure
in the centre, as sufficient. I may here remark, that the nerve
proceeding from the acoustic vesicle in Cirripedes, and apparently
running to the splanchnic nerve, may easily be placed in connexion
with the antennular nerves, by the second plexus (_m_) in figs. 1
and 2, pl. 27. I should infer from Von Siebold's remarks on his
ampoule volumineuse in the higher Crustacea, that my acoustic
vesicle answered to the labyrinth in higher animals.
_Olfactory Sacks._
I can add nothing to the account given of these organs under the
Lepadidæ: I saw them in all the genera which I examined for this
object. In _Coronula diadema_ the orifices are large; they are seated
in the usual position (Pl. 26, fig. 4, _n_), in the confluent segments,
beneath the free part of the outer maxillæ, and somewhat exteriorly,
or as near as possible to the inner maxillæ. In no sessile cirripede
are the orifices produced or tubular, as is the case with several
genera amongst the Lepadidæ. I failed, as heretofore, in tracing with
certainty the nerve, which appears to enter the base of the sack, to
its ganglion.
_Male Organs of Generation._
All the Cirripedes of the family we are now describing, are bisexual
or hermaphrodite; and no instance has been observed of the presence
of males or complemental males. I have very little to add to the
observations made by M. Martin St. Ange and R. Wagner,[44] and to
those given in my former volume. The testes seem always to be confined
to within the thorax, including the prosoma. With their ducts, they
resemble club-moss or stag's horns, with the extremities a little
enlarged: a figure[45] of a small portion from _Balanus perforatus_ is
given in Pl. 25, fig. 2. It is quite surprising how like in structure
and appearance the branching ovarian tubes often are to the testes with
their ducts; but the latter are of smaller diameter. Two main ducts
generally unite just before entering the broad, often reflexed, end of
the vesicula seminalis: in _Coronula balænaris_, however, I observed
four ducts entering this receptacle. The two vesiculæ seminales, lying
within the thorax and prosoma, are usually very long and tortuous: they
are formed of a thin inner tunic, which is strengthened by thicker
reticulated lines, and of an outer layer of transverse fibres, which
are either elastic, or probably muscular, as they serve to expel
the contents with force when the end is cut off. The inner tunic is
prolonged up the probosciformed penis, at the base of which the two
vesiculæ unite.[46] The contents of the vesiculæ are commonly pulpy and
cellular; and from the cells the spermatozoa are developed; soon after
their development, they are, as it appears, expelled.
[44] The 'Report' on M. Martin St. Ange's memoir was laid before
the Academy of Sciences, July 14, 1834, so that I suppose it was
read previously to this date. R. Wagner's paper was published
in 'Müller's Archiv,' 1834, p. 467. Burmeister's 'Beiträge
zur Naturgeschichte der Rankenfüsser,' was published this
same year, 1834; so that these three authors published almost
contemporaneously.
[45] A far better figure is given by Karsten ('Nov. Act. Acad. Cæs.
Nat. Cur.,' 1845, Pl. 20, figs. 2, 3, 4), but under the erroneous
supposition that these organs were hepatic.
[46] In _Conchoderma aurita_, the ducts, as shown by Burmeister
('Beiträge,' &c. tab. ii, fig. 17), unite half way up the
probosciformed penis.
I have seen the spermatozoa in _Balanus crenatus_, _perforatus_, and
_balanoides_, and in _Chthamalus stellatus_. The cells, from which the
spermatozoa are developed, and which are often found in vast numbers
within the vesiculæ, are on an average about 1/5000th of an inch in
diameter. The spermatozoa differ remarkably within the vesicula of
the same individual, according to their state of development. I have
observed in _B. perforatus_ and in the Chthamalus, that the shortest,
and therefore, I presume, the youngest (Pl. 29, fig. 7, _a_), had a
globular head with no projection in front: as they increased in size,
this head became less in diameter, and a short tapering filament, (_a_,
_b_,) like the tail, projected out of it. This anterior filament does
not lie in exactly the same line with the posterior filament, which is
straight as an arrow. In _Bal. crenatus_, the anterior filament was
1/2000th of an inch in length, and the posterior filament 4/2000th,
giving a total length of 5/2000th: in the longest and best developed
specimens of _Chthamalus stellatus_, the nodular enlargement was
much elongated and spindle-shaped, and not above half the diameter
it had in the earliest stage; the posterior filament (measured from
the front of the enlargement, this consequently being included) was
5/2000th in length, and the front part only 1/4000th, giving a total
length of 11/4000ths of an inch. These observations agree pretty well
with Kölliker's;[47] but this author states, that perfectly developed
spermatozoa are absolutely without any nodular enlargement: if this be
the case, I have never chanced to see the spermatozoa in their perfect
condition. Mr. Bate, also, figures some (Pl. 29, fig. 7, _c_) in this
state, without any enlargement.
[47] 'Annales des Sciences Naturelles,' (2d series), tom. xix, p.
348. Kölliker refers to Wagner's paper on the same subject, in
Wiegmann's 'Archiv,' 1835, part ii, pl. iii, fig. 9. He also refers
to Von Siebold's observations. Mr. C. Spence Bate has figured, in
the 'Annals and Magazine of Natural History' (vol. viii, 2d series,
1851), the spermatozoa of _Balanus balanoides_, _perforatus_, and
of _Verruca (Clitia) Strömia_, and of these I have given copies,
Pl. 29, fig. 7.
The probosciformed penis lies adpressed on the under side of the
thorax, with its apex generally projecting between the first and second
pairs of cirri. It presents the same ringed or articulated structure as
in the Lepadidæ: it arises from an unarticulated projection or support,
which also forms the posterior border to the anus. This support
often terminates, as first observed by Poli, in a very sharp point;
but this point cannot be of much functional importance, for though
present in _Balanus balanoides_, it is absent in the closely allied
_B. crenatus_; in Tubicinella there is only a rudiment of this point;
I have not observed it in any member of the Chthamalinæ. The strong,
transverse and longitudinal muscles with which the penis is furnished,
are attached to this support. The apex or orifice of the penis is, I
believe, invariably surrounded by some bristles. Its length varies
much, according to its state of contraction or relaxation; and this
again, I believe, is dependent on the condition of the male secreting
organs. In a small specimen of _Elminius modestus_, the penis was
actually thrice as long as the whole thorax, including the prosoma:
in Pachylasma and in _Octomeris angulosa_, the penis is very short,
being equal only to once and a half the length of the pedicel of the
sixth cirrus: in _Octomeris brunnea_, the unarticulated support is
much elongated, being as long as the pedicel of the sixth cirrus, in
which respect this organ resembles that of _Ibla quadrivalvis_, and of
no other Cirripede. From the attachment of the penis at the posterior
end and on the under side of the anus--from the position of the caudal
appendages (where such occur) over the anus--from the position of
these same appendages in the pupa--and lastly, from the position of
the papilla-like penis in the abnormal _Proteolepas_, I infer that,
homologically, the penis is situated at the apex of the abdomen, on
its ventral surface; and that, consequently, this organ cannot be
considered as the abdomen itself in a modified condition.
_Female Organs of Generation._
I have scarcely anything to add to the statements in my former volume.
These organs consist of the true ovaria, or glandular bodies seated
on each side, not far from the basal edge of the labrum; of the main
or unbranched ovarian ducts; and of the (Pl. 25, fig. 1, _g_) ovarian
branching tubes and cæca. I traced distinctly in Balanus, Tetraclita,
and Coronula, the two main ovarian ducts, running from within the
prosoma to the layer of inosculating, branching, ovarian cæca[48] which
overlie the basis. In _Coronula diadema_ one of these main ducts was
1/100th of an inch in diameter. Though I traced these ducts near to the
grape-like, glandular masses,[49] which I cannot doubt are the true
ovaria, I did not succeed in tracing them into actual connection. As
in the Lepadidæ, these ovarian glands lie on the sides, near the basal
margin of the labrum, and almost under, but rather to the outside of
the antennular nerves. The branching and inosculating ovarian cæca
form a layer, which corresponds with the mass filling up the peduncle
in the Lepadidæ. In Tetraclita they do not cover the whole basis, but
are confined to the circumference; they, however, likewise extend
up between the two layers of corium round the walls of the shell,
and chiefly in the interspaces between the depressor muscles of the
opercular valves. In Chelonobia, they enter between the radiating septa
in the thickness of the walls: in _Coronula diadema_, they extend from
over the basal membrane into the six large square chambers (Pl. 16,
fig. 7, _v_) separating the radii and alæ: in Tubicinella they are
confined to the basis: in Xenobalanus, they form a layer over the basis
and likewise round the upper part of the peduncle-like body, which
answers to the shell of other sessile cirripedes.
[48] These are well described in Lepas, by R. Wagner, in 'Müller's
Archiv,' 1834, p. 467. Von Siebold, I observe, refers to Burmeister
as the first author who discovered the ovarian cæca within the
peduncle; I had thought that M. Martin St. Ange had a prior claim.
[49] These are obscurely figured by Karsten ('Nov. Act. Acad. Cæs.
Nat. Cur.,' 1845, Pl. 20, fig. 1_d_) as salivary glands; they were
so considered by Cuvier and M. Martin St. Ange: I may observe
that salivary glands have not been positively recognised in any
Crustacean.
As after the most careful and repeated examinations of various
Lepadidæ, I was convinced that there were no oviducts, so I have
come to a similar conclusion in regard to the Balanidæ; the ova
being brought to the surface, by the formation of a new membrane
round the sack underneath them, and by the subsequent exuviation of
the old membrane. The ova are united together by a most delicate
tunic investing each egg; the ovigerous lamellæ being thus formed,
as in the Lepadidæ. In the cases of _Chthamalus stellatus_, _Balanus
balanoides_, and _Platylepas decorata_, I saw a pair of very distinct
but fragile lamellæ. In Xenobalanus, the two ovigerous lamellæ form
two sub-cylindrical packets, pointed at their lower ends and often
cohering. There are no ovigerous fræna, for the attachment of the
lamellæ; the ova being sufficiently well retained, as it would appear,
by the well-closed shell. I have elsewhere stated my full belief that
it is the ovigerous fræna which have been metamorphosed into the
branchiæ of the Balanidæ. Most sessile cirripedes breed when very
young; and I have every reason to believe that they breed several times
in the year. The ova are ovate, and vary in length from 14/2000th of
an inch in Chthamalus, to 19/2000th in some species of Balanus, in
which this greater length was owing to a more elongated shape,--up to
25/2000th in some other species of Balanus. The ova are wonderfully
numerous, especially in the genus Coronula.
I may here mention the singular case of some elongated specimens
of _Balanus balanoides_, from Tenby, in South Wales: some of these
presented nothing abnormal; but in no less than seven specimens, the
two, three, or four posterior pairs of cirri, either on one or both
sides, were in an almost rudimentary condition, being of small size
and having a shrunk and wasted appearance. In six out of these seven
specimens, the probosciformed penis was quite short and abruptly
truncated, as if from abortion. By cutting off the truncated apex, and
cleaning the external tissue, I ascertained that it was imperforate,
apparently in all the cases, and I am certain of this fact in several
of the cases. In three of the specimens, I examined the vesiculæ
seminales; in one, I found some spermatozoa, but cohering together
in a peculiar manner; in the second, there were no spermatozoa; and
in the third, the vesiculæ were shrunk, empty, and quite rudimentary
in size. So that these three individuals certainly could not have
impregnated their own eggs; nevertheless, within the shell of these
very three, there were perfectly developed larvæ: I am led to conclude
from this fact, that adjoining specimens in a perfect condition had,
by means of their long probosciformed penis, effected the fecundation
of their imperfect neighbours. I need only further add, that some
out of the above six specimens, with more or less aborted cirri and
imperforate male organs, were infested by a peculiar parasite, allied
to Bopyrus,[50] and that these specimens did not contain ova.
[50] I have given a short notice on this parasite, in my former
volume on the Lepadidæ, in a foot-note to p. 55.
_Metamorphoses and Homologies, throughout the Order of Thoracica._
In my former volume, the metamorphoses were described under three
principal stages or heads; but whether these three included all
the main changes, I was then hardly able to conjecture. But now
I have reason to believe that such is the case, for in the genus
Cryptophialus, belonging to the Abdominalia, the whole course of the
metamorphosis, from the egg to the pupa, takes place within the sack of
the parent; and I found, when having, on the coast of South America,
numerous specimens to examine, that the egg-like larvæ (Pl. 24, fig.
15-18) could be naturally grouped into two main stages, but with many
transitional intermediate grades (answering to the successive moults in
the first stage of ordinary larvæ), before they passed into the third
or pupal stage. And the first two stages in these egg-like larvæ of
Cryptophialus, clearly seem to correspond with the first two stages in
ordinary larvæ; for in both the chief changes are, the shortening of
the terminal projection--the increase in size and approximation on the
ventral surface of the anterior horns or cases for the antennæ--and the
compression of the whole body. In all members of the Thoracica, the
metamorphosis seems to run a remarkably uniform course. The larva in
the first stage undergoes several moults and slighter changes--how many
is not known--before arriving at its second main stage, which has been
observed only in one single instance; and judging from Cryptophialus,
this second stage passes abruptly by one moult into the pupal stage;
and this, certainly, passes abruptly into the Cirripedial or mature
stage.
_Larva, First Stage._
The larvæ in this stage are known, amongst the Balanidæ, in Balanus,
Pyrgoma, Coronula, Platylepas, and Chthamalus; and these genera include
all the principal forms. Amongst the Verrucidæ they are known in its
one genus, Verruca. Amongst the Lepadidæ, in Scalpellum, Ibla, Alcippe,
Lepas, Conchoderma, &c.; and in all these genera the larvæ present no
important difference--hardly any difference which could be viewed as
generic, were these larvæ independent animals,--as may be inferred,
chiefly, from Mr. C. S. Bate's descriptions.[51] The abstract given in
my former volume was not accompanied by any illustrations, and I have
consequently here given (Pl. 29, fig. 8), a view of the larva, in the
first stage before moulting, of _Scalpellum vulgare_: the natatory legs
are not drawn with accuracy, only the relative position of the several
organs having been carefully attended to. I have also had copied from
Mr. Bate's memoir, a figure of the larva (Pl. 29, fig. 9) of _Balanus
balanoides_, in its first stage, _before_ moulting, with its ventral
surface exhibited; and another figure (with a few trifling alterations
made after examining specimens most kindly sent me by Mr. Bate) of
the larva of _Chthamalus stellatus_ (fig. 10), in its first stage, but
_after moulting once_. It should be observed that Mr. Bate has given
a drawing of the larva of this latter cirripede, in the first stage,
_before moulting_; and it does not differ essentially from that just
referred to (fig. 9), of _B. balanoides_, but is rather more fully
developed. These drawings suffice to show the character of the larvæ
in the first stage, both before and after the first moult, and even
after the second moult, throughout the Order of Thoracica. The larvæ
sometimes undergo their first moult within the sack of their parent, as
I have been informed by Mr. Bate, and as I have observed in Coronula.
[51] 'Annals and Magazine of Natural History,' vol. viii (2d
series), 1851, Plates 6, 7, and 8.
I will now make a few remarks on these larvæ in the first stage, before
and after the first moult, supplemental to those in my former volume.
Their shape is oval, and the whole dorsal surface is evidently covered
by a carapace. It is remarkable that the body exhibits no distinct
articulations; those given by Goodsir[52] being certainly erroneous.
Commencing at the anterior extremity, the eye varies considerably in
the state of its development; in _Platylepas decorata_ it is nearly
circular, and in most of the specimens very distinct; whereas in
the allied _Coronula balænaris_, before the first moult, it is very
imperfect, but afterwards square and of considerable size. In _Balanus
galeatus_, in the immature larvæ dissected out of the egg, the cellular
matter which was in process of conversion into the eye, formed a
transverse band, obscurely divided into two portions, and this seems
to indicate that the single eye is in fact formed by the confluence of
two eyes. In _Scalpellum vulgare_, this heart-shaped eye lies between
a V-shaped muscle, the nature of which I cannot understand, and which
has not been represented in (Pl. 29, fig. 8, _a_). I need only further
add, that in _Chthamalus stellatus_, after the first moult, the eye
exhibits, in specimens sent me by Mr. Bate, some appearance of tending
to become double.
[52] 'Edinburgh New Philosophical Journal,' July, 1843, Pl. 3, 4.
Arising posteriorly to the eye, we see, in _Scalpellum vulgare_, a
pair of minute curved horns (_b′_), directed backwards; and within
these horns I distinctly saw an articulated organ. These horns are
difficult to be distinguished, and probably could not be made out
previously to the first moult, in any larva of less size than that of
_Scalpellum vulgare_. But after the first moult, Mr. Bate has seen, in
two species of Balanus, in Verruca and in Chthamalus (fig. 10, _b_),
a pair of articulated organs, in this same position, evidently now
forming antennæ, and directed anteriorly, and free from any envelope.
It is somewhat important, as we shall presently see, to bear in mind
that these antennæ first appear within an envelope or horn; and that I
detected that they included an articulated organ, before I had heard
of Mr. Bate's observations. These antennæ, from their small size, from
being seated internally with respect to the horns containing the other
pair of antennæ, and from the position which the latter assume in the
later stages of the larva, I believe to be the first or anterior pair.
Their position in appearance posteriorly to the large lateral horns,
containing the second pair of antennæ, is probably due to the anterior
cephalic segments having been driven inwards, the truncated outline
of the front of the head, and likewise, probably, the position of the
mouth between the bases of the natatory legs being thus caused.
In this same larva of _Scalpellum vulgare_, within the great lateral
horns just alluded to (fig. 8, _c_), filiform organs, supporting rows
of spines, could be distinguished; and these appeared to me to be
antennæ. These horns or cases resemble in structure the smaller pair
just described; they arise from the ventral surface, and can hardly,
therefore, be considered as prolongations of the carapace. After
the first moult (fig. 10, _c_) they are seen to have increased much
in length: in some cases they are of considerable length before the
first moult, as in Lepas: in the Balanidæ they seem to be generally
shorter than in the Lepadidæ; but in _Balanus galeatus_ I found them
one third of the entire length of the animal. Whilst within the
egg, these horns are adpressed laterally to the body, and so point
posteriorly; afterwards they project rectangularly from the sides,
or, as in _Scalpellum vulgare_, are directed somewhat anteriorly. As
in the larvæ of all ordinary Crustaceans, as yet known, the antennæ
are amongst the earliest developed organs; and as the first pair of
natatory legs (Pl. 29, figs. 8-10, _e_) in these Cirripedial larvæ,
might so very naturally be thought to be antennæ (as has been remarked
to me by Mr. Dana), both from their structure and from their position a
little anteriorly to the mouth, I am well aware that to prove my view
correct, namely, that these horns are the second pair of antennæ in
process of formation, it is not sufficient merely to have seen organs
resembling antennæ within them; nor is it sufficient to advance the
strictly analogical fact of the first-mentioned pair of antennæ, which
in Scalpellum indisputably appear in their earliest condition within an
envelope or horn. Further evidence is required, and this is presented
in Cryptophialus, in which the lateral horns of the egg-like larva, in
its first stage (Pl. 24, fig. 16), can be actually followed step by
step until, in the second stage (fig. 17), just before passing into
the pupa, the horns are seen to have become larger and more nearly
approximated to each other on the ventral surface; and whilst in this
condition, I several times dissected out the prehensile antennæ of the
future pupa with _every character perfectly recognisable_. Hence I
cannot doubt that in the larvæ of Cirripedes the law of development is,
that in their very earliest condition, the small first pair of antennæ
are enclosed in cases; and that the large second pair remains thus
enveloped until the pupal stage. This conclusion, we shall immediately
see, is in harmony with the late development of the succeeding
appendages or organs of the mouth, which certainly do not appear in the
first larval stage, and are not known to appear even till after the
final metamorphosis.[53]
[53] According to M. Joly, ('Annales des Sciences Naturelles,' 2d
series, tom. xix, p. 59) in the larva of the macrourous Caridina,
the natatory legs appear before the gnathites or parts of the
mouth; so that in ordinary Crustaceans there is no invariable order
of development from the anterior towards the posterior end of the
body, as has sometimes been supposed.
The mouth is more or less probosciformed (Pl. 29, figs. 8-10, _d_),
differing considerably in this respect in different species of the
Lepadidæ; and this, probably, is due to the larva being born in a more
or less mature condition. Its exact position likewise varies, for it
arises either between the first or second pairs of natatory legs. It is
known, from Mr. Bate's observations, to have the power of movement. It
is directed posteriorly, the œsophagus extending anteriorly; both these
directions being the same as in the mature cirripede. Certainly during
these early stages there are no jaws or gnathites; but the margin,
answering to the labrum, is furnished with some short, thick, sharp
spines, and with hairs. In _Scalpellum vulgare_ the orifice of the
œsophagus seems to lie rather beneath the upper prominent spinose edge,
which, as just remarked, probably answers to the labrum; but this is
one of the species in which the probosciformed mouth, at least before
the first moult, is not much developed.
We come, now, to the three pairs of natatory legs: the first (Pl. 29,
figs. 8-10, _e_) has throughout the order only one ramus, whereas the
two succeeding pairs (_f_, _g_) are biramous. I must here remark that
the straight and strong, and the curved plumose spines, with which
these limbs, after the first moult, become furnished, now appear to me
as more probably prehensile, rather than masticatory as I imagined in
my former volume. That these spines are important organs to the larvæ
I do not doubt. With regard to the homologies of these three pairs of
limbs, my first impression was that they were the mandibles and the
two pairs of maxillæ in their earliest condition; but I consider this
view as quite untenable, for several reasons; viz., the wide interval
between their bases and the mouth itself,--the somewhat variable
position of the mouth with respect to the legs,--and the position
which the latter occupy in the _second_ larval stage.[54] A far more
tenable view is that these three pairs of legs are the three pairs
of maxillipeds, in their earliest condition, in accordance with the
view of M. Joly[55] on the nature of the three very similar pairs
of natatory legs in the larva of Caridina, a macrourous Crustacean.
But, in Cirripedes, the three pairs of natatory legs, in the larva
in the first stage, are apparently the very same as the first three
pairs, in the larva in the second stage, and in the pupa. And in the
pupa the first three pairs, which certainly correspond with the first
three pairs of cirri in the mature animal, seem to me, for reasons
presently to be assigned, to be the second, third, and fourth thoracic
limbs. Hence I am led to the conclusion that the first pair of legs in
the larva in the first stage, are homologically the second thoracic
(answering to the third pair of maxillipeds in the higher Crustaceans),
and that the two succeeding pairs are the third and fourth thoracic
limbs; to be succeeded, in the _pupal_ stage, by the fifth, sixth, and
seventh thoracic appendages.
[54] Mr. Dana, moreover, has remarked, ('Crustacea: United States
Exploring Expedition,' p. 1386), "that he knows of no instance of
a mandible becoming so completely a leg, as to lose wholly the
mandibular function even of its basal portion."
[55] 'Annales des Sciences Naturelles,' 2d series, tom. xix, 1843,
p. 34. M. Joly's observations were made on the Caridina. I owe to
the great kindness of Mr. C. Spence Bate, an examination of some
larvæ of the allied genus _Hippolyte varians_, and I found, on
dissection, the view of M. Joly, that the three pairs of natatory
legs are the maxillipeds, so far strongly confirmed, that they
followed closely, with equal intervals, the mandibles and two pairs
of maxillæ. The first pair of natatory legs in Caradina, in its
earliest condition within the egg, is uniramous, like the first
pair in the larvæ of Cirripedes. There is one fact which seems
rather strongly opposed to the view of these three pairs of legs
in the larvæ of the macrourous Crustaceans being the maxillipeds,
which is that Capt. Du Cane ('Annals of Nat. Hist.,' 1838, vol.
ii, pl. 6, and 7) observed only three pairs of limbs in process
of formation posteriorly to the first three pairs, whereas there
should be found, in accordance with M. Joly's view, five pairs, _i.
e._ all five pairs of ambulatory legs. This one fact countenances
the view, which I hold on the nature of the legs in the larvæ of
Cirripedes during their early stages, namely, that they are the
second, third, and fourth thoracic limbs, to be succeeded by only
three additional pairs.
Lastly, behind the natatory legs, on the ventral surface, (Pl. 29,
figs. 8, 9, _i_), the body is much produced, and terminates in a
horny fork, which, after the first moult (fig. 10, _i_), becomes much
elongated. Anteriorly to this fork, on the ventral surface, there
is another fork (_l_), and again above this I could distinguish, in
_Chthamalus stellatus_, after the first moult, another fork (_m_),
or at least a pair of short thick spines. From the structure of the
forked abdomen in the known larvæ of the Podophthalmia, I presume
that this portion of the body is the abdomen of the young Cirripede,
but it is not at all plainly articulated. After the first moult, the
posterior end of the carapace (_h_), which is always pointed, becomes
much elongated and serrated on both sides;[56] reminding one of the
structure of the carapace of the so-called Zoea, or larva of certain
Podophthalmia. Situated under this posterior prolongation of the
carapace, there is a swelling (_n_, with long hairs on both sides),
which apparently lies on the dorsal surface of the spinose and forked
abdomen; here, when the larva is compressed, the cellular and oily
contents of the body burst forth; and I suspect that this swelling
is the anus, for it is known from the researches of Rathke,[57] that
the anus in the higher Crustaceans opens during the earliest periods
dorsally.
[56] I suspect that the account given by Goodsir ('Edinburgh New
Phil. Journal,' 1848) of the posterior points of the carapace and
abdomen in the larva of a Balanus, is not quite accurate.
[57] 'Annales des Scienc. Nat.,' tom. xx, p. 451.
_Larva, Second Stage._
I have given, from Burmeister,[58] a lateral view (Pl. 30, fig. 1)
of the one single specimen, ever observed of a larva in this stage,
belonging, as is supposed, to the genus Lepas. The carapace has now
greatly altered its character. The two fleshy projections, as so
called by Burmeister, by which the larva adhered to the sea-weed,
were supposed by this author to include the great prehensile antennæ
of the pupa; from my observations, already alluded to, on the two
projections (Pl. 24, fig. 17) in the closely analogous egg-like larva,
in the second stage, of Cryptophialus, by which it also adheres, I
have not the least doubt that this is the case. The small, internal,
and anterior pairs of antennæ, are, as it would appear, now aborted.
The eye, according to Burmeister, has commenced becoming double;
but the two approximate eyes are not as yet compound. The mouth is
probosciformed (_m_), and does not differ much from its condition in
the first stage; no gnathites were observed by Burmeister, and they
could not be expected to be present, for they are not found even in
the pupa. The mouth, which in the larva in the first stage differs in
different genera, in being more or less advanced forward, here stands
some way anteriorly to the natatory legs, as in the pupal condition.
The first pair of legs is uniramous, and the two other pairs biramous;
this fact, together with the number of the legs in this second stage
being still three, and their structure being not very different,
leaves little doubt on my mind that we here have the same three pairs
as during the first stage. The abdomen has become much shortened, but
still space is left for the development, in the pupa, of the three
posterior pairs of legs. I may here remark that in the pupa the
anterior natatory legs have become, like the others, biramous; but yet,
as it were for the purpose of showing their metamorphosis from the
uniramous legs of the earlier stages, they have their bristles arranged
rather differently from those on the succeeding five pairs of legs.
[58] 'Beiträge zur Naturgeschichte der Rankenfüsser,' tab. 1, figs.
3, 4.
_Larva in the Last or Pupal Stage._
I have given a lateral view of the pupa of _Lepas australis_ (Pl. 30,
fig. 2), illustrative of the description in my former volume: the
specimen is drawn as if transparent, and it was to a certain extent
thus rendered by boiling in caustic potash. A sketch of the position
of the young Cirripede within the pupa, was made by the camera. At
first the drawing will perhaps hardly be comprehended: the darker
shaded portion to the left of the letter (_b_) shows the extent of the
sack, with the included thorax and natatory legs of the pupa: to the
right of the same letter, if we do not consider the young included
Cirripede, the only organs distinguishable in the mass of cellular and
oily matter, are the alimentary canal, the cement-glands (_t_), _i.
e._ the incipient ovaria, and the cement-ducts (_t′_) which enter the
antennæ. A view is also given (fig. 4) of the ventral surface of the
pupa; and a transverse section (fig. 7) of the carapace, taken close
to the eye-apodemes. On comparison with the larva in the second stage,
the changes in external appearance and structure are not very great;
the prehensile antennæ are freed from their cases; the two eyes stand
further apart; the three posterior pairs of legs have been developed,
and a small abdomen has become distinctly separated from the thorax.
Before proceeding to make a few additional remarks and corrections to
my former description of the pupa, it will be advisable, on account of
the importance of the subject, to discuss the homologies of the limbs.
From the presence of eyes and of two pairs of antennæ in the larva,
during its earlier stages, the front of the head consists, in
accordance with all analogy, of three segments; the mouth, likewise,
from being formed of three gnathites (which can be detected by
dissection in the pupal state), consists, also in accordance with all
analogy, of three segments, making altogether six segments--on the
nature of which I apprehend no objection will be raised. In two out
of the three orders into which Cirripedes may be divided, the mouth
is succeeded, in the adult animal, by eleven most distinct segments;
of which the first (_i. e._ the seventh cephalic) differs from the
succeeding seven thoracic segments; and these seven again differ
from the three abdominal and terminal segments. Hence it must be
admitted that, as far as the cephalo-thorax of the archetype Cirripede
is concerned, it consists, like that of the archetype Crustacean,
of fourteen segments, of which eight succeed the first-named six
that form the mouth and front of the head; and that, with the three
abdominal segments, there are altogether seventeen segments. In the
order Thoracica, however, which includes all common Cirripedes, both
in the pupa and in the mature animal, only six thoracic segments
with their appendages, succeed the mouth, two having been lost; and
the question arises which are these two, whether the seventh and
eighth, or the thirteenth and fourteenth (_i. e._ the two terminal
thoracic) segments; for there is no reason to suspect any other
segments of having disappeared. In my former volume, I inferred,
without sufficiently entering into my reasons, that it was the seventh
and eighth, _i. e._ the last cephalic and first thoracic segments,
which had disappeared; but I now find that Mr. Dana[59] believes
that, in ordinary Crustaceans, the abortion of the segments with
their appendages almost always takes place at the posterior end of
the cephalo-thorax. Nevertheless, after due deliberation and fresh
examination of the pupa, I must retain my former opinion, that it
is the last cephalic and first thoracic segments which have either
coalesced with the others, or wholly disappeared. In the pupa, the
mouth, although functionless, has its place most plainly marked by
being slightly prominent, and by the presence of a sort of labrum and
of a shrivelled œsophagus, round which latter the gnathites and the new
œsophagus of the future young cirripede are in process of formation.
Now between the mouth of the pupa and the first pair of natatory legs,
there is a space of membrane, equalling, when stretched out, the three
succeeding thoracic segments in length and breadth: this interspace, I
conceive, must have some homological signification; here then we have
at least an appearance of the abortion of appendages; whereas, at the
posterior end of the cephalo-thorax, no such appearance is presented.
Moreover this interspace of membrane is divided nearly in the middle by
a most conspicuous fold, which, on the view here adopted, would mark
the separation of the seventh (cephalic) from the eighth (thoracic)
segment; and the interspace and fold are thus simply explained. Lastly,
I have shown, in the Introduction (p. 18), that the first and five
succeeding pairs of cirri of the mature Cirripede present certain
small, but significant, resemblances in structure and in the origin
of their nerves, with the outer pair of maxillipeds and with the five
pairs of ambulatory legs in the Podophthalmia; which resemblances are
all futile, if the cirri belong to the 7th, 8th, 9th, 10th, 11th, and
12th segments of the cephalo-thorax, or those immediately succeeding
the mouth; but are full of meaning, if the six pairs of cirri belong,
as I believe, to the 9th, 10th, 11th, 12th, 13th, and 14th segments, or
the six posterior segments of the cephalo-thorax.
[59] 'Crustacea: United States Exploring Expedition,' p. 22.
Before commencing on details, I may premise that I have examined
the pupa of _Lepas australis_, _pectinata_, _fascicularis_, and
_anatifera_, of _Conchoderma virgata_, partially of _Dichelaspis
Warwickii_, of _Ibla quadrivalvis_, and of _Alcippe lampas_; and in
the Balanidæ, of _Balanus balanoides_ and _Hameri_. In the pupæ of all
these genera there is a most close general agreement in structure,
excepting in minute details: I was surprised to find exactly the same
slight differences in the spines on the first pair of natatory legs,
as compared with the succeeding pairs, in _Balanus Hameri_, as in
Lepas. The abdomen and caudal appendages of the pupa in the abnormal
Alcippe, as we shall presently see, offer the only marked exception
to this uniformity of character throughout the Thoracica. The outline
of the carapace or shell is usually not so blunt at the anterior end,
as in the pupa of _Lepas australis_ (Pl. 30, fig. 2); more commonly
the shape is that of the pupa of Alcippe (Pl. 23, fig. 16). In _Lepas
pectinata_ the two posterior points of the carapace are produced into
two short spines. The surface of the carapace in _L. australis_ is
lined, as represented in fig. 4: the colour of this species when alive
was blue:[60] in _L. fascicularis_ the surface is punctured: in _L.
pectinata_ it is marked with curious points of various shapes, often
star-shaped, in parts reticulated, and confluent along the dorsal
margin, and in parts lined: in _B. balanoides_ it is very obscurely
punctured, and in _B. Hameri_ the punctures pass into lines. The whole
of what is externally visible consists of the carapace, for this is
produced not only backwards, so as to enclose the thorax and abdomen
with their appendages, but also forwards, so as to overhang the whole
front of the animal; and the prehensile antennæ, in Lepas, Ibla,
Balanus, and probably in all the genera, can be retracted within its
lower edge. The protection afforded by the carapace to the antennæ
is aided by two crests (Pl. 30, fig. 7, _c_) parallel to this lower
edge. The whole sternal surface is very narrow (fig. 4), and is
likewise protected by the carapace; that is, when the two sides are
drawn together by the adductor muscle. The shell, however, when thus
drawn together, gapes a little at the two ends, at least in the case
of _Lepas australis_. The adductor muscle, if introduced in fig. 4,
would have crossed close anteriorly to the basal margin of the mouth;
and in fig. 2, its end on the near side would have been attached under
the dark cæca, which enter the upper end of the stomach. The adductor
is shaped almost like an hour-glass, and so differs from this muscle
in the mature Lepas, in which it is of the same thickness throughout.
I may here add that the pupa of _Lepas australis_ could swim very
rapidly, and often on one side in a circle; it could walk by the aid
of its antennæ, but often fell over; being thus locomotive, and, as
we shall immediately see, well provided with senses, it cannot be
considered as very lowly organised.
[60] I took this species alive in the Southern Atlantic Ocean; and,
mistaking it for an independent Crustacean, was much perplexed
where to class it. I had overlooked these specimens when publishing
my former volume.
_Acoustic Organs._--Commencing at the anterior end, two small elongated
orifices, 10/6000th of an inch in diameter, (_e_, fig. 4, Pl. 30),
may be seen; these lead, as described in my former volume, into a
sack, with a bag suspended in it, which is provided with a large
nerve, and which I believe to be the acoustic vesicle. These orifices
occur in the carapace, either in the same position, or a little more
posteriorly, in the pupæ of all Cirripedes. In _Balanus balanoides_
they are minute, being only 2/6000th in diameter, but are surrounded
with a border: in _Conchoderma virgata_ they are also surrounded
by a border: in _Lepas pectinata_, the orifices are 3/6000th of an
inch in diameter, and are very singular from being seated on rounded
prominences, causing the carapace to have two short, blunt horns in
front. In _Lepas australis_, and I believe in the other species, the
corium round the acoustic orifices is darkly coloured; and these
coloured marks can be distinguished for some little time on the
peduncle of the young Cirripede, after the metamorphosis, and after
the entire organ, together with the whole pupal carapace and eyes, has
been moulted. Knowing the connection in the higher Crustacea, of the
acoustic organs and the antennæ, and seeing the very backward position
(figs. 2 and 4) of the one great pair of antennæ, I have always
imagined that these orifices probably marked the normal position of the
anterior pair of antennæ, which, since the earlier larval stages, have
disappeared. And I now find[61] that Schödler affirms, that in most,
if not in all Daphnidæ, there is a black spot in front of the eye,
which is connected with an opening in the basal portion of the anterior
antennæ, and he concludes that it is an organ of hearing.
[61] Quoted by Dana, 'Crustacea of United States Exploring
Expedition,' p. 1264.
_Antennæ._--These, from their present position, and from standing, in
their earlier stages whilst within their envelopes or horns, exteriorly
to the small medial pair (since aborted), I believe to be the second
pair; and this is Mr. Dana's opinion. In my former description of these
very singular and important organs (Pl. 30, figs. 4 and 8), I have
fallen into some considerable mistakes: the two plates or segments
(fig. 4, N), of which the posterior margins are inflected as apodemes
(_n_), carrying the eyes, are certainly, as may be clearly seen in the
pupa of Alcippe, Pl. 23, fig. 16, and as affirmed by Burmeister,[62]
the basal segments of the antennæ. The second or main segment
(formerly called by me the basal segment) has in some species an upper
portion of the membrane of which it is composed, next to the body,
excessively thin, and separated from the rest of the membrane composing
the segment, by an oblique line (fig. 8, _o_), which I mistook for
its articulation with the body.[63] We then come to the disc or third
segment; and lastly to the fourth, or ultimate segment. This ultimate
segment, generally, has its external corner projecting up, as a
step; and this sometimes, as in _Dichelaspis Warwickii_, gives the
appearance of its consisting of two segments; but a careful examination
of this part in Ibla, in which the step-like structure is carried to
an extreme, makes me believe that there is only one segment.[64] The
prehensile antennæ, therefore, like the natatory legs, are formed of
four consecutive segments, of which the basal segments give rise to the
singular apodemes, presently to be noticed (fig. 7), that carry the
great compound eyes. This basal segment, in all Cirripedes, is moulted
with the eyes, the three other segments invariably remaining cemented
to the surface of attachment.
[62] 'Beiträge zur Naturgeschichte der Rankenfüsser,' p. 19.
In tab. 1 of this work there are good drawings of the general
structure of the pupa of a species of Lepas, probably _L.
australis_. I believe this author was the first who made out the
structure of the abdomen of the pupa.
[63] In the table of measurements of the antennæ of the several
genera and species of the Lepadidæ (p. 286) of my former volume,
the articulation, called by me _basal_, I now know to be really
the articulation between the basal and second segment. In the
fourth column, headed "Length from end of the disc to the inner
margin of the basal articulation," the term inner margin really
applies to the oblique curved line separating the thin and scarcely
visible membrane from the thicker membrane of that segment. These
corrections do not in the least affect the object for which the
table was given.
[64] In a sketch, sent me by Mr. Dana, of this organ in the pupa
of a Lepas from the Antarctic Ocean, I observe that he divides my
ultimate segment into two segments.
In the Southern Atlantic I took some specimens of the pupa of
_Lepas australis_, not yet attached, and therefore with the muscles
of the antennæ, not having suffered any of that absorption, which
they undergo, as soon as the pupa is permanently cemented to some
floating object. In my former volume I noticed a pair of strong
muscles, attached to the tips of the middle forks (Pl. 30, fig. 7)
of the apodemes, and I now find two pairs attached to the bases of
the two outer forks, and directed dorso-anteriorly; and two other
pairs, also attached to their bases, but directed dorso-posteriorly,
so that altogether there are five pairs of muscles attached to the
apodemes; their chief function, I should think, was to draw the antennæ
posteriorly and upwards within the carapace; but as the apodemes cannot
be moved without the great compound eyes being likewise moved, the
muscles probably serve a double purpose. When the pupæ were alive, I
noticed that their eyes were constantly kept in a state of vibratory
movement. Flexor and extensor muscles are attached at one end to the
posterior margin of the basal segment, and at the other end to the
second or main segment; other powerful muscles attached to this latter
segment, are prolonged by ligaments into the disc. In Cryptophialus I
observed that the disc-segment had a movement almost like that of the
wrist. Whether any muscles enter the small terminal segment, I know not.
The drawing in Pl. 30, fig. 8, of part of the second segment, of the
third or disc segment, and of the fourth or ultimate segment, in
_Lepas australis_, is, I think, very accurate. The second segment
articulates on the upper or dorsal surface of the disc, and has the
articulation on one side constricted and formed of thin and flexible
membrane; the little terminal segment, which is turned outwards at
right angles, also, articulates on the disc. That the disc forms a
true segment is shown clearly in Cryptophialus (Pl. 24, fig. 18),
where the articulation with the second segment is not in such close
contact. The disc is either circular, as in Lepas, or hoof-shaped, as
in Ibla: in _B. balanoides_ the disc is rather hollowed out on the
inner side. It has the power of adhering even to so smooth a substance
as glass, placed vertically. It is surrounded by a rim of transparent
membrane. On the hinder margin some spines arise from the central and
more opaque part: in _Lepas australis_, there are no less than seven of
these spines (fig. 8): in _Conchoderma virgata_ there are only four,
in _Scalpellum_ and _Ibla_ only one. When the disc is placed on the
surface of attachment, these spines lie parallel to it. The middle
part of the disc is, almost always, nearly opaque; and in tracing
the cement-ducts from within the body of the pupa, or of the young
Cirripede, I in many cases traced them as far as this point, but here
lost them. From this same obscure central part of the disc, in most,
if not in all species of the Lepadidæ, spokes radiate, which sometimes
are branched, and are not regular, not always even resembling each
other on the opposite sides of the same individual. Round the proper
membranous border of the disc, a second one may be observed (fig. 8,
_p_), which differs in shape and extent in different specimens: under
favorable circumstances, and very high powers, it may be seen to
have a reticulated structure, and to be of a very pale brown colour;
towards the exterior margin, the reticulations become finer, and are
blended together and lost; on the inner margin, the substance forming
this membrane may be seen to come out of the spokes. This substance
is the cement, which has the power of adhering to whatever substance
it grows against; and thus the disc of each antenna becomes cemented
down, and soon both the antennæ are surrounded by a common border of
cement, which gradually increases, after the metamorphosis, in extent.
Occasionally the cement forms little projections, like short spines, on
the edges opposite to the orifices of the spokes.
The small terminal segment usually bears on its truncated extremity
six spines, some of which are occasionally hooked; in Scalpellum, two
spines, rather longer than the others, are borne on a step some way
down on the inner side of this segment; but in Lepas, two spines (fig.
8), very much longer than the others, arise from the outer corner
of the extremity. These two are very different from the other four
borne by this segment, or indeed any other spines on the body; for
they are quite flexible, and are furnished with a double row of very
long, straight, excessively fine hairs, which seem to be articulated
on them--the whole presenting a very beautiful appearance. These
spines are of considerable length, and in _Conchoderma virgata_ they
even equal in length the whole antenna. I can hardly doubt that these
beautiful, plumose, flexible spines, into the thick bases of which the
coloured corium could sometimes be seen to enter, serve as feelers.
Owing to the facts immediately to be mentioned, I erroneously stated,
in my former volume, that there were three long spines.
In three species of Lepas, in _Dichelaspis Warwickii_, and in
_Scalpellum Peronii_, after having torn the lately-cemented antennæ
from the surface of attachment, I observed proceeding from the end of
the terminal segment, from between the above two groups of spines, what
appeared to be a long narrow ribbon with its end torn off; and which,
in the case of _Lepas australis_, I fancied was one of the plumose,
long, flexible spines ripped open. But now that I have examined some
of the pupæ of this species before their attachment, I find (as
represented in Pl. 30, fig. 8, _v′_) a flattened tube, ending in a
blunt point, and having a peculiar ringed structure. I have noticed
similar appendages to the antennæ of specimens just attached of _Lepas
anatifera_. In the Dichelaspis and the Scalpellum, the tube was very
long, and seemed, from its torn appearance, to have been firmly
attached to the supporting surface. In both these cases, the tube came
out from within another slightly larger tube, which had been broken
off close to the extremity of the terminal segment of the antenna. In
the case of the _Lepas anatifera_, the tube expanded a little after
leaving the antenna. In the Dichelaspis, it had exactly the same
diameter as the cement-duct, which could be clearly distinguished
within the two lower segments. From these several facts, and from the
peculiar appearance of the tube itself, I believe it to be a tube of
cement-tissue which thus, sometimes even before the pupa is attached,
independently grows outwards. That the cement-tissue can grow outwards
and assume definite forms, we know from the singular case of _Lepas
fascicularis_, in which the cement proceeding from several apertures,
forms a vesicular float round the peduncle of not only a single
individual, but often of a group of specimens: we shall presently find
a somewhat analogous fact in the case of Coronula. It is possible
that this tube, proceeding from the extremity of the antenna, may be
the channel through which cement continues to be poured forth during
the continued growth of the above Cirripedes; and the manner in which
this is effected, considering how firmly the end of the peduncle is
cemented down, has always appeared to me a difficulty. In those pupæ
of _Lepas australis_, which I caught swimming about unattached, it is
surprising that the disc should have been edged with cement, and that a
tube, similarly formed, should have grown out of the ultimate segment:
it shows, I presume, that the cement-tissue will grow out, whether or
no the pupa has succeeded in finding a proper object for attachment.
Lastly, I have felt some surprise, in two or three instances in
observing some dark purple pigment-cells, like those in the corium,
within the terminal tube of cement; and likewise within the spokes of
cement in the disc: this is the only fact which causes me the least
doubt, whether I have rightly determined the nature of the terminal
tube, as wholly formed of cement tissue; or whether it may not be
covered by an outer integument, itself lined by true corium, coloured
purple.
Finally, I may add, that, excepting in small details, the prehensile
antennæ present no difference throughout the Order: I have minutely
examined them in several genera of the Lepadidæ; and in the Balanidæ,
I have seen them in Coronula, and in several species of Balanus. In
_B. balanoides_ I have examined them carefully; they are smaller and
thicker than in Lepas, with the second or main segment bowed outwards,
carrying its usual single spine; with the disc excised on its inner
margin and apparently without the spoke-like vessels for the cement;
and with the ultimate segment proportionably longer, and carrying,
I believe, six spines, of which two appeared to be longer and more
flexible than the other four shorter and somewhat hooked spines. In
_Coronula balænaris_, also, the terminal segment is, proportionably
to the others, of large size. Not only throughout the order, but
throughout the whole Class, the antennæ are singularly uniform in
structure, as will be seen, when the last two orders are described.
_Eyes._--These present no difference, except in size, throughout the
class; and have been sufficiently described in my former volume. The
true basal segments of the antennæ (incorrectly designated formerly as
sternal plates or segments) are separated from each other by a deep
fold; and are separated from the edges of the carapace on each side
by a crest and slight fold (Pl. 30, fig. 7, _c_; and 4); these folds
and crests die out posteriorly, and disappear. The hinder, rounded
margins of the basal segments are inflected inwards, and their corners
are produced far up into the body, thus forming the curious UU-like
apodemes. These apodemes exist throughout the whole class; and the
outer arms always carry the great compound eyes. I noticed, in _Lepas
pectinata_, that the two middle arms are proportionably longer than in
_L. australis_. Owing to the presence of these apodemes, and to certain
coloured marks on the adjoining corium, the eyes, though enclosed
fairly within the carapace, yet deceptively appear pedunculated,
so that even J. Vaughan Thompson was thus deceived. I have already
described the several muscles attached to these apodemes, and the
constant vibratory movement of the eyes. Whilst the pupa remains a
freely swimming animal, the eyes are included, not only within the
shell or carapace, but (as would naturally happen) within the corium or
true skin lining the carapace; but after the pupa has become attached,
preparatory to its final metamorphosis (in the state represented at Pl.
30, fig. 2.), not only are the muscles, as before remarked, which are
attached to the apodemes, absorbed, but so is the corium investing the
apodemes and the immediately adjoining parts of the carapace. Hence it
comes that the new corium of the young Cirripede within, is formed in
a deep transverse fold across the whole lower half of the animal, and
the apodemes with the eyes are thus, as it were, rejected from within
the corium, though still remaining within the carapace. Consequently in
this final stage, the eyes and apodemes are very conspicuous from the
outside, being seen only through the transparent carapace. I presume
that the eyes at this period have become functionless, with the optic
nerve divided and absorbed. The eyes, apodemes, and carapace soon
afterwards are all moulted together.
The eyes of Cirripedes certainly undergo a remarkable series of
changes: in the larvæ in the first stage, there is a single eye,
perhaps formed by the confluence of two eyes, occupying the normal
position in the front of the head: in the second stage, according to
Burmeister, the eye has become double, but the two are as yet simple;
they are now situated posteriorly to the second pair of antennæ: in
the third or pupal stage, they remain in the same situation, but have
become compound, of great size, and are attached to the apodemes of
the antennæ: in the mature and fourth stage, they have moved someway
posteriorly, and again have become simple, of minute size, and are
either confluent, as in the Lepadidæ, or tolerably far apart, as in the
Balanidæ. It must not be supposed that the eye of the mature Cirripede
is metamorphosed from the eye of the pupa, for such is not the case;
the new eyes and old eyes being formed someway apart, and frequently
both can be seen within the pupa (as in Alcippe, Pl. 23, fig. 16) at
the same time. It is scarcely possible that the eye of the larva in the
first stage, can be changed into the double eyes of the second stage;
though these latter may possibly be multiplied into the eyes of the
pupa, as both continue to occupy nearly the same position.[65]
[65] Zenker, in his 'Physiological Remarks on the Daphnidæ,'
('Journal of the Microscopical Society,' 1853, p. 274), speaks of
a "tripartite azygous eye" as common amongst Crustacea, and as
occurring "in conjunction with the aggregated eyes in Artemia,
Argulus, &c.; but as appearing regularly in all the Branchiopoda
and Siphonostomata as the _earliest_ visual organ." Hence I
conclude that this azygous eye is the homologue of that single eye
which appears in the earliest larval stage of Cirripedes; and that
the compound eyes of the cirripedial pupa, answer to the aggregated
eyes of Artemia and Argulus, &c., with the difference, that in
these latter genera the single eye is retained. See, also, Von
Siebold, 'Anatomie Comparée,' tom. i, p. 435.
_Mouth_, _thorax_, _limbs_, _abdomen_.--I have nothing to add regarding
the mouth, except to confirm my former account; viz., that it is
functionless, consisting merely of crests, which project inwardly
between the gnathites of the young Cirripede, and of a shrivelled
closed tube representing the œsophagus. In fact the mouth is a model
of the outside of the mouth of the young Cirripede. I may remark that
some little way beneath the membrane answering to the labrum, a pair
of ligamentous apodemes, the use of which I do not know, slightly
penetrate the body. The degree of prominence of the mouth varies, but
it is far less than in the mature animal. On the limbs I have nothing
particular to add: the drawing of the first pair of legs (Pl. 30, fig.
5) is, I think, very accurate: I observed all the spines here figured,
on the corresponding leg of the pupa of _Balanus Hameri_. The five
posterior pairs of legs differ only in the outer ramus having five
plumose spines, instead of four, and one short simple spine at the
exterior angle, making six altogether. The legs, in their natural
position (fig. 2), have only the terminal segments of their two rami
directed posteriorly; and as a consequence the spine (close to _i_
in fig. 5), borne on the penultimate segment of the outer ramus, is
directed in the same line with that segment and with the pedicel,
namely, anteriorly, and at right angles to the natatory plumose spines.
This short spine acts, I imagine, as a defensive weapon; it has been
omitted in fig. 2. Of the thorax I need not give, from my notes, any
more details. The abdomen (fig. 6) is similarly constructed, as far as
I have seen, throughout the order, with the exception of Alcippe (Pl.
23, fig. 17), in which it is composed of only one segment instead of
three. In this genus the caudal appendages likewise consist of only one
segment, with very short spines. In the pupa of _Balanus balanoides_,
the three spines borne on each caudal appendage are very much more
unequal in size than in the pupa of _Lepas australis_, although in the
latter (fig. 6) the inner spine is considerably thicker than the two
outer. Whether the three segments of which the abdomen is composed, are
the three anterior or three posterior, of the normal seven segments,
I know not: on the view that they are the three posterior segments, I
presume, according to analogy, that the caudal appendages are borne on
the penultimate segment, and that the ultimate segment is here quite
aborted.
On the _internal viscera_ I have nothing to add. The cement-duct is
represented in Pl. 30, fig. 2, _t′_, on the near side, running into
the antennæ; and I repeatedly traced it, for the duct is very strong,
as far as the disc segment; at the other end it joins the cement-gland
(_t_) on the same side of the body. This cement-gland is proved, by
the clearest series of facts, to be converted into the incipient
ovaria and ovarian cæca. The cement-glands in the older pupæ could be
traced as far as the cæca of the stomach, exactly where the ovaria lie
in the mature animal; but in some young pupæ, they extended further
posteriorly, past the mouth, between the outer and inner membranes
of the overlapping carapace. I have faintly shown the course of the
stomach, with its two cæca at the upper end; the anus lies between the
caudal appendages, at the extremity (above _b′_) of the abdomen. At
this age there is no trace of the vesiculæ seminales, so conspicuous in
the mature Cirripede.
_Young Cirripede, whilst within the pupa._--I repeatedly succeeded in
dissecting the young _Lepas australis_ out of the pupa; and by the
previous action of boiling potash, and by a strong light, I was enabled
to make a camera sketch (Pl. 30, fig. 2) of the relative positions of
their several parts. The young Cirripede is drawn very faintly, and is
best seen by holding the plate in the same position with the mature
animal, of which a section is given in my volume on the Lepadidæ, Pl.
9, fig. 4. I may just notice how complicated are the membranes in a
longitudinal section taken at this period: we have, 1st, beginning at
the back, the horny tissue of the carapace or bivalve shell of the
pupa; 2d, the primordial valve (_z_, in fig. 3) of the young Cirripede;
3d and 4th, two folds of corium; 5th, the membrane of the sack of the
Cirripede; 6th, the membrane of the sack of the pupa; 7th, the outer
tunic of the thorax of the pupa; 8th, the outer tunic of the thorax
of the young Cirripede; 9th, the corium lining the latter membrane;
and these nine membranes would be repeated on the opposite side of the
section, if it were taken through either side of the shell or carapace,
bordering the orifice.
After the exuviation of the outer membranes of the pupa, certain
pre-existing coloured marks in the corium, such as those round the eyes
and round the acoustic orifices, along the ridge of the back and on the
borders of the orifice, &c., are still retained by the young Cirripede,
either temporarily or permanently; so that the correspondence of part
with part of the external surface admits of no doubt. Moreover, the
three terminal segments of the antennæ are invariably retained by
the young Cirripede, though in a functionless condition, and into
them the outer membrane of the body, and an important organ, viz.,
the cement-ducts are still prolonged; hence these prolongations must
be considered as aborted antennæ. Again, we have seen that the mouth
of the young Cirripede is formed under the rudimentary mouth of the
pupa, with the new œsophagus, round the old œsophagus, leading into
the same alimentary canal. The twenty-four extreme tips, likewise,
of the six pairs of biramous cirri of the Cirripede are formed
within the twenty-four extremities of the six pairs of biramous,
natatory legs of the pupa. Consequently, in the Cirripede and pupa,
thus far, part corresponds with part, notwithstanding that new eyes
are formed posteriorly to the old eyes, and new acoustic organs in a
quite different position from the old ones; but now we come to a most
important diversity in the metamorphosis, or rather to follow Professor
Owen,[66] in the metagenesis, of the young Cirripede. Although, as
just stated, the extremities of the cirri are formed within the legs
of the pupa, yet, from the great length of the cirri, they occupy more
than the whole of the thorax of the pupa; so that the thorax of the
young Cirripede is not formed within the pre-existing thorax of the
pupa, but within that part of the pupa, (homologically a portion of the
first three cephalic segments), which lies anteriorly to the thorax.
As a consequence of this, the pedicels and lower portions of the
cirri, the segments of the thorax and its dorsal surface, all come to
occupy a position at nearly right angles to that of the corresponding
parts in the pupa: this is shown in Pl. 30, fig. 2. And as a further
consequence, (and this is the more important point), the sack, which
both in the young Cirripede and pupa is formed by the overhanging and
produced portion of the carapace, and which is _internally lined by a
reduplication of the membrane of the thorax_, is necessarily, owing to
the changed position of the thorax, altered in extent and carried much
further; namely, from extending merely parallel to the longitudinal
axis of the pupa (from _b_ to _b′_), it is now in the young Cirripede,
in addition, carried (to _s′_) almost quite across the inside of the
animal. Hence it comes that the young Cirripede is, as I have said in
my former volume, internally almost intersected; and its body remains
attached only by a small space, (see the broken line, round _a_ and _b_
in Pl. 25, fig. 1, of a Balanus with the shell, &c., removed from one
side), to the sternal or ventral, inner surface of the carapace,--the
carapace being modified either into the capitulum and peduncle, or into
the shell with its operculum and basis. As a still further consequence
of this change of position of the body of the young Cirripede within
the body of the pupa, the alimentary canal becomes shortened to fully
half its former length. At the same time, the interspace between the
mouth and first pair of legs of the pupa, (consisting of the seventh
and eighth segments of the archetype), is quite lost in the Cirripede
by coalescence. The final cause of the thorax of the young Cirripede
not being developed within the thorax of the pupa, probably is, that
the cirri may be formed of considerable length, so as to be immediately
enabled to seize prey; and that the thorax, which supports the cirri
(and this probably is even more important) should be as free as
possible within the sack, so as to aid the captorial action of the
cirri.
[66] 'Parthenogenesis,' pp. 13 and 26.
After these remarks, more especially with regard to the formation
of the sack, if any one will look at the sectional drawing of a
pedunculated Cirripede in my former volume, or of a sessile Cirripede
(Pl. 25, fig. 1) in this present volume, in which latter the shell adds
to the complexity, he will perceive the cause of the extreme difficulty
in understanding the relative position of the parts throughout the
whole class. Even after I had discovered that the prehensile antennæ
of the pupa might always be found in the centre of the basis or
surface of attachment, and which fact, it might have been thought,
should have convinced me that this was the anterior end of the whole
animal, yet still I fancied that the prominent mouth represented the
entire head, and that the shell was something quite distinct. It is
clear that others have been equally perplexed; for that which is the
anterior end in the eyes of one naturalist, is the posterior end in
the eyes of another; so with the dorsal and ventral surfaces: one
naturalist considers the peduncle of the Lepas as the abdomen; another
considers it as a pair of metamorphosed, thoracic limbs, &c.! The
probable position of the segments of the body of a mature Cirripede,
in relation to the three anterior cephalic segments, or carapace, is
shown in the diagram (Pl. 25, fig. 6) of the supposed position of the
mature Proteolepas within its pupal envelopes. Here, in the diagram,
the two segments immediately succeeding the mouth (_c_), which are the
seventh and eighth of the archetype, (for the mouth consists of three
segments, and all in front of the mouth of three other segments),
have come to adhere by their dorsal surfaces to the internal surface
of the carapace,--that is, of the first three segments, which ought
of course to have stood quite in advance of these two segments, and
these two segments again ought to have stood in advance of the mouth.
The mouth is directed posteriorly, instead of from the body; and the
three segments of which it is formed (closed at their anterior end by
the labrum), and are very small compared to the relatively monstrously
great, three anterior cephalic segments, composing the carapace. To
place the segments of the body of Proteolepas in proper sequence, in
respect to those of the carapace, and in accordance with the sequence
of the archetype Crustacean, it would be necessary, by seizing the
extremity of the abdomen (_a_), to tear the two segments succeeding
the mouth from their dorsal attachment, as far back as the basal
margin of the labrum; and then pull them till they stood posteriorly
to (or in the diagram, above) the mouth; which latter part would,
by the same movement, be made to project out at right angles to the
ventral surface, and would then be preceded only by the first three,
great, confluent segments of the head, which being produced backwards,
form the carapace. All that has just been said on the position, in
Proteolepas, of the segments of the body in relation to those forming
the carapace, I believe to be applicable to all ordinary Cirripedes,
with this difference, that in the latter, after the metamorphosis,
the two segments succeeding the mouth quite disappear on the ventral
surface, and dorsally are either aborted or have coalesced with the
adjoining segments.
_Act of Metamorphosis._
When the due time for the act of metamorphosis has arrived, the pupal
carapace splits along the dorsal ridge, and is cast off, together with
the acoustic sacks, the basal segments of the two antennæ, and the
great, black, compound eyes, hanging to the UU-like apodemes. The three
terminal segments of the antennæ invariably remain cemented to the
surface of attachment. The exuviæ usually continue for a time united to
the cemented antennæ, but are finally washed away. Besides the split
along the dorsal ridge, the carapace separates, all round the orifice,
from the delicate tunic lining the sack and investing the thorax and
natatory legs of the pupa; for these membranes are not moulted for some
considerable time afterwards. Hence all these inner parts retain for a
period the appearance and structure of the natatory pupa, whilst the
exterior resembles, in every respect, a fixed and perfect Cirripede.
In my former volume, I have insisted on the important and curious
results which ensue from the eye-apodemes penetrating so deeply into
the body (see Pl. 30, fig. 7, in which the proportions are more correct
than in fig. 2), with the eyes attached exteriorly to their outer
arms; for as these apodemes have to be ejected, the external membrane
of the young Cirripede (Pl. 30, fig. 2) has to be formed in a deep
fold or arch over them, and consequently the membrane on the sternal
surface is formed considerably longer than on the dorsal surface. From
this it follows, when all the membranes are free and are stretched
fully out after the moult, that the whole animal, posteriorly to the
cemented-down surface, turns vertically up, and assumes its normal
position at right angles to the surface of attachment, and to that
which it held in its pupal condition; for the pupa always adheres with
its sternal surface parallel to the surface of attachment. A young
Lepas, which has just moulted its pupal carapace, and has assumed its
proper vertical position, with the cemented antennæ and the surface of
attachment remaining as before, is shown at fig. 3, but is drawn on a
smaller scale than the pupa fig. 2, out of which it may be supposed
to have been excluded. In this fig. 3, it may be observed that the
natatory legs and caudal appendages of the pupa have not as yet been
moulted. The fact of the stretching out, in the young Cirripede, of the
fold of membrane, which in pupa, just before the metamorphosis passes
over the apodemes and eyes, is well shown by three darkly-coloured
bands in the corium, which in the pupa are curled, but after the moult,
are stretched straight out on the peduncle of the young Lepas.
The pupa, and consequently the young Cirripede, from being attached
at first by the antennæ, does not adhere by the actual anterior
extremity, but by the sternal surface near it; the anterior extremity,
however, soon becomes cemented down, and afterwards, in ordinary
cases, ceases to grow. In Cryptophialus, however, and in certain
genera of the Lepadidæ, as Alcippe, Lithotrya, and Anelasma, the
anterior or basal extremity does continue to grow, and is not cemented
down, and therefore comes to be prolonged beyond the original point
of attachment; in order to allow of this, the surface to which the
Cirripede is attached has to yield, apparently simply to the pressure
exerted in the case of Anelasma, but in the three other genera, to
the rasping action of the roughened surface of the extremity of the
peduncle.
When after a period the pupal membranes of the sack, thorax, and
natatory legs are moulted, the cirri of the young Cirripede are
curled up, and its thorax is raised towards the orifice, and we have
the animal in its ordinary position, and perfect with the exception
of a few parts to be further developed or modified. For, instead of
calcareous valves, we have at this period only the so-called primordial
valves, composed of chitine; and in the case of _Lepas australis_,
some minute spines and some coloured marks on the peduncle, which soon
disappear. The muscles, which enter the three terminal segments of the
antennæ in the pupa, have to be absorbed and converted into ligamentous
threads. In Lepas, the labrum has to become bullate; and the cæca have
to increase in number round the upper end of the stomach, and their
dark colour and that of the whole alimentary canal has to disappear or
be much weakened. The filamentary appendages at the bases of the cirri,
which generally contain some of the testes, have to be developed.
The probosciformed penis, which at first equals only the pedicels of
the posterior cirri in length, and is apparently imperforate, has to
increase greatly in length. The testes and vesiculæ seminales have to
be formed. And lastly, and this is a more important point, the two
gut-formed cement-glands (or incipient ovaria, _t_, fig. 2, Pl. 30)
which, at the period of the moulting of the carapace and eye-apodemes,
and when the whole animal was upturned, came to occupy, together
with the cement-ducts (_t′_), their normal position, _i. e._ nearly
parallel to the sternal surface, now undergo further changes. Their
upper and posterior ends lying near the cæca of the stomach, increase
in size, but retain nearly the same character, and thus form the two
true ovaria; their middle parts become emptied of their cellular
contents, and are converted into the two simple ovarian tubes; and
their lower ends branch out, inosculate, and form the inextricable mass
of ovarian tubes and cæca. The points of junction on each side between
the two cement-ducts and the newly branched ovarian tubes, become now
developed into the two cement-glands. The cement-ducts, which continue
throughout life growing, either still enter the old antennæ and there
pour out the cement-tissue, or they pour it out through special
orifices formed for this purpose in the lower part of the peduncle. The
changes, supervening during the metamorphosis, in the ovaria and in
the cementing apparatus, here described, I have no doubt are general
throughout the Order.
I have above alluded to the _primordial valves_; these are beautiful
objects when viewed under a high power: they are composed of chitine
without a trace of calcareous matter, but prefigure in shape, size,
and direction of growth, the shelly valves soon to be formed under
and round them. They are composed of an outer membrane, with its
margins separated by yellow thickened rims from the membrane uniting
the several primordial valves together; and this outer membrane is
underlaid by a single layer of generally hexagonal, thickish cells (Pl.
30, fig. 3 _a_), varying from 1 to 2/6000th of an inch in diameter.
These cells seem to contain a nucleus; and they are at first separated
from each other by clear interspaces. If a specimen be taken, only
a little before the formation of the calcareous valves, one or more
layers of membrane, marked by an hexagonal reticulation, can be
separated from the lower surface of the main hexagonal network. It
is a singular fact, that in those genera in which there are several
valves, the primordial valves occur only on five, namely, on the two
scuta, two terga, and the carina; and these are the most persistent
valves in the several genera. The other valves are prefigured only by
brownish membrane, without the hexagonal tissue. In the mature _Lepas_,
the membrane connecting the several shelly valves is not moulted, but
disintegrates; in the primordial valves, however, which stand far
separate from each other, this membrane is moulted; and immediately
after the first moult, the first layer of shell appears under and
a little way beyond each primordial valve; shelly matter likewise
appears, at least in some cases, between the cells of the hexagonal
tissue. The young shelly valves are connected together, at each
successive moult, by narrower strips of membrane, till, in the case of
Lepas, the valves when mature come to touch each other (Lepadidæ, Pl.
1, fig. 5). The primordial valves are often preserved for a long time
on the umbones, or centres of growth of the five valves, on which they
occur, in the same manner as the larva-shell is sometimes preserved on
the apex of certain spiral molluscs. Had not Cirripedes gone through
so many and such complicated metamorphoses, this last state, when
furnished only with primordial valves and with several internal organs
only partially or not at all developed, would have deserved to have
ranked as a special stage, and not as merely subordinate to the last or
pupal condition.
In the Balanidæ, or sessile Cirripedes, the young animal, immediately
after the metamorphosis, or still better if dissected out of the
pupal carapace, as I succeeded in doing with _Balanus balanoides_,
may be said to be pedunculated; for it is attached by a little disc
of cement closely surrounding the antennæ, the rest of the membranous
basis forming an almost semi-globular, flexible peduncle. The valves,
at this the earliest period, are all membranous, and do not overlap
each other. In the Balaninæ they do not present the peculiar structure
of the primordial valves of the Lepadidæ; but in the Chthamalinæ, in
Chthamalus, I saw traces of this structure. Calcareous valves are
soon formed under the membranous valves. The opercular valves, at
this early period, are much larger than the valves or compartments
of the shell, which are only four in number, for the carino-lateral
compartments are not yet formed. The compartments from the first are
surprisingly strong, and have their alæ already formed and overlapped
by the adjoining compartments; but of the radii there is as yet no
trace. The four compartments form a narrow but nearly circular hoop,
which, from its relatively large diameter, tends to draw down the
upper or posterior end of the animal, now forming the opercular valves;
and as the basis soon becomes throughout cemented to the surface of
attachment, the young Cirripede is much depressed. Soon the opercular
valves are drawn a little way down within the shell, becoming attached
to the sheath, instead of, as at first, to the very summits of the
compartments. In regard to the changes which take place in the shell,
in the number of the segments in the cirri, and in the number of spines
borne on these segments, &c., during the continued growth of the
animal, as they are chiefly important for the identification of the
species, I will here refer to a discussion on this subject under the
head of the Genus Balanus.
_On the Homologies of the Carapace and Shelly Valves._
In the pupa, the carapace is produced, not only posteriorly, but
anteriorly, so as to cover the entire animal, with the exception of a
narrow sternal surface (Pl. 30, fig. 4): in front it is notched, where
the sternal surface terminates, and from this notch a faint line runs
along the dorsal surface, separating its tergal elements. In the young
Cirripede, after the metamorphosis, there is no trace of this medial
dorsal suture, or of the wider sternal surface. Looking at the several
genera of the Lepadidæ, the external covering of the whole peduncle and
capitulum is so continuous and of so uniform a nature, that I think we
must consider the whole as a carapace, of which the sternal borders
have become completely confluent; formerly I was inclined to look at
the capitulum alone as formed by the carapace, and at the peduncle as
being composed of the two or three anterior cephalic segments, cased
only by their own integuments. As far as can be discerned, the carapace
in the pupa, and consequently in the Cirripede, consists only of the
tergal elements of the segments; and this seems likewise to be the
case with the carapace of the Podophthalmia. Until lately,[67] Prof.
Milne Edwards doubted whether the carapace in the higher Crustaceans
(to which I believe the carapace of Cirripedes must be compared) was
formed by the backward production of the third segment, which bears
the second pair of antennæ, or of the fourth, _i. e._ the mandibular
segment; but from the distribution of the nerves, he now argues that
it must mainly belong to the third segment. In Cirripedes, the course
of the nerves leads to the same conclusion; for the whole shell, sack,
and peduncle are supplied with nerves proceeding from the compounded
ganglion, which belongs to the second and third cephalic segments.[68]
[67] Compare 'Histoire Naturelle des Crustacés,' tom. i, p. 27,
with 'Annales des Sciences Nat.,' 3d series, tom. xvi, 1851, p. 233.
[68] This conclusion is supported by the structure of Proteolepas:
in this Cirripede there is not a vestige of a carapace, and as
the whole of the animal in front of the mouth is almost utterly
aborted, being reduced to a mere covering to the two cement-ducts,
and as, on the other hand, the mouth with the mandibles, though
peculiarly modified, is not at all aborted, there is a strong
probability, that the abortion of the carapace is connected with
the aborted state of the three anterior cephalic segments; and that
the carapace in its origin is not any way related to the fourth or
mandibular segment.
The whole of the head in front of the mouth, together with the
carapace, is, as we know, formed of three segments; and each of these
segments, homologically, ought to consist of eight elements; I recall
to mind these facts, inasmuch as the transverse separation between the
peduncle and capitulum in the Lepadidæ, and between the basis, the
shell, and the opercular valves in the Balanidæ, might be thought to be
connected with the separation of the three cephalic segments. So again,
as in the Balanidæ the shell normally consists of eight compartments,
these might be thought to be related to the eight elements of one or
other of the three segments. But I see no reason for admitting this
view; and in the case of the carina and rostrum, two of the most
persistent and important of the compartments, they exactly cover the
sutures which ought to separate the two tergal and two sternal elements
of the segment. The valves, moreover, often form many more whorls
than three, or the number of the true cephalic segments in front of
the mouth; and in each whorl the valves tend to stand in tile-like or
alternate order, with respect to those in the whorls both above and
below, which would not be the case, if they were the eight elements of
the segments.
For the true homologies of the sclerodermic plates, or Shelly valves,
with which the external covering of Cirripedes is so generally
strengthened, we must, I believe, look to the carapace of the
Podophthalmia. In these latter, we find the carapace composed of
sclerodermic plates, which, though closely joined and only occasionally
separated by sutures, yet in their origin are distinct;[69] they
tend, also, to be arranged in alternate or tile-like order. As the
animal grows, the old sclerodermic plates, all joined together, are
moulted, and new ones, also all joined together, of a larger size,
are formed beneath. Now let us imagine the growth to be more gradual
but yet periodical, and the new and larger sclerodermic plates, when
formed under the old ones, to adhere firmly to them; the older plates
would thus be prevented from becoming confluent, and instead of
being all moulted together, as is now the case, they would be almost
continually separated from each other, owing to the almost continuous
increase in size of the new underlying plates. Consequently lines of
splitting would run between the several plates, however numerous they
might be, instead of there being, as now, a single line of splitting
extending down the back. In fact, we should have the identical manner
of growth of the shell or carapace, which occurs in Cirripedes. It is
on this ground, and from the several points of homological resemblance
incidentally mentioned in the last few paragraphs, that, in the early
part of this Introduction (p. 13), when discussing the whole class, I
stated that I believed that the carapace of Cirripedes presented more
real resemblance with the carapaces of the Podophthalmia, or higher
Crustacea, than with those of the lower Crustacea, though in mere shape
they more nearly resembled the latter.
[69] 'Annales des Sciences Naturelles,' 3d series, tom. xvi, pp.
233, 236, 237.
_Cementing Apparatus._ (Plate 28.)
I have already (p. 128) given an account of the manner in which, in the
pupa of Lepas, the cement-tissue escapes from the prehensile antennæ,
and of the structure of the cement-ducts, and of the cement-glands
or incipient ovaria; and likewise of the changes by which these
organs assume their ultimate form in the mature Cirripede. In my
former volume, on the Lepadidæ, I described the cement-glands and the
cement-tissue in several genera, and I have there shown (singular as
the fact is) that the two cement-glands, with their contents, actually
consist of ovarian tubes with their contents (for there seemed to be
a relation in the state of fulness in both) in a modified condition.
In the Balanidæ, I am not able, from the difficulty of the dissection,
to confirm these conclusions, excepting in so far that the tubes on
which the cement-glands are formed, run into the mass of ovarian cæca;
but, I may add, that in the abnormal Proteolepas, belonging to another
Order (see the section, Pl. 24, fig. 1), nothing could be plainer than
that the membrane of the ovarian sack (_b_) formed the cement-ducts,
and that their cellular contents, which within the sack (_a_) were in
process of conversion into ova, within the ducts were converted into
the cement-tissue. This cement, by some unknown power, travels down the
ducts, and debouches at the antennæ.
In the Lepadidæ, there are only two cement-glands, which are situated
high up in the midst of the ovarian cæca, with one cement-duct
proceeding from each: both the glands and ducts increase in size with
the age of the animal:[70] the cement issues either permanently from
the prehensile antenna, or, after a short period, through apertures
in the peduncle, arranged irregularly or in straight lines,--the last
formed apertures being furthest from the central and basal point of
the peduncle. In the Balaninæ, on the other hand, at each period of
growth, a pair of new cement-glands is developed, larger than those
last formed, and making, with the older glands, a chain, connected
together by what I have called the cement-trunk. The cement-trunk
consists of a tube, which generally becomes enlarged just before
entering each gland. The glands, the cement-trunk, and cement-ducts,
all adhere to the basal membrane or basal shelly plate. Each gland
gives rise to two cement-ducts, these often bifurcate, and sometimes
repeatedly bifurcate and inosculate before pouring out their contents
round the circumference of the basis; and sometimes they all first
enter a circumferential cement-duct. The probable cause of the greater
complexity of the cementing apparatus and of the greater number of the
excretory orifices in the Balanidæ, compared with the Lepadidæ, is
that the entire surface of the broad basis, which answers to the whole
peduncle in the Lepadidæ, is firmly cemented down to the supporting
object, instead of merely the basal end of the peduncle. The cement
issues either in a cellular condition, or more commonly as a fine
network, which, at a short distance from the orifices (Pl. 28, fig.
4 _a_, _z_), becomes so fine as to form a sheet or layer: I may here
recall the fact, that in the cement proceeding from the disc of the
antennæ, in some species of Lepas, a similar structure was observed.
The cement itself presents the same transparent, brown, laminated,
structureless appearance, and the same chemical reaction, as described
in my former volume. The cement has the capacity of occupying and
filling up all inequalities in the supporting surface; I have seen it,
when spread over an encrusting Flustra, present an exact model of every
cell; in the case of Coronula, it seems, as we shall immediately see,
to have the power of penetrating into, and even almost blending with
the epidermis of the supporting Cetacean. The last-formed cement-glands
and cement-ducts present a delicate and transparent appearance, and
contain cellular matter; whereas the old cement-glands, and sometimes
the old cement-ducts, are filled with brownish cement, not acted
on by boiling potash. The foregoing remarks are confined to the
sub-family Balaninæ, for I have not been able to examine thoroughly the
Chthamalinæ, and can only affirm, that in Chthamalus and Pachylasma the
cement-ducts repeatedly bifurcate and inosculate, in the same manner as
in the Balaninæ. I will now proceed to describe, in some detail, the
cementing apparatus in the several following genera.
[70] I had some slight reason to suspect in Pollicipes that new
cement-glands were successively formed: this is more probable in
this genus than in the others, inasmuch as it is the most nearly
related to the Balanidæ.
_Coronula._--The cementing apparatus is here more simple than in any
other genus of the Balaninæ, and I have studied it more carefully.
The basal membrane of _Coronula balænaris_ is figured in Pl. 28, fig.
1 _a_, and must first be described; its relation to the shell will
hardly be understood without looking at the outline of the folded walls
of this species, in Pl. 16, fig. 5. The basal membrane closes the
central circular hollow, and is continuous with rays (not represented
in Pl. 28) extending under the doubled walls and terminal transverse
loops. It has eighteen concave sides, corresponding with the inner
ends of the folded walls, for each of the six compartments is trebly
folded. The membrane consists of successive, conformable slips (_c′_,
_c′_), bordered exteriorly by thickened yellowish rims, and internally
overlapping (when viewed from the inner side) the few last-formed
slips, and then thinning out. The membrane forming each slip is itself
laminated. The middle portion, about 1/50th of an inch in diameter,
is rather opaque, owing to the slips being so close together. Beyond
this central part, the slips suddenly increase in size, but yet have
a different shape from the 18-sided outline, which they ultimately
assume: this difference is owing to the great changes in shape, as
explained under the genus Coronula, which the shell undergoes, when the
walls at first assume their folded structure. The walls are invested
by longitudinally striated membrane (_p_, _p_, _p_, fig. 1 _a_), which
turns in under their basal edges; and this membrane is united with the
basal membrane, by what I shall call the circumferential slip (_b_),
and which is shaded in fig. 1 _a_, simply for the sake of catching
the eye. It is the circumferential slip of membrane which sends
rays under the spoke-like folded walls: thin as it is, this slip is
yet laminated, but is not bordered by thickened edges. The membrane
investing the walls is, like the basal membrane, formed of successive
slips with thickened edges, which overlapping (viewed from the inside)
the last-formed slips, project beyond them, and so face the edges of
the slips in the basal membrane; they are only obscurely indicated in
fig. 1 _a_. The circumferential slip (_b_) lies over (as viewed from
within) both the basal and wall membrane. This whole structure will,
perhaps, be best understood by the sectional diagram (fig. 1 _b_),
in which the letters (_c′_, _c′_) show the slips of basal membrane;
(_p_) the parietal membrane, coating the outside surface of the walls
of the shell, not here represented; (_b_) the circumferential slip
overlying both; and (_z_, _z_) the layers of cement, which may for
the present be disregarded. In order to allow, of the growth of the
shell, the circumferential slip (_b_) periodically splits in the
middle, all round, in a line exactly conformable to the edge of the
last-formed slip of basal membrane; and likewise in straight, medial
lines under the spoke-like (cut off in fig. 1 _a_) doubled walls. I
have seen, under a high power, the line of splitting, very shortly
after its formation, with the two edges ragged and near together, with
an extremely narrow, new circumferential slip just formed, between
and over the edges of the previously formed slip. What causes the
circumferential slip to split so symmetrically, I cannot say: the
opposed edges, after a time, become thickened, apparently from adhering
to the underlying layer of cement, as will presently be described. The
circumferential slip continues increasing in breadth till the period
of its splitting arrives, by which time it has become much broader
than the last-formed slip of basal membrane; and after the splitting
takes place, the interior half towards the basal membrane, forms a
new basal slip all round the basis, and the exterior half adds a new
slip to the membrane investing the walls. This latter membrane being
inflected under the basal edges of the walls, is, during the growth of
their edges, drawn straight down, the newly-formed portion taking the
inflected position.
In the sectional diagram, (1 _b_) the circumferential slip is not yet
broad enough to split; when it has become so, it will split under
the letter (_b_). The slips of basal membrane are, as may be seen in
fig. 1 _a_, narrower towards the circumference; but the two or three
last-formed slips, are out of proportion narrower than the others; and
it is certain, from the comparison of the basal membranes of specimens
of different ages, that these will afterwards increase in width.[71]
I have seen no other instance, in Cirripedes, of growth in membranes,
except at their extreme margins: I suspect that these last-formed slips
are pulled, during the downward and outward growth of the shell, a
little from over the last-formed slips, new and larger laminæ being
all the time thrown down, so as to prevent any fissure being formed. I
also suspect that the gradual increase in width of the circumferential
slip itself, is due to the opposed edges of the underlying and
last-formed circumferential slip being dragged further apart from each
other, new and wider laminæ of membrane being continually thrown down;
the new circumferential slip being thus, also, all the time thickened,
as well as rendered broader.
[71] In the case of one _young_ shell, I found that the
previously-formed circumferential slip must have split, long before
it had assumed its proper and ordinary width; for the last-formed
slip of basal membrane was of extreme narrowness, and would have
to be considerably added to in width, whilst the new and narrow
circumferential slip was likewise being added to in width. This
slip of basal membrane, though so extremely narrow, had its own
cement-ducts and glands.
The central slip or rather disc of membrane, is 3/400ths of an inch in
diameter; and this shows the basal diameter of the shell immediately
after the metamorphosis. In the middle of this little disc I saw,
in several specimens, the prehensile, pupal antennæ; I made out
distinctly the ultimate segment with its bristles, and, as I believe,
the disc-segment, which was 7/2000ths of an inch in diameter; but this
portion was much obscured by the quantity of cement. When the corium
is removed from the inner side of the basal membrane, the two chains
of glands, extending from exactly over the antennæ in the centre about
half way towards the circumference, are conspicuous. The cement-trunk,
connecting the glands, is thin, and at the further end is always broken
off, by the removal of the corium and overlying layer of ovarian cæca,
into which the two cement-trunks enter; and without which removal,
nothing could be seen. The two chains of glands form a very large
angle, open towards the rostral end of the shell, as represented at
fig. 1 _c_, much enlarged; by a mistake in fig. 1 _a_, the two have
been drawn in a straight line. The cement-trunk increases in diameter
in proceeding from the centre to the circumference, and the glands
likewise increase in size, at the same time altering somewhat in shape.
From near the lower side (the basal membrane being viewed from within)
of each gland, two cement-ducts proceed, which pour out their contents
beneath the basal membrane. The orifices of the ducts always exactly
face the middle folds of the two lateral, and two carino-lateral
compartments. In a full-sized specimen, there are from thirty-five to
forty cement-glands on each side, always corresponding exactly with
the number of slips of basal membrane, including the circumferential
slip, to which the last-formed pair of glands and cement-ducts belong.
In correspondence with the great number and narrowness of the central
slips of membrane, so are the cement-glands towards the centre numerous
and very small. All the glands, in the more central parts, consist of
a mere transverse enlargement of the cement-trunk; but the exterior
and larger glands, which are more closely packed together, are more
globular or pear-shaped; and the two ducts (of which the one on the
rostral side is considerably enlarged at its base) do not come out of
the gland exactly at the same level. The trunk, connecting the glands,
runs straight from one to the other. The ducts proceeding from the
outer and older glands, on the carinal side, are much curved (Pl. 28,
fig. 1 _c_). To give an idea of the dimensions of the several parts, I
may state that the largest ducts were 3/1000ths of an inch in diameter,
and the glands belonging to them nearly thrice as much, measured in the
direction of the cement-trunk; on the other hand, some of the ducts
from the small central glands had a diameter eighteen times less than
that of the largest ducts. Towards the circumference, the ducts that
proceed from the older and larger glands are piled one exactly over the
other--the last formed being the topmost, and all are imbedded in the
corium which overlies the basal membrane: this position of the ducts,
one over the other (which could not be well represented in figs. 1 _a_
and 1 _c_), is owing to their all debouching at the same exact point.
But the ducts form the smaller and younger glands, when the shell
had a different shape, are spread out, and are all attached to the
basal membrane. Altogether, the basal membrane of Coronula, when well
cleaned, and examined under a moderately high power, presents the most
singular and elegant appearance.
We now come to the cement-tissue: this lies on the under or outer
side of the basal membrane; it is not represented in figs. 1 _a_ or
1 _c_, but only in the sectional diagram, 1 _b_, by the letters _z_,
_z_: it presents its usual character and appearance, like solid glue
or brown gum, but is obliquely laminated and sub-laminated: it forms
a layer, much thicker than the basal membrane itself, being as much
as .004 or .005 of an inch in thickness. It is, however, difficult
to ascertain its thickness, from the singular manner in which it
penetrates into and almost blends with the epidermis of the whale's
skin; so much so, that for a considerable time I thought (not then
knowing anything about the cement of Cirripedes) that this transparent
horny substance probably answered to a corn on the human foot produced
by pressure. But I soon observed that this horny substance certainly
extended into and up the cement-ducts; and this fact first led me to
the examination of the whole subject in the several genera of Lepadidæ
and Balanidæ. It was not difficult to remove the cement-ducts, leaving
small portions of the contained cement projecting freely up as points
from the general surface of cement. The cement adheres slightly to the
whole basal membrane, but chiefly to the yellowish rims or edges of
the successive slips; and it is indeed this adhesion which, I believe,
produces the rims; for the circumferential slip, when first split, had
very thin ragged edges. The cement also extends under the spoke-like
prolongations of the circumferential slip, and likewise some way up the
sides of the walls.
The cement-glands, the trunk, and the ducts, except the two, three,
or even four last-formed ones, are all filled with an apparently
solid thread of transparent, brownish cement, differing in no respect
from the cement under the central parts of the basal membrane. In
one instance, in which the last-formed pair of glands and ducts had
apparently been only just developed, they were so perfectly transparent
that I could distinguish them only under certain lights, and I could
not perceive any contents. The last-formed glands and ducts always
appear very delicate, and include a tube of very delicate tissue,
containing more or less of granular matter. The next succeeding pair of
glands and ducts are always more opaque, and contain much more granular
matter; which, in the next, or next but one, may be seen passing into
the state of pale brown, transparent chitine. I have seen some most
distinct instances, in which, in the same duct, the part towards the
centre of the basis was filled with homogeneous cement, and the part
towards the circumference with still plainly granular matter. In the
successive circular slips of cement-tissue, lying attached under the
circumferential slip and under the two or three succeeding slips of
basal membrane, an exactly analogous series of changes is presented; as
indeed might have been expected, as the slips of cement are absolutely
continuous with the contents of the ducts. Moreover, if a vertical
section be made across one of the last-formed slips of cement, it
may sometimes be seen to be apparently in the act of separating into
layers, with the lower layers more pulpy, elastic, and white than the
upper layers, which are less coherent, and show as yet even still less
the character of cement. The cement under the circumferential and
adjoining slips, often presents a peculiar wrinkled appearance, in
lines conformable with the outline of the basal membrane; but I do not
believe that these are real wrinkles, though so like them; they seem
to consist of sinuous threads, longer or shorter, sometimes slightly
branched, crossing and inter-joined, and composed apparently of faintly
coloured cement. I suspect that these threads are formed by the union
and subsequent drawing out of aggregations of that matter, which within
the ducts is first granular, and then changes into cement; for at the
orifices of the ducts these wrinkled threads sweep outwards in curved
lines on both sides. The cement in these early stages adheres, with
very little force, to the basal membrane; and with only a little more
force to the underlying layers of cement; in fact, till it assumes the
brown translucent appearance, like solid glue, it hardly seems to act
as cement.
How the cement reaches the skin of the whale, will be best understood
by referring to the sectional diagram (Pl. 28, fig. 1 _b_). When the
circumferential slip of membrane (_b_) splits, a new circumferential
slip will be formed over it, together with new cement-glands and
ducts, and cement (_z_, _z_) will issue from four new orifices, and
will extend on both sides of these orifices, till the ends meet and
become united, thus forming a narrow, 18-sided, continuous, new slip
of cement, with 18 spokes proceeding from it. I have not noticed lines
of union in the cement of any one slip; but the matter forming each
slip, certainly has proceeded from four distinct orifices. Seeing how
perfectly successive layers of cement often become blended together,
lines of union or junction, could hardly be expected to be preserved
in the same individual layer. When the circumferential slip of basal
membrane splits, the underlying slip of cement, which we will call A,
does not split, but becomes stretched, so that the newly formed slip of
cement, which we will call B, does not reach the skin of the whale. As
the new circumferential slip of basal membranes goes on increasing in
width, A continues to be stretched, but does not split, till at least
another circumferential slip of basal membrane has been formed and
has been split, and till B has been also stretched. By this time, the
cement-tissue A has assumed its normal structure, and has the power of
adhering to the whale's skin, which it has now reached, owing to the
splitting of the under and older slips of cement. At the next period
of growth, A itself will split, and B will touch the whale's skin and
adhere to it; and this, also, will ultimately split. It results from
this action, that the cement has a stretched, and sometimes even a
fibrous appearance, with the lower edges of the layers, of which each
slip of cement is formed, thinning out. I have before stated, that the
two or three last-formed slips of basal membrane are formed at first
too narrow, and apparently have to be dragged outwards, over each
other; and it is perhaps owing to this circumstance, and to globules of
cement having first adhered to the under surface of the slips of basal
membrane, that these slips are studded beneath with parallel little
vermiform bodies, sometimes of considerable length, and furnished
with heads, all directed outwards. These tapering, vermiform bodies
have a considerable resemblance to the threads before mentioned,
which give the wrinkled, concentric appearance to the newly-formed
layers of cement, and have probably a closely analogous origin: in
one case, indeed, it appeared as if some of these concentric threads
were in process of being drawn out at right angles to their original
course. Lastly, it should be observed, that as the exterior half of the
membrane of the circumferential slip, after each splitting, is dragged
down, and thus comes to invest the outer surface of the wall of the
shell (the wall not being represented in the diagram, but standing
where the letter (_p_) stands), so it must be with the cement, which
thus likewise comes, in an unusual manner, to invest the outer surface
of the folded walls of the shell, and attaches them to the skin of the
whale,--which latter is always growing upwards, and tending to bury the
shell.
_Platylepas decorata._--This genus is closely allied to Coronula,
and the cementing apparatus is essentially similar. In one specimen,
I counted no less than forty-nine slips of basal membrane, each of
which, of course, had its pair of cement-glands, and each of the
latter its two ducts. The glands consists of a transverse enlargement
of the trunk, as in the early stages of Coronula. Neither the glands
nor the duct, when old, become filled up with cement, but only the
main-trunk. The ducts are very delicate and thin; the larger ones being
only 3/10,000 of an inch in diameter. The glands stand some way apart
on the two cement trunks; and the latter, instead of being straight
as in Coronula, are deeply serpentine; the glands are formed on each
bend, so that, though all on one side are connected on the same trunk,
they form a double row on each side of the basal membrane. The basal
membrane (in the centre of which I distinctly saw the antennæ of the
pupa) has six deep bays or excisions, corresponding with the midribs
(see Pl. 17, 1 _a_, 1 _d_) of the six compartments; and the two ducts
from each gland, on the right and left sides, debouch at the heads of
the four lateral excisions, exactly opposite the midribs of the lateral
and carino-lateral compartments. The later-formed glands, owing to
all of them being situated some way apart from each other on the two
cement-trunks, lie further from the centre of the basis than do the
orifices of their ducts; hence the later-formed ducts are directed a
little backwards, or from near the circumference towards the heads of
the deep excisions.
_Tubicinella._--The cementing apparatus is here less symmetrical; but
this, I believe, is chiefly owing to the basal membrane being formed
of successively larger discs (not slips) of membrane, thrown down not
quite concentrically one over the other; each new disc of membrane
seems to cover the last-formed cement-glands and ducts; and there
are as many ducts and glands as there are discs of membrane, all
adhering together. In one specimen, it appeared that normally there
were four sets of cement-ducts, as in the allied genera of Coronula and
Platylepas; but in other specimens, the ducts were distributed very
irregularly. In one case the two cement-trunks extended parallel and
close together, one of them terminating long before the other. I have
given a figure (Pl. 28, fig. 3) of three of the cement-glands, removed
from the basal membrane, together with their ducts. The cement-trunk
(_f f_) seemed to be a little enlarged, and to be crossed by septa, as
it entered the glands (_h_), but I could not make out this structure
clearly enough to be represented. Whilst young, the cement-glands stand
some little way apart from each other; and in the figure given of some
of the last-formed glands, they are hardly separate enough. Each gland
gives out obliquely, on one side, a cement-duct (_c_) which I traced
in several cases to the margin of one of the discs of basal membrane,
where cement issued from it; and on the opposite side, a tapering spur
(_b_), varying in length, which may be called, and I believe really
is, a rudimentary duct. Of these spurs we shall meet many instances
in other genera. The duct (_c_) and the spur (_b_), close to where
they entered the gland, in some specimens gave off, at about right
angles, short blunt points, or rudimentary branches. This duct and spur
correspond, I believe, with the two ducts in Coronula; but besides
these, a duct (_a_) is given off from one end of the gland, from the
surface opposite to that at which the cement-trunk enters. This duct
(_a_) is very singular, from always forming a loop (_a′_), with two
spurs projecting from it: these two spurs occasionally spring from a
common point: I have seen nothing like this structure in any other
Cirripede. This duct (_a_) runs, like the duct (_c_), to the margin
of its own disc of basal membrane, where it debouches. Besides these
ducts, in the best specimen which I examined, there were two other
sets of ducts, which were slightly zig-zag, and at each angular bend,
a mere knob or point, or at most a short branch, was given off; but
this branch seemed never to run to the margin of the basal membrane
or to give out cement; whereas the main branch did give out cement. I
was not able to trace these ducts to their glands. In these zig-zag
ducts, and in the rudimentary points sometimes observed at the base
of the duct (_c_), and likewise at the base of the spur (_b_), we see
the first indication of that tendency to bifurcation, so strongly
characteristic of the cement-ducts in all the genera, excepting those
already described, which are allied to Coronula.
_Chelonobia patula._--The cementing apparatus is here chiefly
remarkable for the thinness and straightness of the main trunk, (_f f_,
Pl. 28, fig. 2), and from the great distance at which the glands stand
apart; had another gland been drawn, it would, on the scale here used,
have stood exactly under the two upper, (_c′ c′_) in fig. 1 _c_. We
here see that the trunk (_f_), before entering the gland (_h_), has an
enlarged portion (_g_); this, I suspect, is a very general structure.
Each gland gives out, on opposite sides, two ducts (_a a_, _b b_),
larger even than the main trunk; and these ducts bifurcate repeatedly,
and inosculate. By this inosculation it is not improbable that all four
ducts, proceeding from the two glands of the same age, may be connected
together; certainly the bifurcating branches from the same duct thus
become repeatedly connected. For the first two or three bifurcations
the ducts decrease very little or not at all in diameter; but nearer
the circumference they become smaller. The ducts, also, proceeding
from the younger and smaller glands, are, of course, proportionably
smaller. In one case I was able to count four bifurcations in the duct
between the gland and the edge of the basal membrane. It follows from
this structure, that the basal membrane, at each period of growth,
is cemented down by cement issuing from several orifices; but we
shall presently find that in other genera the cement proceeds from
many more orifices. In fig. 2 there is represented, by the aid of the
camera, a small portion (from the outer (_a_) to the outer (_b_) being
12/100ths of an inch in length) of the basal membrane, with all the
several cement-ducts adhering to it, which I could distinguish, and
drawn of their proper relative sizes; this figure also shows some of
the bifurcations, but no inosculation happened to be included in the
space here given; the basal membrane itself has not been represented.
In taking a view of a considerable portion of the basal membrane,
especially towards the circumference, some parallelism in the branches
could be perceived; one set of branches tending to run in the direction
of the ray of the circle, and the other set in the line of the
circumference.
_Elminius Kingii._--The cement-glands here resemble those of
Chelonobia, but the trunk does not seem to be enlarged before
entering the gland. The glands are situated rather far apart; and
the chief peculiarity is, that the trunk connecting the glands is
as tortuous as the track of a worm. Each gland gives out two ducts,
which bifurcate repeatedly, and often inosculate, making, in parts,
an hexagonal mesh-work: some of the branches do not debouch on the
basal membrane, but terminate in blunt points. So numerous are the
ducts, that the basal membrane may be compared to pieces of paper
with the fine fibrous branching roots of some plant dried and heaped
on it. Some of these ducts are very regularly jointed, and resemble
a conferva,--an appearance which I believe is owing to divisions in
the contained cement; other ducts are partially marked by little
wrinkles, as presently to be described under Balanus. The cement,
instead of, as heretofore, invariably forming a slip round and beneath
the circumference of the basal membrane, here often forms little,
independent, circular, and irregularly-shaped discs, each of which has
issued from a single orifice. I may here add that in two species of
_Tetraclita_ I saw the cement-ducts repeatedly bifurcating, with some
of the branches inosculating, as in Elminius and Chelonobia.
In _Balanus balanoides_, which, like all the Cirripedes hitherto
mentioned, has a membranous basis, I could only make out an amazing
number of bifurcating and inosculating cement-ducts, of very various
diameters. The cement-tissue, on the under side of the basal membrane,
generally consisted of little circular discs, on an average from 2 to
5/1000ths of an inch in diameter; but there were also globules and
short tortuous threads of cement. In some very much elongated and
crowded specimens--in which, during the downward growth of the walls,
the basal membrane had ceased to reach the surface of attachment, and
being thus suspended had become, as viewed from the outside, deeply
concave--the cement had apparently continued to try to reach the rock,
and now hung down in the form of two thickish roots, some tenths of
an inch in length. These roots were round, and tapered either to a
fine or blunt point; one was doubled on itself, and so had become
united; in the other, I could perceive five layers or sheaths of the
cement-tissue, one within the other; the innermost of these layers,
which once, no doubt, formed the outside surface of the root, was only
a quarter of its present length.
In _Balanus tintinnabulum_, the basis is calcareous: when its upper
surface is cleaned, dried, and examined under a good light, the
numerous larger cement-ducts can be seen, even by the naked eye, or
under a weak lens, and present an elegant appearance. These larger
ducts run in parallel lines from the two chains of glands towards the
circumference. They are all encrusted with calcareous matter, and in
the more central parts are hidden under it; at each period of growth,
when the basis is added to round the circumference, it would appear
that a layer of excessive tenuity of shell is thrown down over the
whole surface, just in the same way as in Tubicinella, at each period,
a new and larger disc of membrane was thrown down over the pre-existing
membranes with their cementing apparatus. The cement-glands, in the
middle of the basal plate, seem often to give rise to small abnormal
depositions of calcareous matter. When the basis (it is best to take
a young specimen) is slowly dissolved in acid, all the cementing
apparatus is left uninjured, adhering to the delicate tissue which
before existed in a calcified condition. Near the middle I saw the two
antennæ of the pupa; and from them the two cement-trunks extended about
half-way towards the circumference. These two chains of glands are
often placed very irregularly, but they tend to form, as in Coronula,
a large angle, open towards the rostral end of the shell. The glands,
close to the old antennæ, commence abruptly, of rather large size: the
later-formed glands, with their ducts, are in regular order larger than
the younger ones, and stand much closer together. After immersion in
acid all the glands and ducts appeared empty, instead of the older
ones being, as in Coronula, filled with cement. In one case I counted
on each trunk twenty-five glands, besides some smaller obscure ones
close to the centre.
In Pl. 28, fig. 4 _b_, I have given a drawing of two of the cement
glands: the cement-trunk (_f f_) is smooth and apparently cylindrical:
it becomes enlarged (at _g_) before entering the gland: it seems even
to be prolonged across the gland under the form of a narrow bar (not
represented), which apparently serves to keep the two ends of the
trunk, on the two sides of the gland, in their proper relative places
and distances. The gland itself is an elongated bag (_h_), which
properly lies exactly over the enlarged portion (_g_) of the trunk,
but in the drawing has been purposely displaced: it gives rise, in the
later-formed glands, to a sort of neck (see the upper gland), which
is either so long as to deserve rather to be called a duct and which
soon bifurcates, or is quite short (see the lower gland) and gives rise
to two separate ducts. On the opposite side of these glands, there
is a spur (_m_), of greater or shorter length, which is evidently a
rudimentary duct, for in the younger glands it existed as a perfect
duct. Moreover, the first-mentioned duct often gives off branches
(_t′_), having an exactly similar appearance with the spur (_m_). The
membrane of which the cement-trunk (_f_), with the enlargements (_g_),
is composed, is smooth, but that of the glands and of all the ducts,
presents a very peculiar appearance, which at first would be called
scaled, but more properly perhaps notched,--each notch being apparently
formed by a line of thickened membrane, extending obliquely round only
a short portion of the tube, and indenting it. The ducts, which I
measured, were between 1/3000th and 4/3000ths of an inch in diameter.
In fig. 4 _a_, I have given a drawing of the two chains of glands, but
with only those ducts figured which proceeded from the last-formed
pair of glands. The specimen here drawn was old; and it is rare to
find the structure of the ducts so simple. From both glands[72] a neck
or thick duct arises, which soon bifurcates; one branch runs direct
into the circumferential duct, and the other (_t_) bifurcates again;
of the latter, one branch unites with its fellow from the opposite
gland, and then forming a single duct (_t′_) enters, as do the two
other branches, the circumferential duct. Thus, into the latter, five
main ducts enter: the position of their points of entrance, with
respect to the shell, varies considerably; but I think the five points
tend to face the middle of the rostrum, and middle of the two lateral
compartments on each side. In some other specimens, in which the
ducts were nearly as simple, I observed that the neck or main duct at
once divided into three branches, instead of into two, with one soon
bifurcating; and on one side a rudimentary branch or spur was given off
(above _t_), indicating a tendency to an additional bifurcation. In
the later-formed glands, the ducts proceed only from the outer sides
and form the ends of the glands furthest from the centre; but in the
earlier-formed and smaller glands of the same individual, other ducts
proceed from the inner sides, where in the older glands the spurs (_m_)
are situated: moreover, in the younger glands, all the ducts bifurcate
much oftener (how often I was not able to ascertain), before entering
the circumferential duct; many of the branches, however, terminating in
spur-like points. Now if we imagine twenty or thirty repetitions of the
ducts given in fig. 4 _a_, (independently of the greater complication
of the ducts of the younger glands), each a very little smaller than
the other, and placed, with the main branches parallel, one over and
within the other, we shall gain some insight into the wonderfully
complicated structure of the cementing apparatus in this and many other
species of Balanus.
[72] It should be observed that fig. 4 _b_ ought to have been drawn
with its present upper end downwards, to make it correspond in
position with fig. 4 _a_.
I have as yet only alluded to the _circumferential duct_ (_i_, _i_, Pl.
28, fig. 4 _a_): we have not hitherto met with this duct, but I suspect
that the branches which in Chelonobia inosculate, and which seem to
run nearly parallel to the circumference of the basal membrane, answer
the same purpose of connecting the ducts together, and are, perhaps,
strictly homologous. In this, and some other species of Balanus, the
last-formed circumferential duct runs round the margin of the upper
lamina of the basal plate, close to the basal edges of the walls; and
as these latter have projecting longitudinal ribs, the duct curves
a little round each rib; so that the whole duct is formed by as many
short inwardly curved portions as the walls have ribs, or longitudinal
septa. Between the basal extremities of these parietal, longitudinal
septa, the extremities of the radiating septa of the basis project
and enter; and along the crests of the latter, little branch-ducts
(_i′_), proceeding from the circumferential duct, extend. In the basis,
beneath the tubes formed by the just-mentioned radiating septa, there
is a cancellated shelly mass (which, in fig. 4 _a_, was of unusual
thickness), and along the crests of the branching ridges forming this
cancellated mass, the sub-branches of the above branch-ducts (_i′_)
run; these soon become so minute as not to be distinguished by the
highest powers, and thus form a sheet of cement, which attaches the
last-formed zone of the shelly basis to the supporting surface. At
what point the membrane forming any one duct ceases, the cement-tissue
being alone left, I was not able to ascertain; but the lower parts
of the reticulated slip (_z_, _z_, fig. 4 _a_) closely resembled the
cement-tissue which surrounds the disc-segment of the pupal antennæ
in _Lepas australis_. The circumferential duct, here and there, forms
little loops, as may be seen in fig. 4 _a_: and often two branches,
running along the crests of two adjoining basal septa, proceed from
a common point of the circumferential duct. The cement itself, under
different parts of the basis, appears as little separate discs, as
threads, globules, and as a fine network, but most commonly as simple
layers. As each thick zone of shelly matter is added round the basis,
the exterior branches of the ducts, between the circumferential duct
and the new layer of cement beneath, are fairly imbedded in shell, and
are for ever hidden, without, indeed, acid be used for the dissolution
of the calcareous matter: so, also, the pre-existing ducts and glands,
and the main trunk, would all have been hidden, if the layer of
calcareous matter, which, I believe, is thrown down at each period of
growth over the entire surface, had not been of excessive tenuity.
I cursorily examined the cementing apparatus in _Balanus galeatus_,
_improvisus_ and _crenatus_, which have all calcareous bases, and
belong to different sections of the genus; and the structure seemed to
be essentially the same. In _Bal. galeatus_, I found the cement-ducts
varying in diameter from 1/4000th to 1/10,000th of an inch in diameter.
In _B. improvisus_, the cement-glands do not differ much from those
of _B. tintinnabulum_; but the cement-ducts bifurcate often before
entering the circumferential duct; and the little branches, which
proceed from the latter, are very short, and almost immediately, owing
to the thinness of the basis, blend into a slip of cement.
* * * * *
I hope to be excused for describing at such length, the apparatus by
which sessile cirripedes are permanently attached to a supporting
surface; for this is the great leading character of the sub-class,
not hitherto observed in any other Crustacean.[73] It is not easy to
overstate the singularity and complexity of the appearance of the
basal membrane of a Balanus or Coronula: and when we consider the
homological nature of the apparatus, the subject becomes still more
curious: I feel an entire conviction, from what I have repeatedly
seen in several genera of the Lepadidæ, both in their mature and
pupal condition, and from what I have seen in Proteolepas, that the
cement-glands and ducts are continuous with and actually a part of an
ovarian tube, in a modified condition; and that the cellular matter
which, in one part, goes to the formation of ova or new beings, in the
other and modified part, goes to the formation of the cementing tissue.
To conclude with an hypothesis,--those naturalists who believe that
all gaps in the chain of nature would be filled up, if the structure
of every extinct and existing creature were known, will readily admit,
that Cirripedes were once separated by scarcely sensible intervals
from some other, now unknown, Crustaceans. Should these intervening
forms ever be discovered, I imagine they would prove to be Crustaceans,
of not very low rank, with their oviducts opening at or near their
second pair of antennæ, and that their ova escaped, at a period of
exuviation, invested with an adhesive substance or tissue, which served
to cement them, together, probably, with the exuviæ of the parent,
to a supporting surface. In Cirripedes, we may suppose the cementing
apparatus to have been retained; the parent herself, instead of the
exuviæ, being cemented down, whereas the ova have come to escape by a
new and anomalous course.
[73] Rathke has described ('Acta Nova,' 1839, p. 147), in some
siphonostomatous crustaceans, a pair of curious organs, which serve
to secrete a substance that holds the eggs attached together in a
mass to the parent's body: these organs Rathke has designated by
a similar name to that which I have used, namely, the cementing
organs or receptacles; they are distinct from the oviducts, but
enter them near their external orifices. As in Cirripedes, the
cement-glands and ducts are certainly continuous with an ovarian
tube; and as they occupy a quite different position in the animal's
body, these organs of Rathke, though in some degree analogous in
function, must be homologically distinct.
_Affinities, Classification, Variation._
Under the Order it has been stated that the Balanidæ, are, on the
cirripedial type, the highest in the class; that is, they are the
most complicated, but not (to use Professor Owen's term) by mere
vegetative repetition. Amongst the Balanidæ, the first section of the
genus Balanus may be taken as typical; here we have the structure of
the shell extremely complicated, yet beautifully adapted for strength,
and for the protection of the included body. The cementing apparatus
is here, also, most complicated. I have divided the Balanidæ into two
natural sub-families, the Balaninæ and Chthamalinæ, in accordance
with certain differences in the structure of the shell and of the
animal's body: that this division is natural, might almost be inferred
from the one fact, that all the characters by which the Chthamalinæ
differ from the Balaninæ, are those by which the former approaches
the family of the Lepadidæ; moreover, certain anomalous characters
in the Chthamalinæ, as the supplemental whorls of compartments in
Catophragmus, and the presence of caudal appendages in this same genus
and in Pachylasma, reveal this same affinity. The only objection which
I can see to the separation of the sixteen genera into the above
two sub-families, may be drawn from the degree to which they blend
together; thus, as far as the shell is concerned, Chelonobia, in one
important internal point of structure, tends to assume the character
of the Chthamalinæ; and on the other hand, Pachylasma, a member of the
Chthamalinæ, has a shell, which if not examined during its earliest
growth, would be placed without doubt amongst the Balaninæ. But it
fortunately so happens, that in no one character of the body does
Pachylasma approach the Balaninæ more nearly, than do the other members
of its sub-family; or Chelonobia approach, in the same respects, the
Chthamalinæ. It is only in Chthamalus, of which the shell clearly
places it in the sub-family bearing its name, that in some of the
species, the less bullate labrum,--the larger palpi,--the lower teeth
of the mandibles being laterally double,--and the lower segments of
the third pair of cirri being thickly clothed, like the lower segments
of the second pair, with bristles--all show that these species make an
approach in structure to the Balaninæ.
It will be seen that I have divided the Balaninæ into two little
groups, according as whether the branchiæ consist of one or of two
plicated folds of membrane, and as whether or not the scutum and
tergum are articulated together. I have been greatly tempted to
follow Drs. Leach and Gray, who have separated the second of these
groups, containing the genera Coronula, Tubicinella, Xenobalanus,
and Platylepas, into the sub-family of the Coronulinæ. Certainly
these genera have a peculiar aspect in common, and agree in being
parasitic and imbedded in the skin of Cetaceans, as is the case
with the first three genera, or in that of turtles, manatee, and
sea-snakes, as in Platylepas. Though these genera possess several
peculiar characters, yet I can find none common to all four, excepting
their imbedment in the skin of Vertebrata, their double branchiæ, and
their non-articulated opercular valves; and these I do not think of
sufficient importance to serve for the separation of a sub-family; for
in Chthamalus, one species has double branchiæ, one species has no
branchiæ at all, and the other species have single branchiæ; so again
in Chelonobia, the scutum having only a horny articular ridge, makes
an approach to Coronula and its allies. I may further specify that the
folded walls, a singular character common to Coronula, Platylepas and
Xenobalanus, fails in Tubicinella; the open tubes and the imperfect
outer lamina of the parietes towards their bases, are characters which
fail in one species of Platylepas; the muscles running to the opercular
valves being thinly spread out, and partially without transverse striæ,
is also a character which fails in Platylepas; the simplicity of the
cement-ducts partially fails in Tubicinella; and lastly, the existence
of small intermediate teeth on the mandibles, fails in Xenobalanus:
hence, I repeat, I have not thought it prudent to admit the sub-family
of the Coronulinæ though in many respects a very natural group.
The genera in the Balaninæ and Chthamalinæ are founded chiefly on
the number of the compartments (the number being apparently due, as
previously explained, to the fusion or abortion of certain of the
eight typical compartments); and secondarily, on the nature and even
form of the basis, and on the porosity of the walls. In Coronula and
its allies, the non-articulated opercular valves and deeply folded
walls come into play. As a justification for using these characters in
distinguishing the genera, and even to a certain extent in separating
the two sub-families, I must call to mind that the shell, with the
basis, is not merely a dermal envelope, as amongst Molluscs, but
actually consists of the first three segments of the head. The parts of
the mouth and the cirri are of very little service in distinguishing
the genera,--a singular fact, considering that most of the genera
amongst the Lepadidæ could be distinguished by these organs,--though
trifling details in their structure sometimes come in useful as
specific characters. Balanus, with the sub-genus Acasta; Pyrgoma, with
the sub-genus Creusia; Tetraclita, and Elminius, are genera of about
equal value; though perhaps the two latter are rather more nearly
related together than to the others. Chelonobia is more distinct; it
shows some little affinity to the Chthamalinæ, and likewise to the four
following genera. Coronula, Platylepas, Tubicinella, and Xenobalanus,
are genera quite distinct from the foregoing, and from each other;
yet, as we have just seen, palpably allied together. Amongst the
Chthamalinæ, Pachylasma, Octomeris, and Catophragmus, are more closely
related to each other than to the other two genera of the sub-family;
yet Pachylasma, as far as the shell is concerned, leads into the
Balaninæ, and Catophragmus into the Lepadidæ; Octomeris leads towards
Chthamalus, and Chthamalus towards Chamæsipho.
_Variation._--The discrimination of the species in most of the
genera, offers very great difficulties. I cannot too strongly impress
on any one intending to study this class, not to trust to external
characters: he must separate and clean, and carefully examine the
internal structure and form of the compartments, and more especially
of the opercular valves. After considerable experience, when numerous
varieties of a species have been carefully examined, the eye acquires a
sort of instinctive knowledge, by which it can recognise the species,
though the character cannot be defined by language; but I have found
that no amount of experience with some of the commonest species,
will save frequent and grave errors, as long as external characters
alone are trusted to. Not only does every external character vary
greatly in most of the species, but the internal parts very often
vary to a surprising degree; and to add to the difficulty, groups of
specimens not rarely vary in the same manner. After having given up
several years to the study of this class, I must express my deliberate
conviction that it is hopeless to find in any species, _which has a
wide range, and of which numerous specimens from different districts_
are presented for examination, any one part or organ,--which from
differing in the different species is fitted for offering specific
characters,--absolutely invariable in form or structure. I may in one
respect even go further, and affirm, that, if in a species, any part or
organ differs remarkably from the same part in its congeners, then if
many specimens are examined, especially when collected from different
districts, such part or organ will be found _eminently_ variable. I
may instance the antenniformed third pair of cirri in _Chthamalus
antennatus_, the teeth on the posterior cirri in _Acasta sulcata_,
the terga in _Pyrgoma dentatum_, the adductor ridge of the scuta in
_Pyrgoma cancellatum_ and in Creusia, and other such cases: hence it
will not do to found a species on a slight, or sometimes even on a
considerable difference, in _any single point or organ_. On the other
hand, I am far from asserting, that if only half-a-dozen specimens of
some rare species of Cirripede be brought from some one quarter of the
world, characters beautifully defined may not be readily discovered. In
determining what forms to call varieties, I have followed one common
rule; namely, the discovery of such closely-allied, intermediate forms,
that the application of a specific name to any one step in the series,
was obviously impossible; or, when such intermediate forms have not
actually been found, the knowledge that the differences of structure in
question were such as, in _several allied forms_, certainly arose from
variation,--for instance, in the case of two shells otherwise identical,
one being longitudinally ribbed and the other smooth, a character which
we know to vary,--but I have always used this argument from analogy with
great caution. Finally, as in the large genus Balanus, there is an
especial amount of variation, I have there entered in detail on this
subject; and I hope that those interested in it, will refer to that
discussion, which is almost verbally applicable to some other genera of
the family, as Tetraclita and Chthamalus.
_Rate of Growth, Exuviation, Powers of Repairing Injuries._
In my former volume I have shown that the pedunculated cirripedes
grow rapidly; this is likewise the case with the Balanidæ. Mr.
Stutchbury informs me that he has seen numerous specimens of _Balanus
tintinnabulum_ from 2 to 3 inches in height and from 5 to 6 inches
in circumference (and this is nearly the full size which the species
attains), on a vessel which had been to sea only during one year. At
Coquimbo, in Chile, I have seen a specimen of _B. psittacus_ 1.3 of
an inch in basal diameter, and .8 in height, adhering to a chain that
had been only six months under water. Poli, also, gives the case of
some Balani (probably _B. perforatus_), which, in about four months,
had attained a basal diameter of 1 inch, and a height of 1-1/6th of an
inch. _Balanus balanoides_ is a smaller species, and of slower growth;
for the late Mr. W. Thompson, of Belfast, found that in three months
from July 3d, certain marked specimens had increased from 2-1/2-3
lines to 4-1/2 lines, which is the usual maximum size attained in that
locality. From other observations, Mr. Thompson believes that the
extreme duration of life of this species is about two years: whether
the other and apparently quicker-growing species, are shorter-lived, I
have no means of judging.
In accordance with this rapid growth is the frequency of the periods
of exuviation. Mr. Thompson kept twenty specimens of _B. balanoides_
alive, and on the twelfth day he found the twenty-first cast-off
integument, showing that all had moulted once, and one individual
twice, within this period.[74] This frequency of exuviation, together
with the durability of the cast-off integuments, explains the
astonishing masses of exuviæ, which Mr. Peach assures me he annually
has observed off the coast of Cornwall; they are most abundant in
April and May, but he has seen quantities also in September; he could
easily, as he tells me, have filled several quart-measures with them.
The specimens sent to me consisted chiefly of _Balanus balanoides_,
_perforatus_, and _Chthamalus stellatus_. The opercular membrane,
with a narrow strip from between the two scuta, and another narrow
strip from between the two terga, are moulted together, in connection
with the more delicate membrane lining the sack, and investing the
plicated branchiæ. This membrane, likewise, is continuously connected
with that covering the whole body and its appendages. As I have stated
under the Lepadidæ, the inner tunic of the œsophagus, of the rectum,
of the olfactory pouches, and that which enters a little way into the
acoustic meatus, and the apodemes of the maxillæ, are all moulted.
On the cirri and jaws, new spines are formed with their upper ends
enclosed within the old spines, but with their lower ends projecting
inwards, beyond the bases of the old spines, and inverted like the
fingers of a glove hastily pulled off. The membranes of the body, in
the act of exuviation, split, I believe, only over the prosoma. How the
neat separation of the opercular membrane, from all round the sheath
and opercular valves, is effected, I do not fully understand; but it
is, probably, analogous to the splitting of the thick carapace of the
common crab. I suspect in Coronula, in which genus and its allies the
opercular membrane is not periodically moulted, that the membrane
lining the sack is not always thrown off at the same exact time with
that of the body. In _Chthamalus stellatus_, in the act of moulting,
the opercular membrane is the last part that separates from the new
underlying membranes: I find that this species can moult when kept in a
damp box out of water. The new membranes of the body, immediately after
the exuviation, are not lax in any extreme degree. The exuviæ of the
genus Chthamalus, and of some other genera amongst the Chthamalinæ, can
at once be recognised by the divergence of the posterior _four_ pairs
of cirri: in the case of _Chthamalus stellatus_ I have also noticed
that the animal generally dies with these cirri in the same divergent
position. Finally, I cannot doubt[75] that the Triton described by
Linnæus was only the exuviæ of some Balanus (probably _B. porcatus_);
Linnæus mistaking the probosciformed penis for the mouth of his
imagined distinct animal.
[74] In Daphnia, Straus ('Mém. du Museum,' tom. vi, p. 151) found
that the individuals moulted every five or six days.
[75] Linnæus speaks of the included body (inhabitant as he calls
it) of other Cirripedes, as a Triton; and this, I think, shows
that Lesson's conjecture that the Triton was an Alepas cannot be
correct; for Linnæus could hardly have supposed that a pedunculated
cirripede inhabited another pedunculated or sessile cirripede.
I have seen a few specimens showing that when the shell has been broken
it can be repaired; and this I believe is effected by the growth
of a crest of corium between the broken edges, and the subsequent
calcification of this crest. Mr. Stutchbury possesses a monstrous
specimen of _Chelonobia testudinaria_, in which one of the lateral
compartments on one side has not been developed. The cirri not rarely
get cut off, but are, as it appears, soon repaired. I have observed
a singular number of examples of the act of reparation in a group of
the Australian _Balanus vestitus_. The manner in which the cirri are
repaired seems to me curious: the cut-off end is closed by a rounded
scale of yellowish chitine, and then the corium, in the four or five
subjacent segments, separates from the external articulated membrane,
which now serves only as a case or capsule. The tube of corium thus
enclosed, with its contained muscles, shrinks a little, and soon can be
perceived to be in process of dividing into new and smaller segments,
in one instance ten in number,--which at the next exuviation would, no
doubt, be invested with an external membrane, and be freely exposed. In
another instance, the pedicel of a posterior cirrus had been cut off
and subsequently closed; in this instance, a whole, immature, miniature
cirrus, with the two rami, each having fifteen minute segments, was
thus enclosed in what had been the single lower segment of the pedicel.
I have seen several specimens of _Balanus balanoides_, as described
under that species, with several of the cirri and the penis truncated;
but I believe this was owing to monstrosity, which seemed particularly
to affect the male organs of generation; for no reparation seemed to be
in progress. In a specimen of Coronula, however, the penis appeared to
have been really cut off by accident; it had been closed, by a scab,
with concentric lines, like the articular rings on the penis itself;
and within the case thus formed, the corium had healed, and had become
pointed, but _inverted_; I presume that the point would, after another
exuviation, have been everted, and its length thus increased.
_Geographical Range and Habits._
With respect to range, the results arrived at have no particular
interest, for the species are not sufficiently numerous; and, what is
still more adverse, the genera, with unimportant exceptions, range over
the world; so that there is no scale of differences, and it cannot
be said that these two regions differ in their genera, and these two
only in their species. In all the following remarks, I have trusted
exclusively to my own specific identification; and have rejected all
assigned localities which appeared from any cause to be doubtful.
Sessile cirripedes are found in every sea, from lat. 74° 18′ north to
Cape Horn. The area included between the north point of the Philippine
Archipelago and the south point of Australia, extending on the right
hand to New Zealand, and on the left to Sumatra,--an area, which,
though including two distinct Cirripedial regions, is small compared
with the surface of the globe, yet includes a greater number of
species, especially of peculiar species, than the whole rest of the
known world. This is, probably, in chief part due to the broken nature
of the land, affording diversified habitats, and to much of the coast
being rocky. Cirripedes, from requiring to be attached, cannot live
where the shores and adjoining bottom are sandy or muddy or formed
of moving shingle; hence, no doubt, it arises, that there is such a
remarkable contrast in the great number of the species inhabiting
the bold rocky western coast of South America, and the few species
living on the sloping, and often sandy or muddy or shingly, eastern
shores of this continent. Hence, also, I believe, it is that not many
species have been brought from India. Coral-reefs are not favorable to
Cirripedes, consequently but few are known to inhabit the islands of
the Pacific Ocean. Where Cirripedes can live, though the species in no
district are numerous, the individuals abound in infinite numbers: I
have walked over the coast-rocks of the Falkland and Chonos Islands,
of Chile and Van Diemen's Land, fairly encrusted over wide spaces with
a continuous layer of Cirripedes, consisting of only two or three
species; in the same manner as may be observed on many parts of the
shores of Great Britain, and, I believe, of North America.
With respect to the effects of temperature on the range of Cirripedes,
no genus (having more than one species) is confined to the torrid
zones. Pyrgoma, from being always attached to corals, is, of course,
ordinarily found in the hotter seas; but one species ranges from the
Cape de Verde Islands in the torrid zone to the southern shores of
England and Ireland. Tetraclita is not found in the colder seas, but
is numerous in species and in individuals, on the southern shores of
Australia and at the Cape of Good Hope. I may here add, that the two
genera with the most confined ranges, are Chamæsipho and Elminius;
the former has only two species, one inhabiting Australia, and the
other the East Indian Archipelago; Elminius has four species, confined
to the southern hemisphere, and inhabiting Australia, New Zealand,
and South America. To return to the effects of temperature; in Mr.
Dana's great work on Crustacea, an excellent chart is given, in which
the _isocrymal_ lines, or those exhibiting the mean temperature of
the waters along their course, for the coldest thirty consecutive
days in any season of the year, are given; and which lines Mr. Dana
has shown are the most influential in governing the distribution of
marine animals. At the isocryme of 68°, Mr. Dana divides the torrid
and sub-torrid zones from the several temperate zones; and at 44°, the
temperate from the sub-frigid and frigid zones; but as no Cirripedes
are exclusively confined to these frigid zones, we may here disregard
them. From Mr. Dana's[76] table of the areas of the torrid and
temperate ocean-zones, on both sides of the equator, it seems that
they are nearly as 337 to 278, in relative area; and consequently, he
remarks, that the marine species in any class, if distributed equally
over the two, would be one fifth more numerous in the torrid than in
the temperate zones. Now of Cirripedes, taking all the orders, there
are at present known 147 species; of these, 7 have doubtful habitats,
leaving 140 for comparison. Of these 140, nearly one quarter, or
37, inhabit both the torrid and temperate zones, as above defined;
46 are found exclusively in the torrid, and 57 exclusively in the
temperate zones; so that the temperate zones, though less in area,
and having, proportionally, even a considerably lesser length of
coastline, nevertheless have a preponderance in the number of species.
But it should be borne in mind, that there are _two_ great temperate
districts, separated from each other by _one_ great torrid district;
and, inasmuch as the number of species in any region seems to depend in
some degree on the isolation of the sub-regions, we might have expected
(the other conditions now being, and the _past_ conditions having been
alike), that the two great temperate areas would have contained more
species, perhaps doubly more, than the single great torrid area.
[76] 'Crustacea: United States Exploring Expedition,' p. 1476
(corrected).
The proportional numbers, above given, are not very widely different,
whether we take separately the Balanidæ, the Lepadidæ, or all
together. Mr. Dana has shown[77] at length, that generally amongst
the Crustacea, the species which he considers of highest rank, belong
to the extra-torrid zones: there seems to me in all such cases to
be some degree of vagueness in the attempt to determine which are
highest or lowest, but I have already elsewhere stated that Balanus
is, probably, the most eminently Cirripedial form, and exhibits in the
strongest manner all the characters by which Cirripedes differ from
other Crustacea; as this genus is the largest, containing 36 species,
of which the habitats are known, I may state that of these, exactly one
third, or 12, inhabit both zones; 9 exclusively inhabiting the torrid,
and 15 exclusively the temperate zones. According to the proportions of
the whole class, the numbers should have been 9 torrid, to only 11.11
temperate; so that evidently the genus Balanus (in one sense typical)
inclines towards the temperate regions more strongly than does either
the family or the sub-class to which it belongs.
[77] Ibid., p. 1528.
With respect to the relation between the size acquired by the different
species of sessile cirripedes, and the temperature of the localities
inhabited by them, the genera Chthamalus, Tetraclita, and Balanus,
alone can serve for comparison: in Chthamalus much the largest species
is found in the temperate zone: on the other hand, the two largest
species of Tetraclita are from the torrid zone, though one of them
also sometimes ranges into the temperate seas: in Balanus, the largest
species, and six other species having a basal diameter sometimes over
two inches, inhabit the temperate regions; and two out of these seven
species, flourish even in the Arctic seas; whereas, within the torrid
zone, there are only three species with a diameter sometimes exceeding
two inches, but two of these frequently become very large and massive;
so that Balanus, judging from the size of the species, as well as
from their range, does not require for its highest development the
temperature of the torrid zones.
The greater number of the species of the Balanidæ have wide ranges, as
might be inferred from the fact of between one third and one fourth
of the total number inhabiting both the torrid and temperate zones;
but it should not be overlooked, that those species, as _Balanus
tintinnabulum_, _amphitrite_, _improvisus_, and, in a lesser degree,
_B. trigonus_ and _Tetraclita radiata_, which seem to range over
nearly the whole world (excepting the colder seas), are species which
are habitually attached to ships, and which could hardly fail to be
widely transported. Indeed, it appears to me surprising, that such
species as _Balanus psittacus_ and _eburneus_, which often become
attached to vessels, should still be confined, the one to Southern,
and the other to Northern America. But some other Cirripedes, which
I have never seen attached to vessels, have likewise immense ranges:
thus _Tetraclita porosa_ is found in every tropical and warmer sea,
and _Chthamalus stellatus_ ranges round the world in the northern
hemisphere, and, along the eastern side of America, far south of the
equator: _Balanus spongicola_, and _Acasta spongites_, extend from the
shores of Britain to the Cape of Good Hope: _Balanus lævis_ ranges from
Tierra del Fuego to California. I may further remark, that the only two
other species of Balanus, and the one Chthamalus, inhabiting Tierra
del Fuego, are, also, found on the shores of Peru. But to show the
powers of endurance in some species, I may specify the case of _Balanus
improvisus_, which flourishes on the coast of Nova Scotia, amongst
the West Indian Islands, in Southern Patagonia, and near Guayaquil.
Even more striking is the case of _B. crenatus_, of which I have seen
specimens from latitude 74° 48′ north, from the West Indies, and the
Cape of Good Hope! In these two latter localities, however, it seems to
be rare, and may have been first transported to them from the shores of
Europe, on the bottoms of vessels, to which it sometimes adheres.
The several species of Balanidæ live attached either to coast-rocks,
or to objects at various depths, down to, as in the case of _Balanus
crenatus_, 50 fathoms. _Balanus balanoides_ sometimes adheres to rocks
or wood so high up, that it is not covered by water during the neap
tides. Mr. Thompson has informed me, that he once accidentally kept
some specimens of this species out of water for seven days in a warm
room, and that they were then quite lively. This species, is very
easily killed by brackish water, as are some other species, whilst _B.
improvisus_ and _eburneus_ can flourish in it; and at the Falkland
Islands, I saw _Elminius Kingii_ attached to rocks at the mouth of
a fresh-water brook, so as to be covered by pure water during the
ebb of each tide. Sessile cirripedes adhere to all sorts of objects,
floating and fixed, animal and vegetable, living and dead, organic and
inorganic. Chthamalus is, perhaps, more commonly attached to rocks than
are the other genera. Living Mollusca are, I think, the most frequent
objects of attachment: Mr. Cuming has remarked to me, that shells
covered by an epidermis, as Patella, Haliotis, and Mytilus, are the
greatest favorites. Acasta is always imbedded in sponges, or in the
sponge-like bark of Isis; Pyrgoma and Creusia in corals; Chelonobia is
attached to turtles, and one species to crabs or very smooth shells;
Coronula, Tubicinella, and Xenobalanus, are imbedded in the skin of
Cetaceans; and Platylepas in that of manatee, turtles, or sea-snakes.
If we attempt, with our present not very imperfect materials, to
divide the globe into provinces, according to the amount of difference
in their Cirripedial inhabitants, including all orders and families,
and disregarding entirely, as I think we ought, all probabilities or
conclusions deduced from the distribution of other tribes of animals,
we find that the globe may be divided into the four following great
provinces and one sub-province. I should premise, that in the following
remarks and tables,[78] the species of Lepas, Conchoderma, Chelonobia,
Coronula, Platylepas, and Tubicinella, are excluded, owing to their
being attached to floating or swimming objects, and being consequently
widely and irregularly distributed.
[78] As the number of Cirripedes in the whole class is not very
great, I have given lists of the species in the four main provinces
and in the one sub-province.
The first, or North Atlantic province, is that of Europe and the
eastern shores of North America, from the arctic regions to lat. 30°:
the island of Madeira, part of the north-west coast of Africa, and the
whole Mediterranean being included. In this province (the above-named
genera being excluded) we have 31 species, of which 22 are not found in
any other distant quarter of the globe. As some few of these species
range into the West Indies, I have not, on this account, excluded them
from the 22 peculiar forms. Had I included the West Indies[79] in my
first province, the total number of species would have been 42, of
which 28 would have been peculiar. The coast of Brazil, even as far
south as the Rio Plata, might, also, have been included, for I have not
seen from it a single species not included in the above 42 West Indian
species. So also, by adding a single species, might the west coast of
equatorial Africa. The two coasts of South America and Africa, which
face each other within the torrid zone, seem to be remarkably barren
in Cirripedes. Europe has several more species than the United States,
which is inhabited by only ten species, including even the probably
imported _Balanus tintinnabulum_ and _amphitrite_. Of these ten United
States species only two are not found in Europe; and both these two
range into the West Indies, and as far as the northern shores of South
America, and therefore cannot be considered as peculiar to the United
States.
[79] The total number of species which I have seen from the West
Indies, is 19 or 20; of these, only 6 are peculiar to it, or 8, if
the United States be likewise included, the other 12 or 14 species
being found in other quarters of the world. Six peculiar species
out of 19 or 20, has not appeared to me sufficient to institute
even a sub-province.
I have formed my single sub-province for the southern extremity of
Africa; for although I know of only 11 species from this comparatively
short and uniform line of coast, yet I was not able to group these
eleven in any of the main provinces: 5 of the species are peculiar,
1 Australian, 3 European and West Indian, and 2 almost universally
distributed.
The second province includes the west coast of North and South America,
from Tierra del Fuego to Behring's Straits: on this enormously long
line of coast, only 22 species are known to exist, but of these no
less than 15 are peculiar. Of these 15, four are not found south of
the torrid equatorial region, and eight are not known to occur north
of this same region; so that this long line of coast might have been
divided into two sub-provinces, of which the southern would have been
the most peculiar; but as eight species are found both north and south
of the equatorial region, I have not made this sub-division. Two of
the species occurring on the western coast of North America, seem to
represent species found on its eastern coast, and in Europe; thus,
_Balanus glandula_ takes the places of _B. crenatus_, and _B. cariosus_
that of _B. balanoides_. Not a single species, excepting a few which
are also widely distributed over other parts of the world, is known to
be common to the east and west coasts of the two Americas.
The third province is that of the East Indian Archipelago, and includes
the Philippines, Borneo, New Guinea, Sumatra, Java, Malacca, and the
eastern coast of India. Here we find 37 species, of which 24 are
peculiar. I may remark, that I have received no species from Madagascar
or the eastern coast of Africa; few from India, or from the coast of
China; and I suspect, that on most of these coasts, only few exist.
Probably our third province will hereafter be found to include the
whole Indian Ocean.
The fourth province is that of Australia, including New Zealand: it
has 30 species, of which 21 are peculiar. Had the temperate Australian
coasts (_i. e._, those south of the isocryme of 68°) been alone
considered, the number of the species would have been probably 25, of
which 20 would have been peculiar,--that is, if we admit within the
20, several species which range from the temperate into the torrid
zone, but do not extend beyond the Australian shores. Owing to the
widely-extended ranges of most Cirripedes, no Arctic or Antarctic
provinces can be said to exist.
To recapitulate the above results, bearing in mind that, although the
total number of known existing Cirripedes is 147, yet that the habitats
of seven are unknown, and that eighteen are excluded owing to their
being attached to floating or swimming objects, so that there are only
122 species referred to in the following table:
+--------------------------------------------------+--------+---------+
| | Total | Species |
| | number |confined |
| | of | to the |
| |species.|province.|
+--------------------------------------------------+--------+---------+
|(1.) First, or North Atlantic Province, to lat. }| | |
| 30° N. (If the West Indies had been included}| 31 | 22 |
| the numbers would have been 42 and 28) }| | |
|(2.) Sub-province of South Africa | 11 | 5 |
|(3.) Second province, or West Coast of North and }| | |
| South America }| 22 | 15 |
|(4.) Third province, or East Indian Archipelago | 37 | 24 |
|(5.) Fourth, or Australian province | 30 | 21 |
+--------------------------------------------------+--------+---------+
The least prolific of these provinces contains 22 species, or between
1/5th and 1/6th of the total number of species, and the most prolific
between 1/3rd and 1/4th of this same number. In each of these
provinces, it is remarkable that the peculiar species are very nearly
two thirds of the whole of its inhabitants. These facts, I think, show
that the above provinces are natural divisions of the world, as far as
their Cirripedial inhabitants are concerned.
As Cirripedes belong to the great class of Crustacea, and as the
distribution of the latter has lately been fully discussed by Mr. Dana,
it may be worth while briefly to compare my results with his; more
especially as they are so very different. I should premise, as perhaps
accounting to a certain extent for this difference, that, owing to the
wide range of many species, and the almost universal extension of the
same genera, my provinces are founded merely on a certain proportion
of the species, namely, two thirds, being peculiar or confined to a
region of considerable dimensions: whereas, in the case of ordinary
Crustaceans, the greater number of the species are distinct even in
the sub-provinces, and the provinces are founded mainly on generic
differences. Mr. Dana divides the surface of the globe into three great
sections, or provinces, the _Africo-Europæan_, the extent of which is
shown by its title; the _Occidental_, which includes both the east and
west coast of both Americas; and the _Oriental_, including the Indian
and Pacific Oceans, with the East Indian Archipelago, and Australia.
Thus Mr. Dana entirely separates the Eastern shores of North America
from Europe; whereas, as far as their Cirripedial inhabitants are
concerned, they are most intimately allied, and form my first or North
Atlantic province; and to this, as I have shown, even the West Indies,
the coast of Brazil, and equatorial West Africa might have been added.
It follows, from this similarity in the Cirripedes on the two sides of
the Atlantic, and from their dissimilarity with those on the shores of
the Pacific, that the east and west coasts of the two Americas form two
quite distinct Cirripedial provinces; though, in the northern half,
some connection is shown by a few representative species: on the other
hand, Mr. Dana unites both sides of the whole American continent,
into his single Occidental province. The South-African province is not
brought out by Mr. Dana so prominently, as I have found necessary. Mr.
Dana joins the East Indian Archipelago and Australia into his single
Oriental province, and makes New Zealand, as a sub-province, apparently
as distinct from Australia, as Australia is from the East Indian
Archipelago: whereas I find that the Cirripedes of New Zealand clearly
belong to Australia; and that the Australian Cirripedes, especially if
the temperate shores be alone considered, are as distinct from those of
the East Indian Archipelago, as from those of any other quarter of the
whole world. I believe that the provinces deduced from the distribution
of Cirripedes, accord better with the Molluscan provinces, than with
those given by Mr. Dana for the rest of the great class of Crustaceans.
In the following tables, an asterisk means that the species is not
found in any other distinct region of the globe. When found in one
of the five provinces, a corresponding number, within brackets, is
appended, to show in which province or sub-province it has been found.
(1.) FIRST OR NORTH ATLANTIC PROVINCE: _Europe and the Eastern United
States, from the Arctic Regions to 30° north latitude._
Balanus tintinnabulum (1 to 5).
tulipiformis* confined to Europe.
calceolus Europe and India.
{ confined to North America and West
galeatus* { Indies.
spongicola (2) and West Indies (?).
{ confined to Europe, but possibly in the
perforatus* { West Indies.
amphitrite (1 to 5).
eburneus* { confined to North America and West
{ Indies.
improvisus Europe and North America and (3).
porcatus* Europe and North America.
crenatus " " W. Indies and (2).
balanoides* " "
Hameri* " "
Acasta spongites Europe and (2).
cyathus* Madeira and West Indies.
{ confined to Europe, but ranges as far at
Pyrgoma anglicum* { least as the Cape de Verde Islands.
Xenobalanus globicipitis* confined to Europe.
Chthamalus stellatus Europe and North America and (3 and 4).
Pachylasma giganteum* confined to Europe.
Verruca Strömia Europe and Red Sea.
Spengleri* Madeira.
Pœcilasma aurantia* "
crassa* "
Dichelaspis Lowei* "
Oxynaspis celata* "
Alepas minuta* Europe.
parasita* Europe and Atlantic Ocean.
Anelasma squalicola* Europe.
Alcippe lampas* "
Scalpellum vulgare* "
Pollicipes cornucopia* "
Here we have 31 species, of which 22 are not found in any other great
region of the world.
(2.) SUB-PROVINCE: _Africa, South of lat. 30°._
Balanus tintinnabulum (1 to 5).
Capensis*
spongicola (1) and West Indies (?).
amphitrite (1 to 5).
crenatus (1) and West Indies.
Acasta spongites (1).
Tetraclita serrata*
rosea (4).
Chthamalus dentatus* also on West Coast of Africa.
Octomeris angulosa*
Scalpellum ornatum*
In this small region we have only 11 species, of which five are
peculiar: _Balanus Capensis_ and _Tetraclita serrata_, seem to be
representatives of _B. psittacus_ of S. America and of _T. porosa_ of
that and several other regions.
(3.) SECOND PROVINCE: _West Coast of South and North America, from
Tierra del Fuego to Behring's Straits._
Balanus tintinnabulum north and south of the equator (1 to 5).
psittacus* south.
vinaceus* "
trigonus north and south (4 and 5).
lævis* "
concavus " (4 and 5).
pœcilus* south.
improvisus south (and north?) (1).
nubilus* north.
glandula north, and Southern Pacific Ocean.
cariosus* north.
Tetraclita porosa north and south (4 and 5 and W. Indies).
Elminius Kingii* south.
Chthamalus stellatus north.
cirratus* south.
scabrosus* "
fissus* north and south.
Hembeli* north.
Verruca lævigata* south.
Pollicipes elegans* south and north.
polymerus* "
Cryptophialus minutus* south.
Here we have on this long line of coast, 22 species, of which 15 are
peculiar.
(4.) THIRD PROVINCE: _Indian Archipelago (including the Philippines,
Malacca, Borneo, Sumatra, Java, and New Guinea, and eastern coast
of India)._
Balanus tintinnabulum (1 to 5).
Ajax*
navicula*
stultus and West Indies.
trigonus (3 and 5).
concavus (3 and 5).
amphitrite (1 to 5).
patellaris*
amaryllis (5).
quadrivittatus*
Acasta lævigata and Red Sea.
fenestrata*
purpurata*
sporillus*
Tetraclita porosa (3 and 5) and West Indies.
costata*
vitiata (5).
cœrulescens Pacific Ocean.
radiata (5) and West Indies.
Pyrgoma cancellatum*
grande*
milleporæ*
crenatum*
monticulariæ*
Creusia spinulosa and West Indies.
Chthamalus stellatus (1 and 3).
intertextus*
Chamæsipho scutelliformis*
Octomeris brunnea
Pœcilasma fissa*
eburnea*
Dichelaspis Warwickii*
Ibla Cumingii*
Scalpellum rostratum*
Pollicipes mitella*
Lithotrya Nicobarica*
truncata and Pacific Ocean.
Here we have 37 species, of which 24 are peculiar to this province.
(5.) FOURTH PROVINCE: _Australia (including New Zealand)._
Balanus tintinnabulum (1 to 5).
nigrescens*
decorus*
trigonus (3 and 4).
concavus (3 and 4).
amphitrite (1 to 5).
amaryllis (4).
allium*
vestitus*
imperator*
Acasta sulcata*
glans*
Tetraclita porosa (3 and 4).
rosea (2).
purpurascens*
vitiata (4).
radiata (4) and West Indies.
Elminius plicatus*
simplex*
modestus*
Chthamalus antennatus*
Chamæsipho columna* and Pacific Ocean (?).
Pachylasma aurantiacum*
Catophragmus polymerus*
Alepas tubulosa*
Ibla quadrivalvis*
Scalpellum Peronii*
Pollicipes spinosus*
sertus*
Lithotrya cauta*
Here we have 30 species, of which 21 are peculiar.
_Geological History._
The ancient history of the Balanidæ is a brief one. No secondary
species has hitherto been discovered; in my monograph on the fossil
Lepadidæ[80] I have shown that the negative evidence in this case is of
considerable value, and consequently that there is much reason to doubt
whether any member of the family did exist before the Eocene period.
The existence of a Cretaceous Verruca is an apparent exception to the
rule, as this genus has hitherto always been ranked amongst sessile
cirripedes; but Verruca, as we now know, must be placed in a family by
itself, quite distinct from the Balanidæ. Balanus is the oldest genus
as yet known; it first appeared in Europe and North America, during
the deposition of the eocene beds; and was at that time, as far as our
information at present serves, represented by very few species. In
South America, one species of Balanus abounds in individuals in the
ancient Patagonian tertiary formation. I have seen, in the British
Museum, specimens, said to have come from the eocene nummulitic beds
near the mouth of the Indus, belonging to the second section of the
genus. Generally, the extinct forms belong to the last section of
this genus, which has the parietes not permeated by pores. During the
miocene and pliocene ages, sessile cirripedes abounded. No extinct
genus in this family has hitherto been discovered. It is singular,
that though the Chthamalinæ approach much more closely than do the
Balaninæ to the ancient Lepadidæ, of which so many species have been
found fossil even in the older Secondary formations, yet that only one
species of one genus of this sub-family has been hitherto found in any
deposit; and that species is the still existing _Pachylasma giganteum_,
in the modern beds of Sicily. During the epoch of the Glacial deposits
in Scandinavia, Scotland, and Canada, the still existing species seem
to have abounded; and they attained larger average dimensions than
the same species now do on the shores of Great Britain, or even on the
shores of the northern United States, where the average size seems
larger than on this side of the Atlantic.
[80] Since the note to page 5 of that work was written, I have been
informed that the so-called cretaceous _Tubicinella maxima_ is not
a Cirripede.
Under the genus Balanus, I have given my reasons for never naming
species in this large and difficult genus, without examining the
opercular valves: it has been owing to this, as it appears to me,
proper want of caution, that there are so many nominal species. Thus
it is made to appear in catalogues, that the tertiary seas abounded
with species of Balanus to an extent now quite unparalleled in any
quarter of the world. Bronn,[81] for instance, in his invaluable
'Index Palæontologicus,' gives the names of 35 Balani, found fossil in
Europe, and I have not counted those found only in alluvial deposits,
as they would certainly be the same as the still living species. Now
I know only 11 recent Balani on the shores of all Europe, from the
North Pole to lat. 30°; and of these I doubt whether _B. balanoides_
and _improvisus_ have been found fossil. In the Red Crag there is one
extinct Balanus: in the Coralline Crag, which seems to have been very
favorable to the existence of Cirripedes, there are six species of
Balani, of which two are absolutely extinct, and one does not occur
in any neighbouring sea: in the Eocene formations the species seem to
have been rare, and I have seen only one, and that is an extinct form.
Taking these several facts into consideration, and bearing in mind
that Cirripedes usually range widely, I do not believe, if all the
specimens of Balani hitherto found in the several tertiary formations,
from the eocene to the glacial deposits, throughout Europe, were
collected together, they would amount to 20 species. I have myself
seen, in a recognisable state, only 12 fossil species, of which five
are extinct, or not found in any neighbouring sea: I think it probable
that three other recent species, viz. _B. tulipiformis_, _perforatus_,
and _amphitrite_, may occur in the Mediterranean formations; and this
would make 15 species. Therefore in the above estimate of 20 species,
five are allowed for species existing in European collections,
but not hitherto seen by me; and this, I believe, is a very full
allowance. Consequently, even on the supposition that the five
species just admitted as possibly existing in cabinets, and that the
other five extinct species, which I have seen and examined, have all
been previously named by other authors, a supposition excessively
improbable, even then there would be 15 superfluous names in Bronn.
[81] To save any other person, interested in fossil Cirripedia,
going through the several works quoted by Bronn, I have given some
remarks on his list of species, in an appendix at the end of the
Balanidæ.
The following short table shows how Cirripedes, including all three
Families, were represented in Great Britain, throughout the several
TERTIARY STAGES.
A = Living species but found fossil in some tertiary deposit.
B = Mammilliferous crag, and glacial deposits.
C = Red crag.
D = Coralline crag.
E = Eocene.
+----------------------------+---+---+---+---+---+
| Name. | A | B | C | D | E |
+----------------------------+---+---+---+---+---+
| Balanus tintinnabulum | * | | * | | |
| calceolus | * | | | * | |
| spongicola | * | | | * | |
| concavus | * | | * | * | |
| porcatus | * | * | * | | |
| crenatus | * | * | * | * | |
| Hameri | * | * | * | | |
| bisulcatus | | | * | * | |
| dolosus | | * | | | |
| inclusus | | | | * | |
| unguiformis | | | | | * |
| Acasta undulata | | | | * | |
| Pyrgoma anglicum | * | | | * | |
| Coronula barbara | | | * | | |
| Verruca Strömia | * | * | * | * | |
| Scalpellum magnum | | | | * | |
| quadratum | | | | | * |
| Pollicipes reflexus | | | | | * |
+----------------------------+---+---+---+---+---+
| Total, 18, recent and } | | | | | |
| extinct, found fossil in } | 9 | 5 | 8 |10 | 3 |
| Great Britain, in some } | | | | | |
| tertiary deposit } | | | | | |
+----------------------------+---+---+---+---+---+
As affording some standard of comparison by which to compare the number
of fossil species, at any period, in relation to the number of species
probably existing in the neighbouring seas during the same epoch, I
may state that there are now living and propagating on the shores of
Great Britain, 18 species belonging to the three Families included
in the above table. I have not counted three species, in the genera
Alcippe and Conchoderma, which, from the minuteness of their valves,
it is hardly possible would be found fossil. On the other hand, I have
included in the 18, five species of Lepas, which from floating and
being oceanic, are more likely to be cast up on beaches, than to be
imbedded in sedimentary deposits; so that 13 would, perhaps, be a safer
number, as a standard of comparison. Now in the coralline crag, which
seems to have been eminently favorable for the existence and subsequent
preservation of Cirripedes, and which has been so well worked, only
nine fossil species, as may be seen in the table, have been as yet
discovered.
Sub-Family--BALANINÆ.
_Shell with the rostrum having radii, but without alæ; lateral
compartments all having alæ on one side and radii on the other side;
parietes generally either porose, or longitudinally ribbed on their
inner surfaces._
_Mouth with the labrum notched in the middle, not swollen; palpi large,
almost touching each other; mandibles generally with the lower teeth
laterally double; third pair of cirri with their segments resembling
those of the second pair._
_First Section._†
_Scutum and tergum articulated together, or overlapping each other;
each branchia composed of a single plicated fold._
_Genera_--Balanus; Acasta; Elminius; Tetraclita; Pyrgoma; Creusia;
Chelonobia.
_Second Section._††
_Scutum and tergum (when both are present) not overlapping each other;
basis membranous; parietes often deeply folded, with the outer lamina,
towards the basis, generally imperfect; each branchia composed of two
plicated folds; shell attached to living vertebrata._
_Genera_--Coronula; Platylepas; Tubicinella; Xenobalanus.[82]
[82] At the end of the volume a Synopsis is given, which will serve
as a systematic index for the discovery of generic and specific
names.
The Balanidæ may be divided into two sub-families; namely, the Balaninæ
and Chthamalinæ; and, in the former, the genera, as we see, may be
very naturally grouped into two sections. The Balaninæ differ from the
Chthamalinæ, as far as the shell is concerned, in the rostrum having
radii but no alæ, all the lateral compartments having both radii and
alæ; on the other hand, in the Chthamalinæ, the rostrum has alæ, and
the rostro-lateral compartments radii on both sides, and therefore
no alæ. These differences probably arise, as already explained, from
the perfect confluence, in the Balaninæ, of the true rostrum with the
rostro-lateral compartments. In Chelonobia, belonging to the Balaninæ,
we see an intermediate state, with the fusion not quite effected:
on the other hand, in one genus amongst the Chthamalinæ, namely,
Pachylasma, we must look to the shell at a very early age, to find the
rostrum with its alæ, distinct from the rostro-lateral compartments.
In Tetraclita, Elminius, and Creusia, the carino-lateral compartments
are aborted, or possibly confluent with the lateral compartments,
making altogether only four: in Pyrgoma all the compartments are
fused together and form a solid ring. The sub-genus Acasta is, in
one sense, very natural, as it includes species most closely allied:
in another sense it is far from natural, as some of the species can
hardly be distinguished from those species of Balanus, which live
attached to Gorgoniæ: I almost regret I did not merge the species of
Acasta into Balanus. In the Balaninæ generally the parietes are either
porose, or are furnished on their internal surfaces with regular ribs,
representing the longitudinal parietal septa, which in other species
form the tubes or pores; there are, however, many exceptions to this
rule in several species of Balanus, in Acasta and Elminius, all of
which have the parietes of their shells internally quite smooth, or
only irregularly roughened with points.
Looking to the animal's body, in the Balaninæ, the labrum is always
notched in the middle, and is never swollen or bullate, for the outer
and inner folds of membrane of which it is composed lie close together.
The palpi are large, so that their tips almost touch each other. The
mandibles, generally, have their lower main teeth laterally double.
Of the cirri, the third pair invariably much more closely resembles,
in its whole structure, and in its action, the second than the fourth
pairs; and it is also generally separated by a small interval from the
fourth pair.
I have already under the Family sufficiently entered on the relations
of the Balaninæ to the Chthamalinæ, and of the genera, one to the
other, so that I need not here add anything.
I can point out no difference in habits or geographical distribution
between the Balaninæ and Chthamalinæ.
1. _Genus_--BALANUS, Auct.[83]
[83] The name Balanus was used, almost as at present, by Lister and
Hill, before the introduction of the binomial system. Since that
period the first two authors, as far as I know, who used this name,
were Da Costa, in his 'Hist. Nat. Test. Brit.,' in 1778; and Bock,
in the 'Naturforscher,' for the same year; Bock, however, applied
it to a Chelonobia.
CONOPEA (pars generis). _Say._ Journal Nat. Sc. Philadelphia, vol.
ii, part ii, 1822.
MESSULA (do.). _Leach._ Zoological Journal, vol. ii, 1825.
CHIRONA (do.). _J. E. Gray._ Philosoph. Transacts., 1835, p. 37.
_Compartments six; basis calcareous or membranous; opercular valves
sub-triangular._
_Distribution._--Mundane: in the warmer seas.
_General appearance._--The shape of the shell in the different
species varies from depressed conical to cylindrical; the latter
form being generally assumed when specimens are crowded together; but
some species, as _B. balanoides_, _crenatus_, and _lævis_, seem more
subject than others to be thus affected. The colour is either white,
generally tinted by the yellowish or brownish epidermis, or any colour
intermediate between bright pink and rich blue, purple being the
prevailing tint. The persistence of the so-called epidermis is very
different in different species, being even sometimes highly variable in
the same species. The surface is either smooth or more commonly folded
longitudinally, or sharply ribbed. The orifice differs in form from
diamond-shape to trigonal; the carinal end, owing to the shape of the
carina, being always sharper or narrower than the rostral end. The size
of the orifice, in proportion to the shell, varies accordingly as the
latter is more or less conical or cylindrical. The orifice is either
entire or more or less deeply toothed, in proportion to the degree of
obliquity of the summits of the radii and alæ. The radii almost always
have smoother surfaces than the parietes. In some few species the
radii are not developed, the sutures being marked only by fissure-like
lines; in others they are very narrow, and in this case their upper
margins are generally rounded and smooth, instead of being straight
and jagged. The carino-lateral compartments are usually much narrower
than the lateral compartments, occasionally in an extreme degree, as in
_B. allium_. The shell is generally strong, sometimes to a wonderful
degree; but the strength and thickness vary in the individuals of some
of the species. By the action of hot caustic potash, the compartments
in several species, such as _B. Hameri_ and _crenatus_, separate on a
touch; in others, they adhere so strongly as to prove that the sutures
must be calcified together. In this genus we have the largest known
sessile cirripede, viz., the _B. psittacus_, and on the other hand many
small species; but it is very difficult, except in well-known species,
to ascertain the average or even the maximum dimensions.
_Scutum._--This valve is almost triangular, with the basi-tergal corner
more or less rounded off. The prominent lines of growth are sometimes
crossed by longitudinal striæ. Internally, the articular ridge projects
to a very different degree in the different species; its lower end is
sometimes (as in _B. lævis_, Pl. 4, fig. 2 _c_) produced downwards as a
small, sharp, free style; there is always an articular furrow receiving
the inflected margin of the tergum. There is always an impression left
by the attachment of the adductor scutorum muscle; and often its lower
side is bounded more or less closely by a sharp adductor ridge, running
some way down the valve; this ridge is occasionally almost confluent,
in its upper part, with the articular ridge, and in this case sometimes
it forms, together with the inflected tergal margin, a large tubular
cavity, running up, as in _B. psittacus_ (Pl. 2, fig. 3 _c_), almost
to the apex of the valve. Almost invariably there is a slight pit or
depression for the lateral depressor muscle; sometimes within the
depression there is a little ridge, as in _B. perforatus_ and _nubilus_
(Pl. 4, fig. 3 _a_, and Pl. 6, fig. 2 _a_); and in the case of _B.
vestitus_, _flosculus_, and _imperator_ (Pl. 8, figs. 3 _a_, 4 _a_),
there are regular crests for this same purpose. The rostral depressor
muscle is usually attached in a small pit formed by the folding over of
the lower part of the occludent margin: in _B. imperator_ (Pl. 8, fig.
4 _a_) there are regular crests for its attachment, and traces of them
may be discovered in _B. vestitus_.
_Tergum._--This valve is more nearly triangular than any other shape,
with the spur more or less prominent. The apex generally projects a
little above the level of the scutum; in some species it consists of
a triangular and solid, in others (Pl. 2, fig. 3 _b_) of an almost
cylindrical, extremely sharp, inwardly curved, and very prominent beak.
This beak is generally purple; it is sometimes hollow, and occupied by
a thread of corium. Its formation, and the apparent sliding up of the
whole tergum, so as to project above the scutum, has been described
under the family. From an account given to me by a person who kept
_B. porcatus_ alive, the beaks appear to be used, when the operculum
is touched, as an organ of defence,--the animal striking with them.
The tergal margin is more or less inflected; and the carinal margin
is convex in different degrees, and, in some species, is added to
by upturned zones of growth. The basal margin either forms a nearly
straight line on opposite sides of the spur, or more commonly slopes
towards it in various manners. The spur, or basal projection, is
rarely placed in the middle of the basal margin, generally near,
sometimes close to the basi-scutal angle; it varies much in length
and breadth, and is sometimes even half the width of the valve. The
surface of the valve is almost always more or less depressed, sometimes
so much as to form a deepish furrow, the "longitudinal furrow," which
extends from the apex to the extremity of the spur. When the furrow is
deep, its sides, as the specimen grows old, almost always become folded
inwards, so as to touch, and then the furrow becomes converted into a
closed fissure: in this latter case the folded sides generally form a
central crest on the spur. Internally, in the middle of the upper part
of the valve, the articular ridge is more or less prominent, forming
the carinal margin of the articular furrow, in which the articular
ridge of the scutum is lodged; occasionally, however, this articular
ridge can hardly be said to exist. In most species the tergal depressor
muscle is attached to sharp crests on the basi-carinal corner of the
valve, but these are almost obliterated in other species.
_Compartments._--The external appearance of the shell has already been
described. In the most typical species, the parietes consist of an
outer and inner lamina, separated by strong longitudinal septa; these
septa are denticulated on both sides at their bases, but only close
to the inner lamina; in fact the inner lamina is apparently formed by
the union, thickening, and production, of some of the denticuli. As
it is not the innermost of the denticuli on the basal edges of the
longitudinal septa, which thus become united into a solid layer, the
longitudinal septa form slightly projecting, longitudinal ribs on the
inner lamina. These internal ribs are longitudinally striated; in
old specimens they often become obliterated, especially in the upper
part of the shell. The parietal tubes or pores (occupied by threads
of corium) are generally square and large; but in _B. Ajax_ they
are very small, and in _B. glandula_ often extremely minute. In the
upper part of the shell, and sometimes low down, they are generally
crossed by thin, transverse, calcareous septa: in some species, as
in _B. perforatus_, and in some varieties of _B. amphitrite_, the
upper ends of the tubes are filled up solidly with shell. In some
varieties of _B. crenatus_ and of _amphitrite_, the longitudinal septa,
near the outer lamina, divide, thus giving rise to a very imperfect
row of outer short tubes. In _B. vinaceus_ (Pl. 2, fig. 7 _d_) the
inner lamina is cancellated instead of being solid, which is caused
by the basal denticuli of the longitudinal septa being simply united
together by their ends and crossed by transverse septa, instead of
being consolidated into a mass. In several species, as in _B. Hameri_,
the walls consist only of the outer lamina with longitudinal ribs, no
inner lamina having been formed; the ribs here evidently answer to the
longitudinal septa in the foregoing species. In _B. flosculus_ and
_imperator_ the walls are solid, their basal margins being formed of
irregular, elongated points, and little ridges (Pl. 8, fig. 4 _c_),
which apparently prefigure the more regular longitudinal ribs or septa.
In _B. balanoides_ the walls are generally either nearly smooth and
solid, or irregularly cancellated; in _B. cariosus_ (Pl. 7, fig. 3 _b_)
two or three rows of short irregular tubes are formed by unequally
branching septa, almost as in the genus _Tetraclita_.
The _Radii_, in all the species, are constructed essentially on the
same plan as the parietes; thus, in the typical forms, there is an
outer and inner lamina, with septa, which, near the inner lamina, are
furnished with denticuli on both sides; hence the radii are permeated
by pores or tubes, like the parietes; but this holds good only in the
first section of the genus, for, in the other species, the tubes are
filled up quite solidly. The denticuli on the septa often occur only on
one side, or disappear altogether; and, lastly, the septa themselves
often appear merely like little teeth, or disappear altogether as in
_B. Hameri_, or occur only near the bases of the radii, as in _B.
amaryllis_. A slight furrow in the compartment, against which each
radius abuts, is generally marked by the septa and their denticuli. In
regard to the _alæ_, their lateral or sutural edges are either thin
and smooth, or, more commonly, finely crenated or ribbed. The little
transverse crenations are homologous with the septa in the radii and
parietes. The edges of the alæ are usually received in a furrow. The
diametric growth of the shell is effected by the growth of the radii
and alæ, and chiefly by that of the former. The sutural and lateral
edges of both radii and alæ are added to, either quite up to their
summits, or only low down, and during the continued growth of the
shell, lower and lower down; in accordance with this difference in
growth, the summits of the radii and alæ become either very oblique,
or they extend parallel to the basis, that is, from tip to tip of the
adjoining compartments. When the radii and alæ are added to, as is most
usual, above the level of the opercular membrane, and therefore above
the sack, ribbons of corium run up the sutures from the sack, higher
or lower, according to the height to which, in the different species,
the edges of the radii and alæ continue to be added to. The obliquity
of the summits of the radii and alæ varies, in some cases, in the same
species. It often happens that when the summits of the radii are very
oblique, the summits of the alæ are but little so; and the converse;
both, however, are often either equally oblique, or both have square
summits. The _sheath_ extends either one third or more than half down
the shell; its basal margin often (Pl. 25, fig. 1, K′) freely depends
or overhangs the inner lamina of the walls.
_Basis._--In typical species the basis is calcareous, and consists of
an upper and lower lamina, separated by radiating septa, forming pores.
In the same manner as the septa of the parietes sometimes, though
rarely, become irregularly divided near the outer lamina, forming
outer pores, so it is, but in a much more marked degree, with the
basis. The basis in such cases becomes extremely thick, and consists
of an upper, thin lamina, with the regular radiating septa and pores,
and of an underlying, thick, cancellated mass, which seems wholly to
result from the dividing and sub-dividing of the septa. The basal
radiating pores, like the parietal pores, are closed at intervals by
calcareous transverse septa. The basal points of the parietal septa
enter the orifices of the basal pores, and the threads of corium pass
into the latter, between the denticuli of the parietal septa. In some
species, as in _B. crenatus_ and _Hameri_, the basis is perfectly
solid, the upper lamina being absent, just as in some species, the
internal lamina of the parietes is absent. In _B. flosculus_ the basis
is calcareous, but consists of so excessively thin a film as hardly
to be distinguished: it presents, moreover, as also is the case with
_B. imperator_, a beaded structure. Again, in some few species, as
in _B. balanoides_, the basis is simply membranous. When the basis
is thin, it is always flat, and is closely moulded to the irregular
surface of attachment; and in this case, when specimens are crowded
together, their elongation is effected exclusively by the growth of the
walls; but, when the basis is thick, it sometimes becomes, in crowded
groups, deeply, but irregularly, cup-formed, or cylindrical, as in _B.
psittacus_ and _perforatus_. In _B. allium_, however, which inhabits
massive corals, the basis is as regularly concave or cup-formed as
in the genus Pyrgoma: in _B. calceolus_ and its allies, and in some
varieties of the fossil _B. inclusus_, the basis is boat-formed, with
its lower surface deeply grooved longitudinally from clasping the
stem of the Gorgonia or other zoophyte, to which it was attached.
In certain varieties of _B. lævis_ it is very remarkable that the
deeply cup-formed basis becomes, owing apparently to the whole shell
having grown too deep for the animal, half-filled up with irregular,
calcareous, transverse plates (Pl. 4, fig. 2 _a_), resting one upon the
other by irregular points or pillars. The cementing apparatus has been
sufficiently described under the Family.
_Mouth._--The _labrum_ is always notched; sometimes it has no teeth,
but generally there are three on each side; in _B. balanoides_ there
are five or six on each side; and in _B. improvisus_ and _eburneus_
there is a whole row of teeth (Pl. 26, fig. 2, _e′_), graduated in
size, on each side of the notch. The _palpi_ are large, with their
apices nearly touching, and furnished with long spines. The _mandibles_
have, as it appears, normally, five teeth, but the two lower teeth
are always small and often rudimentary, and almost confluent with
the inferior, sometimes spinose angle. The _maxillæ_ have either a
simple edge, or a notch under the pair of large upper spines, or the
lower part forms (Pl. 26, fig. 7) a step-formed projection: there are
generally two lower spines, placed singly or not in pairs, larger than
the others, with the exception of the uppermost pair. The _outer
maxillæ_ are, on their inner faces, obscurely divided into two lobes.
_Cirri._--The rami of the first pair are unequal, the shorter one
sometimes not being more than half the length of the other ramus:
the segments of the shorter ramus are broad, and are, together with
the lower segments of the longer ramus, thickly clothed with spines;
in some species, as in _B. perforatus_, the anterior surfaces of the
segments, more especially of the shorter ramus, and of both rami of the
second pair are produced (Pl. 29, fig. 4), so as sometimes to form very
remarkable projections. The segments of the second and third pairs are
always thickly clothed with spines, as also are their pedicels. The
third pair is rather longer than the second; but in _B. vestitus_ and
_imperator_ it is much longer, and is otherwise somewhat different.
The dorsal and basal margin of the pedicel of the third pair, in some
of the species, as in _B. tintinnabulum_, is produced backwards on the
thorax, and forms a membranous plate fringed with fine spines. The
three posterior and longer pairs of cirri have from three to rarely
eight or ten pairs of long spines on each segment, with generally one
or two minute spines in the middle between each pair: their pedicels
have a regular double row of spines.
The _penis_ is long and hairy: in most of the species there is, at its
dorsal basis, a small, sharp, flattened, imperforate projection; first
observed by Poli: but this is sometimes absent, as in _B. crenatus_,
though present in the closely allied _B. balanoides_; and its presence
is variable in _B. tintinnabulum_. All the species have large plicated
_branchiæ_. The base of the _sack_ in several species is furnished
with inwardly projecting filamentary appendages. In _B. perforatus_,
_crenatus_, and _improvisus_, and I believe in other species, the upper
part of the _stomach_ is furnished with a circle of branching cæca.
_On the variation of the species_; _their arrangement and affinities_;
_value of the characters used_; _changes during growth_.--Owing to the
great variation in external characters, to which almost all the species
are subject, and likewise to the genus being a very natural one, that
is, to the species following each other in close and natural order, it
is not easy to exaggerate the difficulty of identifying the species,
except by a deliberate examination of the internal and external
structure of each individual specimen. Every one who has collected
sessile cirripedes must have perceived to what an extent their shape
depends on their position and grouping. The surface of attachment has
a great effect on that of the shell; for as the walls are added to at
their bases, every portion has at one time been in close contact with
the supporting surface; thus I have seen a strongly-ribbed species
(_B. porcatus_) and a nearly smooth species (_B. crenatus_) closely
resembling each other, and both having a peculiar appearance, owing
to their having been attached to a pecten. Dr. Gray has pointed out
to me specimens of _B. patellaris_, curiously pitted like the wood
to which they had adhered; and numberless other instances might be
added. Quite independently of the effect produced by the surface of
attachment, the degree to which the longitudinal folds and ribs are
developed on the parietes, is variable in most of the species, as in
_B. tintinnabulum_, _vestitus_, and even in _B. porcatus_; the presence
or entire absence of these ribs often surprisingly alters the whole
aspect of the shell. The persistence of the so-called epidermis is in
some degree variable; and in _B. lævis_ we have groups of specimens
absolutely naked, and others uniformly clothed with a brown membrane.
Again, some species in certain localities are all subject to the
disintegration of the entire outer lamina of the walls; and in such
cases (as with _B. perforatus_) there is not the smallest resemblance
between the corroded and perfect specimens. The size of the orifice,
and consequently of the operculum, compared with the shell itself,
varies accordingly as the shell is more or less conical or cylindrical;
in the latter case, the summits of the radii are generally more oblique
and the aperture consequently more deeply toothed than in the more
conical varieties. Size is a serviceable character in some cases, but
very many specimens are required to ascertain the average or maximum
size of each species, for there is no method of distinguishing a
half-grown from a full-grown specimen; and I believe, as long as the
individual lives, so long does it go on moulting and growing. _Colour_
is of very considerable service; though the precise tint varies greatly
in almost every species; and what is a far more serious evil, the
majority of the species have their white or nearly white varieties, the
latter sometimes as numerous as the coloured ones: in _B. perforatus_,
_lævis_, _flosculus_, _amphitrite_, and in several other species, the
common white varieties are eminently deceptive.
Besides the slight variation in the obliquity of the summits of the
radii and alæ, dependent on the more or less cylindrical form of
the shell, in some species, as in _B. tintinnabulum_, _amphitrite_,
_improvisus_, _trigonus_, and _porcatus_, their obliquity also varies
occasionally from unknown causes, and thus greatly affects the general
appearance of the shell. In some few species, as in _B. perforatus_,
the radii are often either not at all developed, or are of very
variable width; in others, when the shell has become cylindrical, or
when very old, the radii cease to grow, and from the disintegration of
the whole upper part of the shell, with the continued growth of the
lower part, the radii at last come to exist as mere fissures: I have
seen instances of this in _B. psittacus_, _nigrescens_, and _porcatus_.
Nevertheless, the obliquity of the upper margin, and the breadth of the
radii are useful characters; and still more useful is the fact whether
the upper margins are smooth and arched, or straight and jagged. The
fact of the terga being more or less beaked is useful: as is, likewise,
the presence of striæ, or furrows, or rows of pits, radiating from the
apices of the scuta; but to ascertain the presence of these marks,
it is almost invariably necessary to take out the scuta, clean, and
examine them with a lens; these ridges and furrows, moreover, in some
species, as is strikingly the case with _B. tintinnabulum_, and in
less degree with _B. trigonus_, _lævis_, and _concavus_, appear and
disappear, and vary without any apparent cause.
Now if we reflect that form, size, state and nature of the surface,
presence of epidermis, relative size of the orifice, presence of
longitudinal ribs, tint, and often the existence of any colour, are
all highly variable in most of the species; and that the obliquity
of the summits of the radii, and the presence of longitudinal striæ
on the scuta, are variable in several species; we shall perceive how
difficult it must ever be to distinguish the species from external
characters. As some evidence of this, I may mention that, after having
described nearly 40 species, and when my eye was naturally able to
appreciate small differences, I began carefully to examine varieties of
_B. tintinnabulum_, _amphitrite_, _improvisus_, _porcatus_, _vestitus_,
&c., without even a _suspicion_ that they belonged to these species, at
that time thoroughly well known to me; yet in the cases here referred
to, there could be no doubt, when a perfect series was examined,
that the specimens were only varieties. From this cause the labour
of naming a collection is great. Let no one attempt to identify the
species of this genus, without being prepared to disarticulate, clean,
and carefully examine with a microscope the basis and parietes, and
both the under and upper surfaces of the opercular valves; for I feel
convinced, that he will otherwise throw away much labour. Moreover, in
many cases, it is almost necessary, on account of the variability of
the characters, to possess several specimens. From these facts, I have
not hesitated to form my sections on characters which require close
examination, though I would gladly have seized on external characters,
could I have found such even moderately constant.
The least varying, and therefore most important characters, must
be taken from the internal structure of the parietes, radii, and
basis: not that these characters are absolutely invariable; thus the
porosity of the parietes is slightly variable in _B. glandula_, and
highly variable in the fossil _B. unguiformis_; it is also highly
variable in _B. balanoides_, but under a systematic point of view
this is unimportant, as the section including this latter species is
well defined by the membranous basis. The porosity of the basis is in
some degree variable in _B. nubilus_, _improvisus_, and _patellaris_;
and in _B. flosculus_ we see signs of a passage from a calcareous to
a membranous basis. Characters derived from the general shape, and
from the ridges and depressions on the under side of the scuta and
terga, especially of the scuta, are highly serviceable. The terga,
indeed, in many species, as in _B. amphitrite_, vary considerably,
and are affected by the general shape of the shell. Unfortunately the
differences are not very great between the scuta of the different
species. The cause of the opercular valves offering more useful
characters, as far as outline is concerned, than do the walls of
the shell, is no doubt due to their being almost independent of any
influence from the nature of the surface of attachment. Even the ridges
and depressions on the under side of the scuta, which are in direct
connection with the muscles and soft parts of the animal, vary to a
certain extent: thus the length and prominence of the adductor ridge
is decidedly variable in _B. concavus_ and _tintinnabulum_, and in a
less degree in _B. lævis_; the size and form of the little cavity for
the lateral depressor muscle varies in many species; so does the exact
shape and degree of prominence of the articular ridge. There is one
character in the terga, which at first would be thought very useful,
namely, whether an open longitudinal furrow, or a closed fissure
runs down the valve from the apex to the spur; but it is found that
the furrow almost always gradually closes up during growth; and as a
consequence of this, the width of the spur compared to that of the
whole valve, as well as its distance from the basi-scutal angle, and
the form of its basal extremity, all vary in some degree. The length
of the spur sometimes varies considerably, as in _B. concavus_ and
_amphitrite_. The parts of the mouth are only occasionally serviceable;
for the teeth on the labrum, and the state of the lower teeth on the
mandibles, and the presence of a step-formed projection at the lower
angle of the maxillæ, are all often variable. The relative lengths of
the two rami of the first pair of cirri, the degree of protuberance
of the segments, and the number of pairs of spines on the segments of
the posterior pairs of cirri, are sometimes useful; but the relative
lengths of the cirri, and more especially the numbers of pairs of
spines on the posterior cirri, are apt to vary. Finally, I must
express my deliberate opinion, that every part and organ, internal and
external, in Cirripedes, is liable to some amount of variation in some
of the species.
I must now point out the principal changes which supervene during
growth, and which cannot properly be called variations. In the first
place, I think, it is scarcely possible to recognise a species when
under the 1/10th of an inch in diameter. In some cases, as in _var.
d'Orbignii_ of _B. tintinnabulum_, the shell is invariably coloured
when old, but quite white when very young. Generally the tints become
very much darker with age. Some species, which usually or invariably
have, when mature, longitudinally folded walls, as _B. flosculus_
and _balanoides_, are perfectly smooth in youth. The walls in _B.
eburneus_, when young, have white, hyaline, longitudinal lines, and
are naked; whereas, with advancing age, these lines disappear, and the
subsequently formed shell becomes covered with membrane. The summits
of the radii are apt to be oblique in the young of _B. Capensis_,
_psittacus_, and _tintinnabulum_, whereas they are generally quite
square in old specimens. In _B. eburneus_, _cariosus_, and in a lesser
degree in _B. psittacus_, the scuta become longitudinally striated
only with age. On the other hand, in very young specimens of _B.
tintinnabulum_, the scuta sometimes are deeply impressed by little
pits placed in rows. I have already alluded to the longitudinal
furrow on the tergum so entirely changing its character, owing to the
edges becoming, during growth, folded inwards; this likewise causes a
decrease in the proportional breadth of the spur. In old specimens of
_B. flosculus, var. sordidus_, the whole tergum is much more elongated
than in young specimens. The basal margin of the sheath is hollow
beneath in the young of _B. cariosus_ and of some other species, but
in the old it is continuous with the inner surface of the walls. The
inner lamina of the parietes generally loses, to a certain extent, its
longitudinally ribbed character in old age. The basis is solid, instead
of being porose, in very young specimens of _B. improvisus_. In all the
species, the carino-lateral compartments, in early age, are very narrow
in proportion to the width of the lateral compartments; and in all, at
this early period, the operculum is large in proportion to the whole
shell. The number of spines on the edge of the maxillæ, the number of
segments in all the cirri, and the number of spines on each segment,
are few in early youth, and go on increasing with each successive
exuviation: the pedicels of the cirri are long in proportion to the
rami at this same early age.[84]
[84] In some specimens of _Balanus perforatus_ I made the following
enumeration of the number of segments in the cirri:--
+--------------+--------------+--------------+------------------+
| |Basal diameter|Basal diameter| Medium sized |
| | of shell | of shell | specimen about |
| | 1/20th of | 1/5th of |3/4ths of an inch |
| | an inch. | an inch. |in basal diameter.|
+--------------+--------------+--------------+------------------+
|First cirrus }| ? | 11 | 17 |
|shorter ramus}| | | |
| | | | |
|Second cirrus | 4 or 5 | 9 | 13 |
| | | | |
|Sixth cirrus | 9 or 10 | 19 |31, in another 36 |
+--------------+--------------+--------------+------------------+
In the specimen 1/5th of an inch in basal diameter, each segment
of the posterior cirri carried five pairs of spines; whereas,
in full-grown specimens, there are six or seven pairs. In the
1/20th of an inch specimen, on the inner maxillæ, there were
no spines between the upper large and the lower large pair of
spines; whereas, in the 1/5th of an inch specimen, there were five
intermediate spines, and in larger specimens nine or ten spines.
Notwithstanding the difficulties now enumerated, I hope that, owing to
having examined a vast number of specimens of the most varying species,
I have not fallen into very many errors. I have endeavoured to err
on the side of making too few instead of too many species. In those
cases, however, in which I have seen only a few specimens, I have been
sometimes compelled to decide without sufficient evidence.
I would gladly have divided this genus, already including 45 species,
into smaller genera; but so far from being enabled to do so, I
have been compelled to form my Sections (immediately to be given)
on characters not absolutely invariable, and far from obvious. I
was particularly anxious to separate the elongated species with a
boat-formed basis, which are attached to Gorgoniæ, and which form the
genus Conopea of Say, but I was unable to effect their separation even
as a sub-genus; for _B. navicula_ and _cymbiformis_ graduate in the
most insensible manner through _B. galeatus_ (the type of Say's genus)
and _B. calceolus_ into _B. stultus_, and this into _B. Ajax_; yet
this latter species has even been described as a mere variety of the
typical _B. tintinnabulum_! Indeed, so insensible is this graduation,
that the first and second sections of the genus are hardly distinct. I
fully admit, that if _B. stultus_ and _Ajax_ had never existed, _B.
calceolus_ and its three allies might have formed as natural a little
group, though difficult to be characterised, as does the sub-genus
Acasta; or perhaps this group and Acasta might have been combined
together. These same species, viz., _B. calceolus_ and its allies,
are intimately allied to _B. terebratus_ and _inclusus_, which are
contained in the last section (F) of the genus; and this shows that Dr.
Gray's proposed genus Chirona, including the species with non-porose
parietes in sections (E) and (F), could hardly have been instituted,
even if the porosity of the parietes had not been variable in _B.
unguiformis_, _balanoides_, and _glandula_. My fourth section (D),
founded on the basis not being porose, is perhaps the weakest of the
divisions, as may be seen in the list of exceptions appended to the
sectional headings.
The arrangement of the species is, I think, as natural as a linear
one could be made: every one knows how irregularly and in how many
directions the lines of affinity in every natural genus branch out.
Some few species stand rather isolated, as _B. declivis_; and _B.
allium_, _cepa_, and _quadrivittatus_ in a little group by themselves.
I have already shown how the species in the first and second sections
(A and B) blend into each other; and that the blending species are
likewise allied to some in the last section (F); furthermore, I shall
have occasion to show that these same species can hardly be separated
naturally from the sub-genus Acasta. The first section, moreover,
passes into the third (C) by _B. tulipiformis_; and the third into
the fourth (D) by _B. improvisus_, _nubilus_, _corrugatus_, and
_patellaris_: the fifth and sixth (E and F) sections are closely
connected together by _B. cariosus_ and _flosculus_; and these two
sections blend into the fourth (D) through _B. unguiformis_ and
_balanoides_, and lastly, into the third (C) section by _B. dolosus_
and _improvisus_.
The genus, as we have just seen, is hardly separated from the sub-genus
Acasta; by _B. allium_ it tends to pass through the sub-genus Creusia
into Pyrgoma; and by _B. imperator_ and _flosculus_ it graduates into
Elminius and Tetraclita.
_Geographical Distribution._--This, which is much the largest genus of
sessile cirripedes, has its species scattered over the whole world,
from the arctic regions, in lat. 74° 48′, where we have _B. crenatus_
and _porcatus_, throughout the tropical seas, to Cape Horn, where
_B. flosculus_ adheres to the coast-rocks. Many of the species have
individually very wide ranges; thus _B. tintinnabulum_ and _amphitrite_
are found throughout the warmer seas; but the wide distribution of
these species may be partly due to their frequent adhesion to ships'
bottoms: _B. crenatus_ ranges from the frozen seas, in lat. 74° 48′
north, to the West Indies and Cape of Good Hope--a wonderful endurance
of the most opposite climates. _Balanus improvisus_, again, extends
from Europe to Nova Scotia, thence southward to Patagonia, and up
the western coast of S. America, someway north of the Equator. Most
of the species have considerable ranges; thus of the six species
found on the eastern shores of northern America, five of them occur
in Great Britain. Of the thirty-six species of which the habitats
are known, exactly one third, or twelve, inhabit both the torrid and
temperate zones, these being divided by the isocryme of 68°; nine
are found exclusively in the torrid, and fifteen exclusively in the
temperate zones. Within the warmer latitudes, and especially in the
southern hemisphere, Tetraclita and Elminius to a certain extent
supplant Balanus. In depth, the species range from the upper limits
of the tidal zone to even fifty fathoms. _Balanus improvisus_ and
_eburneus_ are able to survive in brackish water. The different species
are attached to various surfaces--rocks, shells, timber, floating
objects, sea-weed, lamelliform corals, Milleporæ, Gorgoniæ, and even to
sponges. Mr. G. B. Sowerby has remarked[85] that in the species from
the southern hemisphere it is the basis, and in the species from the
northern hemisphere it is the parietes, which are elongated, when the
individuals, from being crowded together, become cylindrical; but this
is erroneous; _B. perforatus_, in the northern hemisphere, sometimes
has an elongated basis; but no doubt the basis of our commonest
species, as _B. balanoides_, _crenatus_, and _porcatus_, from being
either membranous or thin, does not become cup-shaped; whereas this
structure is conspicuous in _B. psittacus_ and _lævis_, the two
commonest species in southern South America.
[85] Darwin's 'Geology of S. America,' p, 264.
_Fossil Species._--Having already given, under the Family, some account
of the geological history of sessile cirripedes, short as it is, I here
only allude to the subject in order to state my conviction that species
cannot be satisfactorily distinguished in a fossil state, and rarely
in a recent state, without an examination of the opercular valves.
Nothing, indeed, could have been easier than to have affixed names
to many groups of specimens, having different aspects, but to feel
sure that these were really distinct species requires better evidence
than can be afforded by the shell, without the operculum. No doubt,
in some of the smaller sections of the genus--for instance, in that
characterised by a membranous basis--it would have been possible to
have distinguished some or several fossil species; but such have not as
yet been found. When the specimens are much fossilised, it is, indeed,
difficult to make out the primary points of structure--namely, whether
the parietes, radii, and basis are porose: to do this it is sometimes
necessary to rub down, polish, and carefully examine, a transverse
section of a piece of the shell.
_Sections of the Genus._
[A]
_Parietes, and basis, and radii permeated by pores._
[B]
_Parietes and basis sometimes permeated by pores, sometimes not; radii
not permeated by pores; shell elongated in its rostro-carinal axis;
basis boat-shaped, attached to Gorgoniæ and Milleporæ._
[C]
_Parietes and basis permeated by pores; radii not permeated by pores._
[D]
_Parietes permeated by pores. Basis and radii not permeated by pores._
[E]
_Basis membranous._
[F]
_Parietes and radii not permeated by pores. Basis sometimes permeated
by pores, sometimes not, sometimes excessively thin and hardly
distinguishable._[86]
[86] The following species might, owing to variation, or to the
obscurity of their structure, without care, be classed in the wrong
sections:--in A, _Balanus Ajax_, from living attached to Milleporæ,
is sometimes elongated in its rostro-carinal axis, and from having
its radii only finely porose, it might be classed in B: on the
other hand, _Bal. stultus_ is sometimes but little elongated, and
the basis hardly boat-formed, and hence might be classed in A. The
distinction between all the species in (B) and the sub-genus Acasta
is artificial.
In sections C and D, I have seen one specimen of _B. spongicola_
with a solid basis, and very young specimens of _B. improvisus_
are thus characterised, and therefore these species are liable to
be placed in the wrong section, D: _Bal. nubilus_, also, has part
of its base non-porose, and therefore likewise might be placed
in D: on the other hand, the circumference of the basis in _B.
patellaris_ is often porose, and hence this species, which belongs
to D, might be placed in C.
In _Bal. glandula_, in D, the parietes of the compartments, without
several were examined, might be thought to be solid, and therefore
this species might be wrongly placed in F; on the other hand, the
fossil _B. unguiformis_, in F, often has porose parietes, and such
specimens would naturally be placed in D. Lastly, without care, _B.
flosculus_ might be thought to have a membranous basis, and so get
placed in E.
The genus Pachylasma, without an examination of the animal's body,
might easily get misplaced in the wrong genus, amongst the species
in the last section (F) of Balanus, yet there can be no doubt that
Pachylasma belongs to the Chthamalinæ.
_Section_ A.
1. BALANUS TINTINNABULUM. Pl. 1, fig. _a_-_l_; Pl. 2, fig. 1 _a_-1 _o_.
LEPAS TINTINNABULUM. _Linn._ Syst. Naturæ, 1767.
---- ---- _Ellis._ Phil. Transact., vol. 50, 1758, Tab. 34, figs. 8
and 9.
---- ---- _Chemnitz._ Neues. Syst. Conch., 8 B. (1785),
Tab. 97, figs. 828-831.
BALANUS TULIPA. _Bruguière._ Encyclop. Meth., 1789; sed non _B.
tulipa alba_, in _Chemnitz_; neenon _B. tulipa_, _O. F.
Müller_, Zoolog. Dan.; neenon _B. tulipa_, Poli, Test. ut.
Siciliæ.
---- ---- _G. B. Sowerby._ Genera of Recent and Fossil Shells, Tab.
Genus Balanus.
LEPAS CRISPATA (_var._) _Schröter._ Einleitung Conch. vol. iii, Tab.
9, fig. 21.
---- SPINOSA (_var._) _Gmelin._ Linn. Syst. Nat.
---- TINTINNABULUM, SPINOSA, CRISPATA ET PORCATA. _W. Wood._ General
Conchology, 1815, Pl. 6, figs. 1, 2. Pl. 7, figs. 4, 5. Pl. 8,
figs. 1-5.
BALANUS TINTINNABULUM. _Chenu._ Illust. Conch.[87]
---- D'ORBIGNII (_var._) _Chenu._ Illust. Conch., Tab. 6, fig. 10,
sed non Tab. 4, fig. 13.
---- CRASSUS (Foss.) _Sowerby_ (!). Min. Conch., 1818. Tab. 84.
[87] Chenu gives several admirable figures of this species; but
he confounds some forms certainly distinct under this name, for
instance the _B. tulipiformis_ of this work.
_Shell varying from pink to blackish purple, often striped and ribbed
longitudinally: orifice generally entire, sometimes toothed. Scutum
with the articular ridge broad and reflexed. Tergum with the basal
margin generally forming a straight line on opposite sides of the spur._
_Var._ (1) communis (Pl. 1, figs. _a_, _b_, _f supra_; Pl. 2, figs. 1
_a_, 1 _c_, 1 _d_, 1 _e_, 1 _i_, 1 _k_): _conical or tubulo-conical;
smooth or moderately ribbed longitudinally; colours varying from
purplish-pink to blackish-purple; often in obscure longitudinal
stripes; orifice of shell rounded-trigonal_.
_Var._ (2) vesiculosus (_young_) (Pl. 2, fig. 1 _h_): _exterior
surface of the scuta impressed with small square holes, arranged in
two or more rows, radiating from the apex of the valve_.
_Var._ (3) validus (Pl. 1, figs. _c_, _f infra_): _globulo-convex;
coarsely ribbed, ribs flexuous; either smooth or rugged; pale
chocolate purple or pink; shell extremely strong; orifice almost
circular_.
_Var._ (4) zebra (Pl. 1, fig. _g_): _conical; rich chocolate purple
with broad snow-white ribs; sheath bright chesnut colour; summits of
alæ oblique; orifice almost circular_.
_Var._ (5) crispatus (_Schröter_) (Pl. 1, fig. _h_): _pale blueish
or pinkish-purple, with irregular rough projections, or with short,
sharp, needle-like points; scuta with their exterior surface either
plain, or with radiating lines formed of hood-like projecting points_.
_Var._ (6) spinosus (_Gmelin_) (Pl. 1, fig. _i_): _globulo-conical
or cylindrical; shell rather thin, with long, upcurved, nearly
cylindrical, very sharp points; colours very pale; attached to other
specimens, and to Lepas anatifera_.
_Var._ (7) coccopoma (Pl. 1, fig. _d_; Pl. 2, fig. 1 _f_, 1 _l_,
1 _o_): _globulo-conical; orifice small, rounded; walls generally
smooth, thick; intense rose-colour, sometimes most faintly striped
longitudinally with varying shades of pink; radii tinged with
purple; scutum sometimes as in_ var. communis, _sometimes with its
basi-tergal corner much cut off, with the adductor ridge prominent,
the pit for the depressor muscle deep, and the articular ridge broad
and hooked; tergum sometimes as in_ var. communis, _sometimes with a
broader spur, placed nearer to the basi-scutal corner of the valve_.
_Var._ (8) concinnus (Pl. 1, fig. _e_; Pl. 2, fig. 1 _g_):
_globulo-conical; walls finely ribbed; dull purple, tinged and
freckled with white; scutum, with a broad, hooked, articular ridge,
with an extremely sharp plate-like adductor ridge, and with a cavity,
bordered by a plate, for the rostral depressor muscle; tergum as in_
var. 1.
_Var._ (9) intermedius: _radii with their summits slightly oblique;
parietes pale blueish purple, with narrow dark purplish-blue
longitudinal lines; sheath with the internal surface of the rostrum
and lateral compartments much darker colored than the internal
surface of the carina and carino-lateral compartments; scuta and
terga as in_ var. communis.
_Var._ (10) occator (Pl. 1, fig. _k_; Pl. 2, 1 _b_): _radii with
their summits slightly oblique; parietes smooth, or ribbed, or
spinose; very pale blueish-purple, with narrow darker longitudinal
lines; sheath with the internal surface of the rostrum and lateral
compartments dull blue, whilst the corresponding parts of the carina
and carino-lateral compartments are white; scuta with small, sharp,
hood-formed points, arranged in straight radiating lines; terga with
the spur placed at either its own width, or less than its own width,
from the basi-scutal angle_.
_Var._ (11) d'Orbignii (_Chenu_) (Pl. 1, fig. _l_; Pl. 2, 1 _m_, 1
_n_): _radii with their summits oblique, and the orifice of the shell
rather deeply toothed; shell conical or tubulo-conical, smooth, or
rugged; colour dull purplish-lilac, with the tips of the parietes and
a band along one side of the radii quite white; sheath rather darker
at the rostral than at the carinal end; scuta as in_ var. 1; _terga
as in_ var. occator.
_Habitat._--West coast of Africa, on mytili; Madeira, on rocks; West
Indies; Cape of Good Hope, on a patella and on kelp; mouth of the
Indus; East Indian Archipelago; Sydney, Australia, attached to Lepas
anserifera, adhering to a floating cane; Peru; Galapagos Islands;
West Mexico; California. Extremely common on ships' bottoms arriving
from West Africa, India, and China, often associated with _B.
amphitrite_.
_Fossil_ Red Crag, England; Mus. S. Wood and J. de C.
Sowerby.--Touraine (?); Mus. Lyell.
_General Remarks._--This, the first species of Balanus, is, perhaps
with the exception of _B. amphitrite_, the most difficult and
variable in the genus. There are some other species which vary quite
as much in external appearance; for instance, _B. perforatus_; but
_B. tintinnabulum_ also varies in far more important points, as in
the proportions and structure of the opercular valves. The difficulty
in determining whether or not the differences are specific, is
wonderfully increased by whole groups of individuals varying in
exactly the same manner. I have seen three most distinct varieties
taken from the bottom of the same vessel, so that I did not at first
entertain the least doubt that they were three distinct species. I
may mention, as showing the vacillations which I have experienced
on this subject, that beginning with the impression, that the above
three varieties were really distinct species, after going over the
several immense collections of specimens placed at my disposal,
I came to the conclusion that the above three, and several other
forms presently to be described, were only varieties; yet after an
interval of some months, having to look at some of these specimens
again, I could not but think that I had come to a false conclusion,
and so went into all the details again, and satisfied myself that I
had followed a right course; after another interval, I had to repeat
the same process, and even now I can never look at a group of the
beautifully coloured shells with their small rounded orifices of
_var. coccopoma_ attached to the _Avicula margaritifera_, or again
at _var. d'Orbignii_, with its toothed orifice and white tips to
the compartments, without thinking that they must be specifically
distinct from the dull-coloured specimens with large entire orifices
so common on ships' bottoms; yet I can produce a full series of
intermediate forms, and I can further show, in each variety, that
the several points of difference by which each is characterised, are
variable. I may be permitted to add, in order to show that it has
not been from indolence that I have combined so many forms, that I
had named and already written out full descriptions of most of the
varieties, before determining to sacrifice them.
Seeing that _B. tintinnabulum_ and _amphitrite_ are the two most
variable species in the genus, more especially in the important
characters derived from the opercular valves, and knowing that these
species are attached so very frequently to ships' bottoms, one is led
to suspect that their extreme tendency to vary may be due to their
being exposed to varying and peculiar conditions, whilst transported
to new and distant localities. It is even just possible, as may be
inferred from the facts given in the Introduction (p. 102) in regard
to certain monstrous specimens of _Bal. balanoides_ having been
apparently impregnated by adjoining individuals, that the varieties
may interbreed, and so produce numerous intermediate forms. Whether
or not this could take place, I am inclined to look at these two
species, as in an almost analogous condition with our domestic
animals, which give rise to such infinitely numerous varieties. It
appears to me probable, that several of the varieties keep true
to their peculiarities, as long as they continue to breed in the
same locality; but that when their larvæ become attached to ships'
bottoms, and are thus transported and exposed to new conditions, they
give rise to new and ever-varying varieties. I will first give a full
description of the more common forms of _B. tintinnabulum_, which
undoubtedly belong to the same species, only alluding to the less
frequent points of difference, and then separately describe the more
marked varieties.
_General Appearance._--Shape of shell generally tubulo-conical, or
conical, or globulo-conical, rarely depressed. Orifice either large
and rounded-trigonal, or small and oval, either entire or less
frequently toothed. Surface quite smooth, or longitudinally ribbed;
ribs of variable strength, not unfrequently flexuous or branching,
sometimes roughened with blunt or sharp projecting, irregular points,
or more rarely with almost cylindrical, upturned, long spines; the
simple longitudinal ribs are generally most strongly marked in
young specimens. _Colour_, generally varying from pink, to pink
tinged with purple, to dark, inky purple, more or less striped,
longitudinally, with white or pale tints; rarely the shell is of the
brightest rose-colour, either uniform or longitudinally striped;
sometimes it is pale purplish, or dark blue; and sometimes dark
chocolate-purple: the ribs, when present, are generally more or less
white, sometimes snow-white. That there is much variation in colour,
and in the prominence of the longitudinal ribs, is quite certain,
as the two sides of the same individual sometimes differ greatly in
these respects. The radii are generally rather darker coloured than
the parietes, but sometimes they are lighter, even in the darkest
tinged specimens. The surfaces of the radii are occasionally finely
plaited in lines parallel to the basis. In some infrequent varieties
the radii have oblique summits, making the orifice of the shell to
be toothed. The sheath is generally feebly coloured, but sometimes
bright chesnut-brown, and sometimes blueish. The _strength_ of the
shell varies considerably; some of the globulo-conical varieties are
extremely massive. _Size_; basal diameter of largest specimen very
nearly three inches; height of the highest specimen three inches.
Young specimens are apt to have a peculiar aspect; for their shell
is often strongly ribbed longitudinally, and the summits of their
radii are sometimes oblique. Their scuta are sometimes deeply
pitted in radiating lines. Their colours are generally pale. I
have seen specimens attached to kelp from the Cape of Good Hope,
with their parietes white and ribbed, and their radii mottled with
pinkish-purple; I have seen other young specimens from the Galapagos
Archipelago, of a uniform grayish-blue.
The _Scuta_ generally have their lines of growth moderately
prominent; occasionally they are longitudinally striated, with the
lines of growth flexuous and upturned at intervals into small,
sharp, hood-liked projections, which are symmetrically arranged in
straight lines radiating from the apex of the valve; I have seen
this structure in some specimens of _var. crispatus_ and in _var.
occator_ (Pl. 2, fig. 1 _b_); and I have noticed an intermediate
state in _var. communis_. The degree to which the basi-tergal corner
of the valve is rounded off varies much even in _var. communis_ (Pl.
2, fig. 1 _a_, 1 _d_, 1 _e_). The articular ridge (1 _c_, 1 _e_)
is broad and much reflexed; and often, but not always, distinctly
hooked (1 _f_). The adductor ridge is confluent with the articular
ridge, and runs straight down the valve, bounding the cavity for
the depressor muscle; generally the adductor ridge is blunt, and so
little prominent as barely to deserve notice; but I have seen it
sharp and prominent in one specimen of _var. communis_, and it is
generally prominent in _var. coccopoma_ (1 _f_), and most remarkably
so in _var. concinnus_ (1 _g_). The cavity for the lateral depressor
muscle is generally very slight; but in the two vars. just mentioned,
and sometimes in _var. communis_, it is deep. In _var. concinnus_ (1
_g_) there is a remarkable plate developed for the attachment of the
rostral depressor muscle. The scuta are coloured either dull-purple
or reddish, or striped longitudinally white and blue. The surface is
sometimes externally depressed in the line of the adductor ridge; and
in young specimens there is sometimes, along this line, a chain of
pits (1 _h_), as in full-grown specimens of _B. trigonus_ and _lævis_.
The _Tergum_ (Pl. 2, figs. 1 _i_ to 1 _o_) is broad, with a generally
closed longitudinal furrow; this furrow is open in young specimens,
and it is often, but not always, open in rather large specimens of
_var. occator_; it is always open in _var. d'Orbignii_ (Pl. 2, fig. 1
_m_), and sometimes in _var. concinnus_. Apex barely beaked, except
in _var. spinosus_, in which it is sensibly produced. Spur placed
either very nearly in the middle of the basal margin, or when least
medial, it stands at above its own width from the basi-scutal angle;
yet in some specimens of _var. occator_ the spur is less than its own
width from this angle. The basal margin (1 _i_), on opposite sides of
the spur, either forms a nearly straight line, or the scutal portion
descends lower than the carinal portion, and curves very regularly
towards the spur; this is the case in _var. d'Orbignii_ (1 _n_), and
in some specimens of _var. occator_. The carinal half of the basal
margin generally forms an angle with the spur of only a little above
a rectangle. The spur varies a little in length and breadth, but
never exceeds one fourth of the greatest breadth of the valve. The
scutal margin is broadly inflected, the inflected portion forming
either a right angle, or somewhat less than a right angle, with the
exterior surface of the valve. Internally the articular ridge is
prominent, and is either considerably or slightly curved; it extends
down either about half, or three fourths, of the length of the valve.
The spur is produced for a considerable distance up the internal
surface of the valve as a prominence. The crests for the tergal
depressor muscle are very feeble. In one specimen, in which both the
shell and operculum had undergone much disintegration, the scuta and
terga were calcified together.
_Compartments_: their exterior appearance has been already described.
The parietal tubes are not large; they are generally crossed by
transverse septa in their uppermost part; but they are sometimes
almost solidly filled up by dark shelly layers. The internal surface
of the parietes is more or less plainly ribbed; in old specimens,
however, it generally becomes smooth. The _radii_ have their septa
denticulated on both sides; and they are porose, that is, the
interspaces between the septa are not filled up solidly. The radii
generally extend from tip to tip of the adjoining compartments, that
is, their summits are parallel to the basis; but in three not common
varieties, viz., _vars. intermedius_, _occator_, and _d'Orbignii_
(the former of which at least must, without the smallest doubt,
be ranked as a mere variety), their summits are oblique. I have
occasionally met with specimens of _var. communis_ with oblique
radii; and this is not very infrequent in young shells. Exteriorly
the radii are generally smooth, but sometimes finely ribbed
horizontally, owing to the projection of the septa. The _alæ_ have
their sutural edges smooth; their summits are usually parallel to
the basis, but they are often much broken; in _var. zebra_, however,
in every specimen which I examined, the summits were oblique. The
sheath varies much in colour: in _var. occator_, and in a less
degree in _var. intermedius_ and _var. d'Orbignii_, the portion
lining the rostrum and lateral compartments is much darker than
the other parts of the sheath. The _Basis_ generally has a thick,
underlying, cancellated layer. Sometimes the basis (Pl. 1, fig. _b_)
is irregularly cup-formed.
_Mouth_: labrum with four or six minute teeth: mandibles with five
graduated teeth; inferior point more or less spinose. Maxillæ, either
with or without a small notch, beneath the upper pair of spines; in
the lower part there are two spines longer than those immediately
above them. _Cirri_, the first pair has the rami unequal, in the
proportion of about 19 segments in the longer ramus, to 16 in the
shorter. The segments in the latter have their anterior surfaces very
protuberant. The second pair is short, with the anterior surfaces of
the segments protuberant. On the thorax (Pl. 25, fig. 1), on each
side, at the bases of third pair of cirri, there is a projecting
membranous plate fringed with fine bristles. The three posterior
pairs have their segments shield-shaped in front, generally bearing
four pairs of spines, of which the lower pair is minute; between
these pairs there are some minute spines. In some young specimens
from the Cape of Good Hope, and in _var. concinnus_, I found six
pairs of spines on the segments of the posterior cirri.
_Geographical Distribution._--This species is extremely common over
the whole of the warmer seas. It ranges from the Island of Madeira
to the Cape of Good Hope, and on the west coast of America, from
Monterey, in lat. 37° N., in California, to Peru. It is attached to
rocks and sub-littoral shells, to floating timber, to kelp, and to
_Lepas anatifera_. It is attached in wonderful numbers to ships'
bottoms arriving at our ports, from West Africa, the West Indies,
the East Indian Archipelago, and China. It is generally associated
with _B. amphitrite_ and _amaryllis_. I have already stated that,
on the bottoms of vessels, the different varieties are generally
grouped together; and this makes me believe that they are local. In
Mr. Stutchbury's collection there are numerous specimens taken from
a ship which first went to the west coast of Africa for guano, and
then to Patagonia for the same object, and it was interesting to see
the manner in which numbers of _B. psittacus_, a Patagonian species,
had become attached on the African _B. tintinnabulum_. The varieties
from the west coast of America seem eminently peculiar; we there find
_var. coccopoma_ and _concinnus_; and a blueish, rugged variety, with
peculiar opercular valves.
_Geological History._--I have seen specimens in Mr. S. Wood's
collection from the Red Crag of England, which, though not
accompanied by opercular valves, I cannot doubt belong to this
species. The specimens named by Mr. Sowerby, in the 'Mineral
Conchology,' as _B. crassus_, and which I have seen through the
kindness of Mr. J. de C. Sowerby, also belong to this species. I
further believe that a specimen in Sir C. Lyell's collection, given
to him by M. Dujardin under the name of _B. fasciatus_ from Touraine,
is likewise _B. tintinnabulum_.
_Varieties._
With respect to _var. communis_, I have nothing to remark. The
second, _var. vesiculosus_ (Pl. 2, fig. 1 _h_), is confined to young
specimens, and may, perhaps, be due to a want of calcareous matter.
With respect to _var. validus_ (Pl. 1, figs. 1 _c_, 1 _f infra_),
I may observe that some of the coarsest and strongest specimens
which I have seen were said to have been attached to a surface of
iron. I have seen two large lots of _var. zebra_ (1 _g_), taken by
Mr. Stutchbury, from the bottoms of ships, arriving from Bengal and
China, and in both cases associated with _var. communis_, and in one
case with _var. coccopoma_. I at first thought that this variety,
_zebra_, was specifically distinct, but now I feel no doubt, that it
is a mere variety; its body was in every respect identical with that
of _var. communis_.
Nor have I any doubt that _var. crispatus_, of Schröter (Pl. 1, fig.
_h_), is only a variety, although the scuta in some specimens have a
peculiar appearance, externally like these valves in _var. occator_
(Pl. 2, fig. 1 _b_): the scutum is here broader and flatter than in
_var. communis_, and the adductor ridge is very feebly developed,
but we shall see how variable this ridge is in all the varieties:
externally, the sharp, hood-like points formed by the upturned lines
of growth, have a very remarkable appearance, from being arranged in
quite straight radiating lines. This structure is evidently caused by
the same tendency which produces on the walls the sharp, upturned,
irregular points; but it is singular that the scuta are smooth in
some specimens with very rough parietes; and, on the other hand,
bristling with the symmetrically arranged, hood-like projections, in
other specimens on which I could with difficulty detect only a few
exceedingly minute points on the walls. In _var. communis_ I have
seen a few specimens with a slight tendency in the scuta to become
striated longitudinally. The tergum in _var. crispatus_ presents no
difference from that in _var. communis_. Some of the roughest and
best characterised specimens of _var. crispatus_ appear to have come
off copper-sheathed vessels.
I believe _var. spinosus_ of Gmelin (Pl. 1, fig. _i_), has been
correctly considered by me as a variety, but I have unfortunately
seen only one set of specimens with their opercular valves preserved:
these were attached to _Lepas anatifera_. The colour of the shell
varies from reddish-purple to nearly white; the radii are sometimes
quite white; the walls are slightly ribbed. The scuta in the above
specimens, externally were smooth; the adductor ridge was rather more
distinct from the articular ridge than in any other variety; and the
terga rather more plainly beaked. The tubular, up-curved, calcareous
spines sometimes occur only on one side of the shell, and often only
in the lower part. These spines are often coloured brighter than
the walls. Their presence cannot be accounted for (any more than
the state of the scuta in the foregoing _var. crispatus_) by the
nature of the surface to which they are attached; for I have seen one
set attached to a large rugged specimen of _B. tintinnabulum_, and
another to the very smooth valves of _Lepas anatifera_. I believe
that this form is almost always associated with _var. communis_,
which is an argument that it is only a variety.
To _var. coccopoma_ (Pl. 1, fig. _d_) I alluded in my introductory
remarks as having so strongly the aspect of a distinct species. I
possess a beautiful group, with a globulo-conical, smooth shell, of
the finest rose colour, with a rather small, rounded orifice. These
specimens were attached (mingled with _B. trigonus_) to _Avicula
margaritifera_, from, as Mr. Cuming believes, Panama. I can never
look at this set of specimens without doubting the correctness of the
determination at which I have arrived. In the British Museum there
are two sets of specimens taken off a vessel, on the west coast of
South America, almost identical in external appearance with those in
my possession, but rather more rugged. Mr. Stutchbury has sent me
some specimens from a ship, direct from China, which are rather paler
pink, and more striped, and come near to some ordinary varieties of
_B. tintinnabulum_. The scuta (Pl. 2, fig. 1 _f_) in the above three
sets of specimens agree in having the adductor ridge more developed,
and the pit for the lateral depressor muscle deeper than is usual.
The tergum (1 _l_, 1 _o_) in most, but not in all these specimens,
has a rather broader spur; and some of the specimens have the
carinal portion of the basal margin considerably hollowed out; the
spur, also, is placed nearer the basi-scutal angle than in ordinary
cases. On the other hand, in Mr. Cuming's collection, there are two
specimens taken off a vessel, identical in external appearance with
the foregoing, but which have scuta and terga in every character
exactly as in _var. communis_; hence I am compelled to consider all
these specimens as mere varieties.
_Var. concinnus_ (Pl. 1, fig. _e_) is, perhaps, the most remarkable
of all the varieties; I have seen three sets of specimens from the
west coast of South America,--all identical in appearance, having
longitudinally-ribbed walls, either rosy or of a dull purple, striped
and freckled in a peculiar manner with white. I have, however,
seen an approach to this colouring in some few specimens of _var.
communis_; and the shell itself offers no other peculiarities. The
scutum (Pl. 2, fig. 1 _g_) resembles, in general shape, that of _var.
coccopoma_; but the adductor ridge is here much sharper and more
prominent; and the rostral depressor muscle, instead of being lodged
in a little cavity formed by the folding over the occludent margin,
has, in addition, a small plate on the under side, which tends to
convert the pit into a tube. The tergum exactly resembles that of
_var. communis_. The segments in the sixth cirrus bear six, instead
of four, pairs of spines,--a circumstance which I have noticed
only in some young specimens of _var. communis_, from the Cape of
Good Hope. From these several peculiarities, until quite lately, I
resolved to keep this form specifically distinct; but I have finally
concluded that they are not sufficient. For firstly, I have seen a
scutum in _var. communis_ (Pl. 2, fig. 1 _d_), with the adductor
ridge nearly as sharp; and this ridge is always strongly pronounced
in _var. coccopoma_; secondly, with respect to the plate for the
rostral depressor muscle, although I have not seen this in any other
variety, yet in _Bal. concavus_ a closely analogous plate, situated
in the lateral depressor cavity, is highly variable, and I am not
willing to found a new species on one minute point of structure,--a
structure which is variable in another species of the same genus.
I have seen some cylindrical and conical specimens of _B.
tintinnabulum_, from the coast of Mexico and California, only
noticeable, as far as the shell was concerned, from being rugged,
and of a dull blueish-purple; but which had opercular valves
exactly like those of _var. coccopoma_, and therefore, as far as
the scutum is concerned, approaching closely in structure to _var.
concinnus_,--all three from the west coast of America. Hence I was
at one time led to believe that there existed a species on this line
of coast, which represented _B. tintinnabulum_, and which varied in
external shape and colour in an analogous manner to that species. But
as the opercular valves in _var. coccopoma_ are sometimes identical
with those of _var. communis_, and as this is always the case with
the tergum of _var. concinnus_, and as the shell itself presents
no differences, it is scarcely possible to admit the existence on
the west coast of America of this supposed representative of _B.
tintinnabulum_.
With respect to _var. intermedius_ I have little to say in addition
to the character given above: I have seen only two groups of
specimens in Mr. Cuming's collection: the chief interest in this
variety is that it shows that the next form must be ranked as a
variety, and not as a distinct species.
Of _var. occator_ (Pl. 1, fig. _k_) I have seen several specimens,
mostly taken off the bottoms of vessels, and one specimen, marked
in Mr. Cuming's collection "South Seas." After having carefully
examined these specimens, I came to the conclusion, that the slightly
oblique radii--the general colouring, and more especially that of the
sheath--the scuta (Pl. 2, fig. 1 _b_), with their sharp hood-like
points, in radiating lines--and the terga, with the spur so near to
the basi-scutal angle, were amply sufficient to distinguish it as
a good species. Subsequently, however, I found that the scuta in
_var. crispatus_ presented, both externally and internally, exactly
the same peculiar appearance. In _var. intermedius_, I found the
summits of the radii equally oblique, and the general colouring
nearly the same, and more especially a close approach to the singular
circumstance of the sheath differing in colour towards the opposite
ends of the shell. So that the position of the spur of the tergum was
the chief remaining character; and this evidently varied considerably
in the four or five specimens examined by me, being either its own
width, or much less than its own width, from the basi-scutal angle:
the outline, also, of the small portion of basal margin, between the
spur and the basi-scutal angle, likewise varied much, being either
angularly indented, or gradually curved down towards the spur: so
also the tip of the spur varied in shape. The longitudinal furrow
is unusually apt, in this variety, to remain open. We know that the
position of the spur varies considerably in _var. communis_. Hence,
although the spur, on an average, lies closer to the basi-scutal
angle in this than in any other variety, even than in _var.
d'Orbignii_, it would, I conceive, be preposterous to found a species
on this one character. In the animal's body, every part agrees
perfectly with that of _var. communis_.
Lastly, we come to _var. d'Orbignii_ (Pl. 1, fig. _l_): until quite
recently I did not even suspect that this form was only a variety
of _B. tintinnabulum_: I have examined a great number of specimens
in Mr. Stutchbury's collection, which had come attached on a vessel
from Java, and likewise a few other specimens in other collections.
They all closely resemble each other in shape, and even in size,
and differ only in tint of colour, and in the surface being either
very smooth, or longitudinally ribbed, sometimes with rugged, sharp
points. From this circumstance--from the peculiarity of the tint,
with the tips of the parietes and one side of the radii perfectly
white--and from the obliquity of the summits of the radii, I was led
to think this form specifically distinct. But the colour does not
differ from that of some other varieties of _B. tintinnabulum_; the
circumstance of the colour being uniform or not striped, is common
to the sub-varieties of several varieties, and the white tips to
the parietes, and the white borders to the radii, result simply
from the shell, whilst young, having been wholly white, and this is
not rarely the case with _var. communis_. Dismissing, therefore,
colour, it will be found that hardly any other characters remain by
which this form can be separated from _var. occator_; in both the
summits of the radii are oblique, in both the sheath is coloured in
nearly the same manner, in both the opercular valves, especially the
terga (Pl. 2, figs. 1 _m_, 1 _n_), resemble each other; the scuta,
however, are smooth in _var. d'Orbignii_ and rough in _var. occator_.
This latter form, certainly, cannot be specifically separated from
_var. intermedius_, and this assuredly is only a variety of _B.
tintinnabulum_. Hence I am led to conclude that _Balanus d'Orbignii_
of Chenu, peculiar as its whole aspect is, must be ranked only as a
variety of _B. tintinnabulum_; its oblique radii resulting from the
same cause, whatever that may be, which has given this structure
to _var. intermedius_ and _occator_; and its peculiar colouring to
having been exposed (owing probably to having been transported on
vessels) to different conditions, whilst young and old.
2. BALANUS TULIPIFORMIS. Pl. 2, fig. 2 _a_-2 _d_.
BALANUS TULIPIFORMIS EX CORALLIO RUBRO. _Ellis._[88] Philosoph.
Transactions, vol. 50 (1758), tab. 30, fig. 10.
LEPAS TULIPA. _Poli._ Test. utriusque Siciliæ, tab. 5, fig. 1. et 6
(1791).
BALANUS TINTINNABULUM (_var._) _Chenu._ Illust. Conch., tab. 3, f. 5.
[88] According to the letter of the Rules of the British
Association, Ellis's name ought to be retained, as it was published
in 1758, the same year during which the 10th edition of the
'Systema Naturæ' appeared, in which edition the binomial method
was first used. But as Ellis himself did not then know of, or
follow this method, it might be disputed whether, according to the
_spirit_ of the law, his name ought to stand. The only other name
given to this species is that of _tulipa_, affixed by Poli in 1791,
but this name had been previously used by Müller in 1776, and by
Chemnitz in 1785, for another species, the _B. Hameri_ of this
work; and likewise, also previously to Poli, by Bruguière in 1789,
for still another species, viz., _B. tintinnabulum_ of this work:
under these complicated causes of confusion, I think it is highly
advisable to adopt Ellis's name. I may add that the _B. tulipa_ of
Mr. G. B. Sowerby is the _B. tintinnabulum_ of this work. It is
possible that the _B. conoides_ of Brown, 'Illustrations Conch.'
(1st edit. pl. 6, fig. 7), may be our present species; but without
details of structure it is hardly possible to identify, in many
cases, the species of Balanus.
_Shell dark rose-coloured, sometimes tinged with purple; orifice
toothed. Scutum externally very smooth, covered by membrane. Tergum
with distinct crests for the depressor muscles._
_Hab._--Sicily, Malta, Malaga, (associated with _B. perforatus_),
Madeira. Often growing in clusters and associated with _Pachylasma
giganteum_. Attached to _Millepora aspera_, oysters, and other
shells. According to Poli, an inhabitant of deep water; yet in mus.
Cuming there are two fine specimens attached to the always floating
_Lepas anatifera._ Mus. Lowe, Macandrew, Stutchbury.
_General Appearance._--Shell tubulo-conical or conical: orifice
large, toothed, approaching to pentagonal. Surface moderately smooth,
naked. Colour rosy, or tile-red, with a slight tinge of purple; or
beautiful rich purple. Radii nearly as dark as, or darker than, the
parietes. The portion of the alæ seen externally is generally white.
Internally the whole shell is nearly white. Generally the tints
outside vary in transverse fasciæ; sometimes there are very fine,
dark, longitudinal lines. Largest specimen (from Malta), 1.4 of an
inch in basal diameter; usually full-sized specimens are about three
quarters of an inch in basal diameter.
_Scuta_ (Pl. 2, fig. 2 _a_, 2 _c_) very smooth, with the
growth-ridges very little prominent, sometimes there are obscure
traces of longitudinal striæ; surface covered by an unusually thick
and persistent yellow membrane: valve narrow, with the upper part
commonly reflexed: the basal margin forms, with the occludent margin,
a smaller angle than is usual: the tergal margin of the valve is
rectangularly inflected, instead of being, as is usual, merely
bowed inwards. Internally, the articular ridge is rather prominent.
The depth of the _slight_ pit for the lateral depressor muscle is
variable; it sometimes includes a minute, central, longitudinal ridge.
_Terga_ (2 _b_, 2 _d_): the longitudinal furrow is deep, with the
sides folded in; the spur is placed at about its own width from
the basi-scutal angle; it is moderately long, with its lower end
obliquely rounded off; but the length, breadth, and precise outline
of the lower end varies a little. The basal margin on the opposite
sides of the spur, forms a nearly straight line, but with the
portion on the carinal side very slightly hollowed out. Crests for
the depressor muscle are well developed.
_Compartments._--The radii and alæ always have their summits oblique:
the sutural edges of the radii are deeply penetrated by pores between
the strongly denticulated septa: the sutural edges of the alæ are
quite smooth: the tubes in the parietes are crossed in the upper
part of the shell by septa. _Basis_ tubular, with an underlying
cancellated mass.
_Mouth._--Labrum with the teeth either absent or very small:
mandibles with the fourth and fifth teeth rudimentary: maxillæ with
a small notch under the two upper spines; near the lower angle,
two spines, one beneath the other, are larger even than the upper
pair; beneath the lower pair, there is a tuft of fine spines.
_Cirri_, segments protuberant in one ramus of the first cirrus and
in both rami of the second cirrus; posterior cirri with the segments
short and broad, each bearing three pairs of spines, with a small
intermediate tuft.
_Affinities._--This species in all essential respects comes very near
to the three last varieties of _B. tintinnabulum_, which have the
orifices of their shells toothed. The smoothness of the scutum, with
its persistent epidermis,--its peculiar shape,--its small and not
reflexed articular ridge,--together with the crests on the tergum for
the depressor muscles, are sufficient diagnostic characters. Even in
general habit and tint of colour, this species has a different aspect
from _B. tintinnabulum_. In some respects _B. tulipiformis_ leads
into the species included in the third section of the genus.
3. BALANUS PSITTACUS. Pl. 2, fig. 3 _a_-3 _d_.
LEPAS PSITTACUS. _Molina._ Hist. Nat. Chile (1788), vol. i, p. 223.
BALANUS PICOS. _Lesson._ Zoolog. Voyage de la Coquille (1829).
---- TINTINNABULUM (var. c). _Ranzani._ Mem. di Storia Nat. tab. 3,
fig. 1-3 (1820).
---- CYLINDRACEUS. _Lamarck_, in _Chenu_. Illust. Conch. Tab. 4,
fig. 17, Tab. 5, fig. 7, sed non var. (_c._) in Lamarck,
Animaux sans Vert., (1818).
---- PSITTACUS. _King_ and _Broderip_. Zoolog. Journal, vol. v
(1832-1834), p. 332.
_Shell, pale dirty pink; orifice hexagonal. Scutum with the articular
ridge very small, confluent with the very prominent adductor ridge,
forming a tubular cavity, which extends up to the apex of the valve.
Tergum with the apex produced, needle-like, purple: spur placed at less
than its own width from the basi-scutal angle._
_Hab._--Peru, Chile, Chiloe, Patagonia. Fossil in an ancient tertiary
deposit, Coquimbo; and in a superficial, recent bed at S. Josef, in
Patagonia.
_General appearance._--Shell either almost cylindrical or steeply
conical, generally flesh-coloured, sometimes pale pink; surface
either smooth (when not disintegrated) or sometimes with the parietes
distinctly and rather strongly ribbed, with the ribs distant from
each other: I have seen six or seven ribs on the rostrum alone. The
orifice in the most perfect specimens is nearly equilateral and
hexagonal. The radii generally are very broad, but occasionally
quite narrow, and even linear. The basis is generally deeply and
irregularly cup-formed.
_Size._--This is the largest species in the family: I have seen a
specimen six inches in length and three and a half in diameter; and
another specimen no less than nine in length, though only two and a
half inches in diameter.
_Scuta._--In full-sized specimens the surface is finely striated
longitudinally, caused by the lines of growth being minutely sinuous;
but in young specimens, until they attain a basal diameter of above
half an inch, the surface is smooth. The valve is transversely
arched, a line of flexure running from the apex to the basal margin,
at about one third of the width of the valve from the tergal margin.
The basal margin is curved nearly continuously, and extends nearly
half-way up the valve; hence the basi-tergal corner is largely
rounded off. The articular ridge is but little prominent, and is not
reflexed: the articular furrow is very narrow. The adductor ridge
consists of a sharp, much-projecting plate, running down close to the
basal margin, and is confluent with the lower part of the articular
ridge. This plate and the inflected tergal margin of the valve,
together form a large and deep cavity, which extends up almost to the
apex of the valve. The depressor muscle is attached in the middle, at
the lower, open end of this cavity.
_Terga._--These are strongly beaked, the beak being from one third
to one fourth of the total length of the valve, including the spur:
the beak is very sharp, somewhat flattened, and bowed; when young,
and when well preserved, it is coloured purple: it is penetrated
by a fine tubular cavity, occupied by a thread of corium, which
extends about half-way up it. The whole valve is narrow, being about
thrice as long as wide. The spur is also long and narrow; it is
seated at less than its own width from the basi-scutal angle. The
scutal margin is not much inflected. The longitudinal furrow has its
sides, in full-grown specimens, closely folded together. The basal
margin slopes down on both sides to the spur. There are no crests,
or only traces of them, for the attachment of the depressor muscle.
Internally, the spur is prolonged, as a prominent ridge, upwards to
the beak, and serves as an articular ridge. In the middle, in the
upper part (Pl. 2, fig. 3 _d_), between this articular ridge and the
carinal margin, there is a second narrow ridge, which extends from
the lower part of the beak half-way down the valve, and then dies
out. The space between these two ridges, and the ridges themselves,
are coloured purple, and consist of harder shell than the rest of
the valve; hence, when the outer surface and the adjoining scutal and
carinal margins disintegrate, this part remains, and so forms the
beaked, purple apex.
_Compartments._--The parietal tubes are unusually large in proportion
to the size of the shell, and run up to the summit without any
transverse septa: the longitudinal septa are strongly denticulated.
The radii are penetrated by large tubes; their septa are very
strongly denticulated, and the denticuli themselves often subdivide
and branch out at their extremities. The sutural edges of the alæ
are smooth, or with a high power can just be seen to be crenated.
The radii are generally very highly developed, so that their summits
are even wider than the bases of the parietes; but, on the other
hand, in some few large specimens, the radii are either very narrow
or absolutely linear. In these latter cases, the diametric growth
has nearly or altogether ceased, whilst the walls of the shell have
continued to be added to at their bases, their summits at the same
time suffering disintegration; and thus the orifice has increased in
size.
_Basis_ generally, and occasionally very deeply, cup-formed. An
unusually thick cancellated layer in most cases forms the under side
of the basis.
_Mouth._--Labrum apparently without teeth, or with very minute ones:
mandibles with three teeth, of which the third is thicker than the
first or upper one: the fourth and fifth teeth are confluent with the
inferior angle. The maxillæ have a small notch under the upper pair
of spines; inferior part projecting and supporting two spines, placed
one below the other, and equalling in size the upper pair. _Cirri_:
the rami of the first cirrus are unequal by four or five segments;
shorter ramus and both rami of the second cirrus with the segments
extremely protuberant: posterior cirri not much elongated, with the
segments rather broad, supporting six pairs of spines.
_General Remarks._--This, which is much the largest known species
of the genus, ranges from Peru (Arica being the most northern spot,
whence I have seen specimens), along the coast of Chile, where it
is very abundant at a few fathoms' depth, at least as far south as
Southern Chiloe; it is said by Captain King to attain the largest
size at Conception. On the coast of Eastern Patagonia, I dredged
up this species from nineteen fathoms, in lat. 49°. In lat. 42°
(S. Josef), on the same eastern coast, I found fossil specimens
in beds of sand upraised between eighty and one hundred feet. In
the tertiary formation at Coquimbo, in Chile, it occurs in the
middle bed, associated with the recent _B. lævis_, and with various
mollusca, all of which are apparently extinct, indicating that the
formation is of considerable antiquity. In the living state, on
the coast of Chile, it is often associated with _B. lævis_. As it
frequently adheres to large specimens of the Concholepas, it must
sometimes be an inhabitant of shallow water. I have seen one specimen
attached to _Mytilus Magellanicus_. Mr. Cuming believes that about
six fathoms is the usual depth at which it lives. Numerous specimens
are often congregated together into great masses. Mr. Stutchbury has
some interesting specimens which he procured from a ship that had
first sailed to Ichaboe, on the coast of Africa, and afterwards
to Patagonia; consequently numerous specimens of _B. psittacus_
had become attached on _B. tintinnabulum_, and subsequently during
the voyage home, some few of the latter again had adhered on _B.
psittacus_: the contrast in the paler colour and hexagonal orifice
of this species, with the darker tints and more trigonal orifice of
_B. tintinnabulum_ was striking. At Coquimbo, in Chile, I procured
a specimen of _B. psittacus_, attached to a chain cable which had
been in the water only six months; this specimen measured 1.3 of an
inch in basal diameter, and .8 in height: this shows a rapid rate of
growth. Lastly, I may mention that it is asserted by Molina, and I am
assured by Mr. Cuming that the statement is perfectly correct, that
this Balanus, when cooked, is universally esteemed as a delicious
article of food.
4. BALANUS CAPENSIS. Pl. 2, fig. 4 _a_, 4 _b_.
BALANUS CAPENSIS ORE OBLIQUO. _Ellis._ Phil. Transact., vol. 50
(1758), Tab. 34, fig. 14.
_Shell shaded, and often longitudinally striped with bright pink.
Scutum as in B. psittacus. Tergum with the apex produced and
needle-like, white: spur placed at its own width from the basi-scutal
angle._
_Hab._--Cape of Good Hope. Attached to stems of Fuci, Algoa Bay. Mus.
Brit. and Bowerbank. Attached to a Patella, Mus. Darwin, Mus. Cuming,
and Stutchbury. Attached to floating kelp, Lagulhas Bank, Mus. James
Ross, associated with _B. tintinnabulum_ and _spongicola_.
This species comes extremely close to the South American _B.
psittacus_, and I should hardly have attached a specific name to
it, had I not examined many specimens, young and old, of the true
_B. psittacus_, from Peru, Chile, and Eastern Patagonia, and found
them all identical in the few, apparently trifling points, in
which that species differs from _B. capensis_. The animal's body
and the shell agree in every respect, excepting that the shell is
decidedly pinker, being often most distinctly and prettily striped
longitudinally with pale and bright pink. In some of the specimens
the basis is cup-formed: in some, the broad radii are pale pink,
in others they are quite white, and in this latter case a singular
aspect is given to the pinkish varieties. In very large specimens
(and I have seen one fully two inches in basal diameter) the pink
colour is extremely feeble, and the whole shell has a very rugged,
disintegrated, coarse, and sometimes dirty appearance: in most
of these large specimens the walls are more massive than in _B.
psittacus_, and the orifice of the shell rather smaller; in some,
however, the walls certainly are of unusual thinness.
The _Scuta_ differ from those of _B. psittacus_ only in the
basi-tergal corner not being so much rounded off, and consequently
in the articular ridge, which is rather more reflexed, descending in
proportion lower down the valve: the cavity at the basi-tergal corner
is in proportion broader. The valves in the two species differ, also,
but only in young specimens, in the occludent half being tinted,
both externally and internally, purple, whereas in _B. psittacus_
the whole valve, at all ages, is white. In the _terga_ the spur is
removed fully its own width from the basi-scutal angle, whereas it
is not half this distance in _B. psittacus_. The scutal margin is
here much more inflected. In _B. psittacus_ there is a remarkable
patch of purple on the inside of the valve, between the articular
ridge and a second special ridge; of this purple patch there is here
no trace, consequently the beak or apex is white. The beak, also, is
less prominent. The special ridge, just alluded to, here runs much
nearer to the articular ridge, and is less prominent: indeed, in old
specimens, it is often almost obliterated. Finally, the whole valve,
in proportion to the Scutum, is rather broader.
I have seen a young specimen, about a quarter of an inch in basal
diameter, with the orifice of the shell toothed owing to the
obliquity of the summits of the radii; and this gave the shell a very
peculiar aspect. The largest well-coloured specimen which I have seen
is 1.2 of an inch in basal diameter; but in Mr. Cuming's collection
there are two rugged, disintegrated specimens, two inches in basal
diameter, and two and a half in height. Some specimens, 1.3 in basal
diameter, in Mr. Stutchbury's collection, are remarkable from the
radii having been obliterated--the shell being merely divided by six
sutures, as we have seen is likewise sometimes the case with large
specimens of _B. psittacus_.
This species is evidently a South African representative of the South
American _B. psittacus_.
5. BALANUS NIGRESCENS. Pl. 2, fig. 5 _a_, 5 _b_.
BALANUS NIGRESCENS. _Lamarck_, (1818) in _Chenu_. Illust. Conch.,
Tab. 4, fig. 16.
---- GIGAS. _Ranzani._ Memoire di Storia Nat., 1820, Tab. 3, fig. 5,
6, 7.
---- ---- _De Blainville._ Dict. des Sc. Nat., Tab. 116, fig. 2, 2
_a_.
_Shell cinereous, tinted with pale or blackish blue, or wholly white.
Scutum with the articular ridge terminating downwards in a small,
sharp, free point: adductor ridge prominent. Tergum with the apex
produced and needle-like._
_Hab._--Swan River, West Australia, Mus. Brit., attached to
sandstone. Attached to sandstone and to each other in a group, Mus.
Cuming. Twofold Bay, S. E. Australia, attached to tidal rocks and
Patellæ, Mus. Darwin.
There can be no doubt that this species is the _B. nigrescens_ in
Chenu, who had access to Lamarck's original specimens; and there can be
equally little doubt that it is the _B. gigas_ of Ranzani, collected,
during Baudin's expedition, at King George's Sound: it is essentially
allied to _B. psittacus_, but in external appearance strikingly
resembles some of the varieties of _B. tintinnabulum_.
_General Appearance._--Shape tubulo-conical: walls smooth, sometimes
longitudinally ribbed: colour ashy-gray tinged with blue, but many
specimens are dark purplish-blue, owing to the disintegration of the
outer lamina, and consequent exposure of the almost solidly filled
up, dark blueish parietal tubes; on the other hand, some specimens
are quite white. Ranzani describes the colour as earthy-violet, which
is very characteristic of some of the specimens. The orifice is apt
to be rather small, compared to the size of the specimens, and tends
to be hexagonal. The radii are often rather narrow. The opercular
valves are tinted pale blue. The basal diameter of the largest
specimen is two inches, and its height two and a quarter.
The _Scuta_ have their basi-tergal corner much rounded off, as in
_B. psittacus_, so that the tergal margin does not extend more than
half down the valve. The surface is somewhat prominent, along a line
running from the apex to the point of chief curvature in the basal
margin. The surface is not striated. Internally, the articular ridge
is little prominent, and not reflexed; the lower end depends as a
free, sharp style or point. The adductor ridge is moderately sharp,
and stands some little way distant from the articular ridge: it is
produced downwards, and forms a moderately deep and large cavity for
the depressor muscle; but this cavity is not closed, and does not
extend up, as in the two last species, to the apex of the valve.
_Terga_, narrow, with a sharp, prominent, needle-like beak. Spur,
long, narrow, placed at less than its own width from the basi-scutal
angle: the basal margin on both sides slopes down to the spur: the
scutal margin is not inflected. Internally, the articular ridge is
very feebly developed, but extends down close to the basi-scutal
angle. On the under surface in the upper part of the valve, there is
a short, very slight ridge, extending on the carinal side, near and
parallel to the articular ridge; this slight ridge plays an important
part, as in the two foregoing species, in the formation of the beak
or apex. Crests for the depressor muscle are hardly distinguishable.
The _Walls_ appear to vary in some degree in strength and thickness;
as is likewise the case with the opercular valves. In some of the
thinner specimens, the parietal tubes are large, and the longitudinal
septa are furnished with small, sharp denticula. The tubes are often
thickly lined or almost filled up solidly with blue shell; they are
not crossed by transverse septa.
The _Radii_ vary in width; externally they are often finely ribbed
transversely, at other times they are smooth; their septa are fine
and thin, with their delicate denticuli not extending to the outer
lamina: they are very porose. The _alæ_ have their summits parallel
to the basis; their sutural edges are most finely crenated. The
sheath is blueish, excepting the wedge-like portions of the alæ which
have been added during the diametric growth, and these are white.
_Mouth_: labrum without teeth: mandibles with five sharp teeth:
maxillæ with the edge straight. _Cirri_, first pair with the rami
very slightly unequal; segments of the shorter ramus and of both rami
of the second pair protuberant: posterior cirri with the segments
shield-shaped in front, bearing four pairs of spines, of which the
upper pair is much longer than the lower pairs; each pair has a small
intermediate tuft of minute spines.
6. BALANUS DECORUS. Pl. 2, fig. 6 _a_, 6 _b_.
_Parietes pale pink; radii rather darker. Scutum with a small articular
ridge. Tergum with the longitudinal furrow very shallow and open; basal
margin on both sides sloping towards the spur._
_Hab._--New Zealand. Mus. Brit., and Flower: attached to shells.
_General Appearance._--Shell conical or tubular, with a large
rhomboidal orifice; very pale pink, but tinted yellowish from the
persistent epidermis, and sometimes faintly striped longitudinally;
radii and sheath of rather a darker pink; scuta in themselves white,
though lined by purple corium; the carinal half of the tergum pink.
Walls extremely smooth. Largest specimen above one inch in basal
diameter.
_Scuta_, with the finest striæ radiating from the apex; growth-ridges
moderately prominent; articular ridge small; there is a very slight
and blunt adductor ridge: the hollow for the lateral depressor muscle
is rather narrow and deep.
_Terga_, with the apex slightly prominent or beaked; the longitudinal
furrow is of very little depth; on its scutal margin there is a
narrow, rounded, slightly prominent ridge, which, however, appears
more like a furrow than a ridge. Spur moderately long and blunt;
placed at half its own width from the basi-scutal angle; the
basal margin on both sides of the spur, slopes gently towards it.
Internally, the articular ridge is pretty well developed; the scutal
margin is not much inflected; the carinal portion of the under
surface of the valve is rough; the crests for the carinal depressor
muscle are entirely absent.
_Compartments._--Walls moderately strong; parietal tubes small, with
transverse septa in their upper ends; inner surface of the walls much
less strongly ribbed than is usual. _Radii_ broad, with their summits
parallel to the basis; their septa are strongly denticulated. _Alæ_
with their summits oblique; their sutural edges are barely crenated.
_Basis_, thin, flat, or cup-formed. _Body_ unknown.
_Affinities._--In general appearance this species comes near to _B.
psittacus_; but in all essential characters it comes much closer
to the following species, from which, however, it can easily be
distinguished by colour, and by the inner lamina of the parietes not
being cancellated.
7. BALANUS VINACEUS. Pl. 2, fig. 7 _a_-7 _d_.
_Shell purplish dark brown: inner lamina of the parietes cancellated.
Scutum finely striated longitudinally. Tergum with the longitudinal
furrow shallow and open; basal margin on both sides sloping towards the
spur._
_Hab._--West Coast of South America. Mus. Cuming.
_General Appearance._--Shell conical, with a large, rhomboidal
orifice; walls rather thin, coloured, together with the radii and
operculum, dark purplish-brown; sheath nearly colourless. Walls
smooth, slightly irregular, very finely striated longitudinally.
Basal diameter of largest specimen .8 of an inch.
_Opercular Valves_, unusually smooth, that is without prominent
growth-ridges. _Scuta_, finely striated longitudinally, with the
sharp striæ closely approximate. The teeth on the occludent margin
are sharp, and stand some way apart from each other. Internally, the
whole surface is remarkably flat and smooth: the articular ridge is
of moderate breadth, and slightly reflexed: there is no adductor
ridge, and the oval depression for the lateral depressor muscle is
extremely slight. _Terga_, with the longitudinal furrow very slight;
the bottom of this furrow is feebly striated longitudinally, and
there is a trace of a fine, rounded ridge on the scutal margin, as in
_B. decorus_. The basal margin slopes on both sides towards the spur,
which is of moderate length and breadth, with its lower end truncated
and parallel to the carino-basal margin; the spur stands at about
once and a half its own width from the basi-scutal angle. Internally,
the valve is lined by very dark, purplish-brown corium; the articular
ridge is prominent; in the upper part of the valve, parallel to the
articular ridge, there are two or three feeble ridges; there are no
crests for the tergal depressores.
The _Parietes_, though moderately thick, yet are light and fragile;
the denticuli at the bases of the longitudinal septa are prominent,
and those on the adjoining septa are united together, making a
network (Pl. 2, fig. 7 _d_), but the interspaces between them are not
filled up by solid calcareous matter (as is the case with every other
species of the genus), but are only crossed at successive levels by
fine transverse calcareous septa; the internal lamina thus becoming
cancellated, and, though thick, fragile. Hence, in a transverse
section of the parietes, the ordinary parietal tubes or pores are
seen to be lined on their inner sides by five or six rows of very
minute pores. I have not seen any other instance of this structure.
The internal lamina is ribbed, as usual, on its inner surface, by
the projection of the longitudinal septa. The ordinary parietal
tubes are open, to nearly the summit of the shell. The _radii_ are
rather thin, and unusually fragile; their summits are parallel to
the basis: their septa, as seen on the sutural edges, are extremely
thin and denticulated on both their upper and lower surfaces, on the
side towards the internal lamina: towards the external lamina, the
septa are simple, and the small square pores thus formed, are open
or not filled up. The _alæ_ have their summits extremely oblique,
being added to very little during the diametric growth of the shell;
the narrow margin, however, which is thus added, is coloured red,
the rest of the sheath being nearly colourless: the sutural edges
of the alæ are smooth. The _basis_ has a thick, underlying, finely
cancellated layer of shell.
_Animal's body_ unknown.
A young specimen, .2 of an inch in basal diameter, differed from
the above in being of a much paler purplish-brown. This species is
distinct from all its congeners, in its peculiar colour, and likewise
in the structure of the inner lamina of the parietes. As already
stated, it comes nearer to _B. decorus_ than to any other species.
8. BALANUS AJAX. Pl. 3, fig. 1 _a_-1 _d_.
BALANUS TINTINNABULUM (_var._) _Chenu._ Illust. Conch., Tab. 2, fig.
8.
_Shell globulo-conical, often elongated in the rostro-carinal axis,
pale pink, smooth, extremely massive: parietal pores, close to the
basal margin, circular and very small. Scutum with the articular ridge
broad and reflexed._
_Hab._--Philippine Archipelago, attached to _Millepora complanata_,
Mus. Cuming. Mus. Brit. and Stutchbury.
_General Appearance._--Shell globulo-convex, sometimes much elongated
in its rostro-carinal axis; smooth; walls excessively strong,
massive, and heavy. Orifice oval, rather small in proportion to the
size of shell, this being chiefly due to the infolding of the upper
part of the rostral compartment. Parietes pale pink, feebly tinted
with purple: radii either paler, or tinted of a bright chesnut-brown:
sheath rich purplish chesnut-brown. Basal diameter of the largest
specimen nearly 3-1/2 of an inch; height 2-3/4: another specimen had
a basal longitudinal diameter of 2.9 of an inch, and a transverse
diameter of only 1.6; this great difference in the two diameters
being caused by the prolongation of the basal portion of the rostrum
in the line of the branch of the Millepora, to which the shell had
adhered; the height of this same specimen was 1.5; and the diameter
of the orifice, both transversely and longitudinally, .75 of an inch.
_Scuta_, broad, feebly tinted with pink; exterior surface rough,
with sharp hood-formed projections, arranged in straight lines
radiating from the apex; an inflected portion of the valve along the
tergal margin is not roughened. Internally (Pl. 3, fig. 1 _d_), the
articular ridge is broad and reflexed. An adductor ridge can hardly
be said to exist, but a slight prominence borders the gentle hollow
in which the lateral depressor muscle is attached. The basal margin,
on its inner face, is slightly toothed. _Tergum_ white, with the
narrow part of the valve, on the scutal side of the spur, rough with
the little projecting hoods, like those on the scutum; the other and
larger half is smooth: spur rather long, narrow, placed at twice its
own width from the basi-scutal angle; on the carinal side, about half
of the basal margin slopes down towards the spur. The longitudinal
furrow is either quite or nearly closed. Internally, the spur is
produced upwards on the valve, as a prominence: the articular ridge
is not very prominent. There are no crests for the tergal depressor
muscle.
Altogether the opercular valves strikingly resemble those of _B.
tintinnabulum_, but all the characters above mentioned have not been
observed in any one variety of this species; perhaps _var. coccopoma_
comes nearest, both in the external appearance of the shell and in
the structure of the opercular valves, to _B. Ajax_.
The _Compartments_ are remarkably compact and solid; the parietal
tubes are cylindrical and quite minute even close to the basis; they
extend, however, nearly up to the top of the shell; the parietal
septa at the basis are thick, and with blunt denticuli; the thickness
of the walls in the upper part of the shell is excessive; in the
lower part, it is also unusually great, owing to the thickness of the
inner lamina, and hence the ribs, generally formed by the projection
of the longitudinal septa on the inner lamina, are here visible only
close to the basis. The _radii_ are rather wide; their summits are
parallel to the basis; the septa on their sutural edges are thin,
straight, and closely approximate, and most symmetrically furnished
with little denticuli of equal sizes on both sides: the interspaces
are nearly filled up solidly, but with some pores still left open.
In the upper part of the shell, the radii, like the walls, are of
extraordinary thickness: the septa are transverse and horizontal,
as seen externally by slight variations in the colour of the radii;
internally, as seen in a vertical section of the shell, the septa
dip inwards at an angle of above 45°. The _alæ_ are thin, and have
their summits oblique: their sutural edges are smooth. The pores in
the basis are crossed by numerous transverse septa, and there is an
underlying cancellated layer: the internal surface is very smooth.
_Animal's body_ unknown.
The strength of this Balanus is truly remarkable; and when, by
repeated blows, a specimen which I was examining at last yielded,
the radii broke sooner than separate at their sutures. In most of
its characters, this species approaches _B. tintinnabulum_, and I
believe has been included by Chenu as one of its varieties; but
it comes almost equally near to _B. stultus_, to which it is much
more closely allied in its habit of being attached to Milleporæ.
By a close and unbroken chain of affinities, _B. Ajax_, through _B.
stultus_, is connected with _B. calceolus_ and its allies in section
(B), which live attached to Gorgoniæ. Some of the specimens of _B.
Ajax_, are almost as much elongated in their rostro-carinal axis, as
are the species in section (B); and there is an affinity in the same
direction in the smallness of the pores in the radii of _B. Ajax_;
indeed, had the basis in this species been generally more boat-or
cup-formed, I should have placed it as the first species in section
(B), instead of, as at present, the last species in section (A). The
intermediateness of the characters of _B. Ajax_ has been one chief
cause why I have rejected the genus Conopea, which was instituted by
Say for the species living attached to Gorgoniæ.
_Section_ B.
_Parietes and basis sometimes permeated by pores, sometimes not: radii
not permeated by pores: shell elongated in its rostro-carinal axis:
basis boat-shaped: attached to Gorgoniæ and Milleporæ._
9. BALANUS STULTUS. Pl. 3, fig. 2 _a_-2 _d_.
_Parietes and base porose: shell white, or faintly tinged with purple.
Scutum with the basal margin protuberant in the middle. Tergum with
the longitudinal furrow closed in the upper part: spur not closely
adjoining the basi-scutal angle._
_Hab._--Attached to Milleporæ, Singapore, Mus. Cuming. West
Indies,[89] Mus. Brit.--Mus. Stutchbury.
[89] This specimen in the British Museum was purchased at the sale
of the Rev. L. Guilding's collection, and therefore it is not
certain that this habitat is correct; but as it was sold in the
same lot with a Cirripede certainly West Indian, and as the main
collection was made in the West Indies, this habitat may, I think,
be trusted.
I have considerable doubts whether it would not have been more correct
to have placed this species in the last section, instead of where
it now stands; it certainly is more closely allied to _B. Ajax_,
especially in its operculum, than to the following species; yet the
fact of the radii not being permeated by pores does not permit of its
admission into the last section; and both in habits and structure it
undoubtedly comes very near to the following species. Those varieties
which are not much elongated, and which have the basis nearly flat,
would certainly, if considered by themselves alone, not have gained
admission into our present section.
_General Appearance._--Shell conical, somewhat globular, more or less
elongated in the rostro-carinal axis, owing to the basal production
of the rostrum. Orifice, rather small, entire, oval, pointed at the
carinal end. Radii moderately broad, with their summits parallel to
the basis. Colour dirty white, often faintly tinged with purple;
sheath, pale purplish-blue. Surface extremely smooth; the parietes
are generally covered (as viewed through a lens) by a very thin,
yellowish epidermis, giving to the whole a glistening, granular
aspect: the radii are generally destitute of this epidermis, and are
therefore of a dead white. The basis is concave, and sometimes deeply
cup-formed; it is, however, not symmetrical; sometimes it is flat.
Basal diameter of largest specimen, including the basis itself, 1.5
of an inch in the longitudinal axis; transverse diameter, 1 inch; the
inequality in the length of the two diameters is rarely so great as
in this unusually large specimen.
_Scuta_, externally very convex, with the growth-ridges extremely
prominent; basal margin sinuous, the middle portion being prominent;
this is best seen in young specimens (Pl. 3, fig. 2 _d_). Internally,
the articular ridge is broad and reflexed. The adductor ridge in the
upper part is almost confluent with the articular ridge; it runs down
to the most prominent point of the basal margin; in young specimens
it is sharp and prominent; in old specimens it is very blunt and
little prominent. There is a rather deep hollow for the lateral
depressor muscle. In young specimens there is a small, depending,
blunt tooth at the basi-tergal angle, which helps to make the basal
margin more deeply sinuous.
_Terga_, with the longitudinal furrow closed, except on the spur
itself, where it is open. The spur is moderately long and broad, but
varies in breadth; it is placed at rather less than its own width
from the basi-scutal angle; its lower end is obliquely rounded; the
basal margin on the opposite sides of the spur, together form a
nearly straight line. The whole valve is rather broad. The crests for
the tergal depressores are barely developed.
The _Compartments_ have rather large parietal tubes; the septa are
coarsely denticulated at their bases; the internal lamina is smooth,
except close to the basis. The _radii_ have their summits parallel
to the basis; their sutural edges are formed of rather thick septa,
which stand at an unusual distance apart from each other, and have
perfectly symmetrical, minute denticuli on each side. The interspaces
between the septa are filled up solidly to within a short distance of
the surface; but yet not so completely as in the following species,
and as in those in the succeeding sections of the genus; this is what
might have been expected from the close affinity of _B. stultus_ to
_B. Ajax_, in which latter the radii are still permeated by pores,
though smaller than is general in the species of our first section
(A). The _alæ_ have their summits extremely oblique, and their
sutural edges, I believe, smooth. _Basis_ porose, with an underlying,
finely-cancellated layer.
_Mouth_: labrum with six small teeth; mandibles with the 3d tooth
blunt; the 4th minute, and the 5th almost confluent with the inferior
angle. Maxillæ with the edge straight and simple. _Cirri_ partly
destroyed; on each segment of the sixth pair there were five pairs of
spines.
10. BALANUS CALCEOLUS. Pl. 3, fig. 3 _a_-3 _e_.
BALANUS CALCEOLUS KERATOPHYTO INVOLUTUS (?) _Ellis._ Phil. Trans.,
vol. 50 (1758), Tab. 34, fig. 19.
LEPAS CALCEOLUS (?) _Pallas._ Elench. Zooph., p. 198, (sine
descript.) (1766).
CONOPEA OVATA (?) _J. E. Gray._ Annals of Philosophy, vol. x, 1825.
_Parietes and basis porose. Scutum with the pit for the lateral
depressor muscle small and deep._
_Hab._--Attached to Gorgoniæ, West Coast of Africa. Tubicoreen, near
Madras, (Dr. Johnston), associated with _B. navicula_. Mediterranean
(?). Mus. Brit., Cuming, Stutchbury.
_Fossil._ Coralline Crag; Mus. S. Wood.
I must premise, with respect to the nomenclature of this and the three
following quite distinct species, that in the published descriptions
no allusion is made to any one of the characters by which alone
they can be distinguished: hence I have been guided by geographical
probabilities in assigning the specific name of _calceolus_ to the
present species, as Ellis's specimens came from the Mediterranean; and
that of _galeatus_ to the North American and West Indian specimens, as
Linnæus' original specimens (according to a statement by Spengler) came
from the West Indies. I have assigned new names to the two remaining
East Indian species. I may here add that Spengler ('Skrifter af
Naturhist.' 1 B, tab. 6, fig. 3, 1790) has described, under the name of
_B. cassis_, an allied form attached to the _Gorgonia placomus_ from
the seas of Norway; but I do not believe that it is the same with our
present species.
_General Appearance._--The degree of elongation of the shell in
its rostro-carinal axis varies considerably (3 _a_, 3 _b_): the
elongation is due to the production of the rostrum and of the
corresponding end of the basal cup. These two portions of the
shell always form together an angle, and sometimes an acute angle,
whereas in all the many specimens which I have seen, the carina and
the carinal end (or heel) of the basis together form a straight
line; yet I should not be surprised if this end of the shell was
sometimes produced. The surface of the shell is smooth, or sometimes
marked with very minute projecting points: it is almost always
covered by the horny bark of the Gorgonia. The colour is either dull
purplish-red or dull purple, with obscure longitudinal stripes,
and often more or less transversely banded with white. The rostrum
is either white or very feebly tinted, being always paler than the
rest of the shell: the radii are usually paler than the parietes,
and are sometimes white: the carinal end of the basal cup is tinted
of the same colour with, but rather paler than, the compartments.
The orifice is rather small compared to the shell, and nearly
heart-shaped. The carino-lateral compartments are about one-third of
the width of the lateral compartments. The shell is very strong, and
the sutures resist the action of boiling caustic potash. The largest
specimen which I have seen was .7 of an inch in extreme length, and
under .25 in extreme breadth.
_Structure of the shell and basis._--The parietes are permeated by
quite distinct pores,--a character sufficient by itself to separate
this from the following species; the longitudinal septa forming the
tubes are slightly denticulated at their bases. The radii have their
summits quite square, extending from apex to apex of the adjoining
compartments. The alæ have oblique summits. The sutural edges of
the radii have approximate septa, which are obscurely denticulated:
the interspaces are filled up solidly, so that the radii are not
porose. The basis is distinctly porose, by which this species can be
distinguished from _B. navicula_ and _cymbiformis_. The basis has a
deep furrow on the under side, from clasping the thin horny axis of
the Gorgonia: the basal point of the rostrum is also notched from the
same cause, and, as a consequence, its upper surface becomes slightly
furrowed along its whole length.
The _Scuta_ have an articular ridge but moderately prominent, and
only slightly reflexed; the basi-tergal corner is rounded off; there
is no adductor ridge; there is a small, rather deep, distinct pit
for the lateral depressor muscle. _Terga_; externally the surface is
considerably depressed in the line of the spur. The spur is between
half and one-third of the width of the valve: its lower end is square
and truncated, or in some degree rounded; it is sometimes (3 _e_)
dentated with a few, minute, sharp teeth. The articular ridge is but
slightly developed; the crests for the depressor muscle are very
feeble.
_Animal's body_ unknown.
11. BALANUS GALEATUS. Pl. 3, fig. 4 _a_-4 _c_.
LEPAS GALEATA (?) _Linnæus._ Mantissa altera Holmiæ, 1771.
CONOPEA ELONGATA. _Say._[90] Journal of Acad. Nat. Sci. Philadelphia,
vol. ii, part 2, p. 323, 1822.
[90] If I have assigned the specific title of _galeatus_ to
the wrong species, yet Say's name of _elongatus_ ought not
strictly to be admitted; as the _Lepas elongata_ of Gmelin is a
Balanus,--probably a variety of _Balanus crenatus_. I may add, that
as the _Lepas galeata_ of Schröter ('Einleitung in die Conch.'
&c.), was attached to a Gorgonia from the East Indies, it cannot be
our present species, but probably is one of the three other allied
species, which all occur in India.
_Parietes not porose; basis porose. Tergum, with the apex square,
caused by the great development of the articular ridge._
_Hab._--Charlestown, South Carolina; Florida; West Indies; Central
America; attached to Gorgoniæ; Mus. Brit., Agassiz, Cuming,
Stutchbury.
_General Appearance._--This and the two following species come so
close in general appearance to the last, that it will be quite
superfluous to do more than describe the few points of difference.
The shell and basis are generally quite as much elongated as in the
last species, and sometimes much more so, owing to the carinal end
(fig. 4 _a_), with the corresponding portion of the basal cup, being
produced like the rostral end, into a flattened, sharp point: I have
seen a specimen in this state .9 of an inch in length, and only .25
in breadth in the broadest part. In many specimens, however, the
shape is exactly as in _B. calceolus_; but the rostrum seems less
usually furrowed from clasping the stem of the Gorgonia. The colour
is paler, pinker, and more distinctly striped longitudinally than
in _B. calceolus_; I have, however, seen some not-striped, purple
specimens (and one transversely freckled with white) from the West
Indies. The parietes are strongly-ribbed internally, and are not
permeated by pores. The radii have their sutural edges crenated. The
basal cup is permeated by pores.
The _Scutum_ differs from that in the last species, only in the pit
for the lateral depressor muscle, being much shallower, and less
defined, and in the apex being truncated. The _Tergum_ is remarkable
from its broad, square, truncated summit, which underlies the whole
broad apex of the scutum: the square summit of the tergum is formed
by a great and peculiar development of the uppermost part of the
articular ridge. The spur is a little narrower than in _B. calceolus_.
_Mouth_: on the crest of the labrum there are two teeth on each side
of the central notch. The mandibles have five teeth, of which the two
lower are very small. The maxillæ show a trace of a notch under the
upper large pair of spines; near the inferior angle there are two
long spines. _Cirri_: in the first pair, one ramus is nearly twice as
long as the other: the segments are not very protuberant. There is
a sharp point at the dorsal basis of the penis. The branchiæ are of
moderate size, and plicated on one side.
12. BALANUS CYMBIFORMIS. Pl. 3, fig. 5 _a_, 5 _b_.
_Parietes and basis not porose. Scutum and Tergum with very small
articular ridges. Tergum broad, almost equilateral._
_Hab._--Attached to a Gorgonia, Tubicoreen, near Madras, (Dr.
Johnston). Hab. unknown, Mus. Cuming.
_General Appearance._--I have seen only two specimens, kindly sent me
by Dr. Johnston, and a single specimen in Mr. Cuming's collection.
In most points this species agrees with the two last species. The
shell (excepting the rostrum), and even the opercular valves in Mr.
Cuming's specimen were of a very fine purplish-red; in the other
specimens they were feebly tinted purple. The parietes are strongly
ribbed internally, and are not permeated by pores. The basal cup is
not porose, but its inner surface is ribbed in lines radiating from
the centre, and in both these respects this species differs from the
two foregoing. The Radii are rather narrow; they are paler coloured
than the parietes; they have their sutural edges plainly crenated.
The alæ have extremely oblique summits; the narrow rim added during
the diametric growth of the shell is white, the rest of the sheath
being, in Mr. Cuming's specimen, finely coloured like the parietes.
Basal diameter of the longer axis of the largest specimen, .4 of an
inch.
_Scutum_, rather narrow, with the basi-tergal corner much rounded
off; externally the lines of growth are little prominent. Internally,
the articular ridge is extremely little developed, and not at all
reflexed; there is no adductor ridge; there is a minute pit for the
lateral depressor muscle, placed almost on the edge of the valve.
The _Tergum_ is broad, forming (the spur being excepted) an almost
equilateral triangle. The articular ridge is remarkably little
prominent, and placed close to the scutal margin. The spur is nearly
half as broad as the valve, with its extremity or basal margin in one
case obliquely truncated, and in another case nearly square.
_Animal's body_ unknown.
13. BALANUS NAVICULA. Pl. 3, fig. 6 _a_-6 _d_.
_Parietes and basis not porose: carino-lateral compartments very
narrow, and of nearly the same width from top to bottom: radii with
their sutural edges smooth. Scutum externally striated longitudinally._
_Hab._--Attached to Gorgoniæ, Tubicoreen, Madras (associated with _B.
calceolus_), Dr. Johnston. Hab. unknown, Mus. Brit. and Darwin.
This is a very distinct form, though nearer to the foregoing than to
the other species. Its separation from the sub-genus Acasta is quite
artificial; its affinity to this sub-genus is shown by its weaker
shell, non-porose parietes and basis; by the radii having their sutural
edges smooth, and their summits not quite square; by the carino-lateral
compartments being very narrow; by the less elongated basis, not
furrowed, from not clasping the branches of the Gorgonia; and by the
longitudinally striated scuta; nevertheless, from the similar habits,
and from the graduated structure in the five foregoing species, it
cannot be removed out of the genus Balanus. I have seen three sets of
specimens of this species.
_General Appearance._--Shell, sometimes with the rostrum, and
sometimes with the carina, and corresponding portions of the basal
cup, elongated; but not, apparently, to so great a degree as in
the foregoing species; basis not furrowed, from not clasping the
branches of the Gorgonia. Colour pale blueish-purple, with the
radii whiter. The surface is studded with small calcareous points.
The carino-lateral compartments are very narrow, not more than one
tenth of the width of the lateral compartments; they are, moreover,
scarcely wider at the base than at the summit. The summits of the
radii are, apparently, a little oblique, or at least not so square as
in the foregoing species. The shell is not nearly so strong as in the
last three species; and the compartments separate by gentle force,
and from the action of caustic potash. The largest specimen was .4 of
an inch in basal diameter.
Internally, the parietes are not very strongly ribbed, or they are
almost smooth, and there are no pores. The basis is concave and
smooth within, and is not porose. The sutural edges of the radii are
quite smooth, or sometimes they exhibit, in the lower part, mere
traces of septa,--a character by itself sufficient to separate this
from the foregoing species. The alæ have oblique summits, and the
rather narrow portion added during the diametric growth of the shell,
is white.
The _Scutum_, externally (6 _d_), has raised striæ, radiating from
the apex; valve rather thick; internally, the articular ridge is but
slightly prominent, and its lower end is rounded off: the depression
for the lateral depressor muscle is slight; between this depression
and that for the adductor muscle, the surface of the valve is
prominent. _Tergum_, somewhat beaked; externally, the surface is
depressed in the line of the spur: the carino-basal margin slopes
towards the spur.
_Animal's body_ unknown.
_Section_ C.
_Parietes and basis permeated by pores. Radii not permeated by pores._
14. BALANUS TRIGONUS. Pl. 3, fig. 7 _a_-7 _f_.
_Parietes ribbed, mottled purplish-red; orifice broad, trigonal, hardly
toothed. Scutum thick, with from one to six longitudinal rows of little
pits. Tergum without a longitudinal furrow; spur truncated, fully one
third of width of valve._
_Hab._--Java; East-Indian Archipelago; Peru; West Columbia;
California; Sydney; New Zealand. Mus. Brit., Cuming, Stutchbury,
Dunker, &c.
_General Appearance._--Shell conical, generally depressed; orifice
broad, triangular, almost equilateral; walls coloured or only mottled
with purplish-pink, having either irregularly branching, or regular,
longitudinal ribs, which are generally white. The radii are pale
pink, or nearly white: the opercular valves have either their upper
parts, or nearly their whole surface, clouded with pinkish-purple:
the epidermis is not persistent: the walls are moderately strong: the
largest specimen was one inch, but generally full-grown specimens are
about half an inch in basal diameter.
The _Scuta_ have the lines of growth highly prominent. From one to
five or six rows (7 _b_, 7 _c_) of nearly circular, or transversely
oblong, deep pits, extend down the middle of the valve; rarely there
is not even one row; in this latter case, the valve is not striated
longitudinally. These little pits are caused by one or more deep
longitudinal furrows, crossed by the lines, or rather ridges, of
growth. In the same group of specimens, I have seen individuals with
three, five, and six rows; and even a few specimens with only one
row, or none at all. The outline of the valve is elongated, with the
apex slightly reflexed: the inner surface is protuberant, sometimes
to a remarkable, but variable degree. The articular ridge is not
very prominent, but it extends fully half-way down the valve, and
generally ends in a small free point. There is a short adductor
ridge, and a deep narrow pit or cleft for the lateral-depressor
muscle. _Terga_, externally smooth, flat, with scarcely a trace of
a longitudinal furrow; spur broad (7 _e_, 7 _f_), varying from half
to one third of the width of the valve, with the end truncated,
situated either near or quite close to the basi-scutal angle. The
crests for the depressor muscles are moderately well developed.
_Compartments._--The parietal tubes are, in their upper parts, filled
up solidly, without transverse septa. The radii generally have their
summits slightly oblique, and this is almost always the case with the
radii of the rostrum; the other radii sometimes extend from tip to
tip of the parietes, and are parallel to the basis; rarely the radii
are considerably oblique. The septa of the radii are very obscurely
denticulated, and the interspaces between them are filled up solidly.
The alæ have their sutural edges thin and smooth.
_Mouth._--Labrum with three teeth close together on each side of the
central notch: mandibles with four teeth, the fourth being small, the
fifth either absent or scarcely distinguishable from the inferior
angle: maxillæ without any notch, with the two lower spines rather
longer than the others. _Cirri_: In the first pair, one ramus is only
half the length of the other; in the second pair, both rami are short
and about equal in length; in the posterior pairs, the segments,
which are not protuberant, bear four pairs of spines, of which the
three lower pairs are short.
This species is widely-distributed, and where found seems to be common.
It is generally attached to shells of mollusca, but I have seen it also
attached to wood. I have found it associated with _B. tintinnabulum,
var. concinnus_, and _coccopoma_, with _B. psittacus_, _improvisus_ and
_amphitrite_, and with _Elminius modestus_.
Young specimens bear a considerable resemblance to certain young
varieties of _B. tintinnabulum_, and can indeed be distinguished
from them only by a careful examination of the opercular valves; for
it should be borne in mind, that in certain cases the scuta in _B.
tintinnabulum_ are pitted with little cavities. This species in some
respects is, I think, allied to _B. porcatus_, but it is far more
closely related to _B. spongicola_, and can be discriminated with
difficulty from certain varieties of this latter species. In Mr.
Cuming's collection, there is a group of small specimens, crowded
between some older specimens, which are remarkable from the shell being
oval in a transverse section,--from the smoothness of the walls,--and
from the absence of pits on the scuta; yet there could be no doubt that
these specimens belonged to our present species.
15. BALANUS SPONGICOLA. Pl. 4, fig. 1 _a_-1 _d_.
BALANUS SPONGICOLA. _Brown's_ Illustrations of the Conchology of
Great Britain (1827), pl. 7, fig. 6: 2d edit. (1844), pl. 53,
figs. 14-16.
_Parietes generally smooth, sometimes longitudinally folded; coloured
pink: orifice toothed. Scutum longitudinally striated. Tergum, with the
apex produced, without a longitudinal furrow; spur truncated, about one
third of width of valve._
_Var. with the walls slightly folded longitudinally._
_Hab._--South coast of England, and Tenby in South Wales, often
imbedded in sponges; attached also to shells and rocks in deep
water; Mus. Brit., Jeffreys. Algiers, on Mytili and Serpulæ, with
_B. perforatus_, Mus. Mac Andrew. Madeira, with _B. tulipiformis_,
Mus. Lowe. Lagulhas Bank, Cape of Good Hope, on detached kelp, with
_B. Capensis_, Mus. Sir J. Ross. Imbedded in sponge with _Acasta
spongites_, Mus. Bowerbank. _Var._ West Indies.
_Fossil_ in Coralline Crag, Mus. S. Wood.
_General Description._--Shell tubulo-conical; orifice of moderate
size, rather deeply toothed; colour dull pink, or purplish, or dark
flesh-colour; sometimes the radii are paler, sometimes of the same
colour with the parietes. Surface smooth when well preserved, having
transverse rows of minute spines. In the West Indian variety the
walls are slightly or much folded, but I will describe this form
separately. Size of largest specimen (Mus. Jeffreys), .6 of an inch
in basal diameter.
_Scutum_, with fine ridges radiating from the apex, and with the
lines of growth, crenated: internally, the articular ridge is small,
adductor ridge short and barely distinct: there is a rather deep
and narrow pit for the lateral depressor muscle. The whole valve
is much thinner than in _B. trigonus_, which in most respects it
closely resembles. _Tergum_, with the apex pinkish purple, produced
or beaked, but the beak is not needle-like, as in _B. psittacus_
and its allies, for the carinal margin is perfectly preserved up to
the tip. Externally the valve is nearly flat, for the longitudinal
furrow is very shallow. The spur is about one third of the width of
the valve; its lower end is abruptly truncated: in European specimens
(1 _b_) the whole basal margin, on the carinal side, slopes down to
the spur in a straight line, which, together with the sharpness and
production of the basi-scutal angle of the spur itself, gives to the
whole valve a peculiar appearance: in the specimen (1 _c_) from the
Lagulhas Bank, the basal margin on the carinal side is a little more
hollowed out, but it is quite impossible to doubt about the specific
identity of these specimens: in the West Indian variety (1 _d_) the
basal margin on the carinal side forms a distinct but obtuse angle
with the spur. In all cases the crests for the depressor muscles are
very feebly developed.
The _Compartments_ have their radii developed to a rather varying
degree, with their summits oblique; hence the orifice is toothed:
the sutural edges of the radii have their septa barely denticulated;
the sutural edges of the alæ are smooth. The basis, as with the
other species of this section, is permeated by pores; yet I found
one specimen, from the Cape of Good Hope, with the basis apparently
solid, thus offering a very singular anomaly. In the specimen
imbedded in sponge, the basis, as viewed externally, is concave;
whereas in Acasta, which always inhabits sponges, the basis is highly
convex or hemispherical.
The _Mouth_ and _Cirri_ resemble those of _B. trigonus_, and I can
point out no distinguishing character.
With respect to the variety from the West Indies, I have seen two
sets of specimens differing somewhat in external appearance, one set
attached to a coral from St. Vincent's, and another set to an Avicula
from an unknown locality; at first I described these specimens, with
some hesitation, as a distinct species, and I am very far from sure
whether this would not have been the more correct course, although
I am unable to point out any sufficient diagnostic characters. This
form differs from the ordinary _B. spongicola_, in the walls being
more rugged, stronger, and slightly or deeply folded longitudinally;
in this latter case the shell in external aspect differs much from
ordinary specimens of _B. spongicola_; but this is a variation so
common that I dare not place any reliance on it. The colour is more
purple; the summits of the radii perhaps rather less oblique. In the
scuta the only difference is that the articular ridge seems rather
longer, and the adductor ridge perhaps more prominent: in the terga,
as already remarked, the basal margin on the carinal side does not
slope so straight into the spur. These differences I consider all too
slight to be of specific value. The difficulty in determining the
nature of this variety is added to by its approach to _B. trigonus_
in all those points in which it departs from the ordinary _B.
spongicola_, so that for a short time I was even tempted to consider
both these species as varieties of one form. But until _B. trigonus_
is found with its scutum longitudinally striated, and with its tergum
beaked, it can hardly be confounded with _B. spongicola_; for it
should be observed that when in _B. trigonus_ the rows of little pits
disappear from the scuta, as sometimes happens, though rarely, yet
these valves do not become longitudinally striated.
_Balanus spongicola_ occurs, mingled with _B. tulipiformis_, in the
Mediterranean, and by the external characters of the shell alone
cannot be distinguished from that species; but the striated scuta and
beaked terga suffice to separate them. Again, this species, at the
Cape of Good Hope, occurs mingled with _B. Capensis_, and from the
non-striped young varieties of that species, it can, externally, be
distinguished only by the beak of the tergum not being sharp like a
needle. I have seen a single, perfectly characterised specimen, with
its opercular valves preserved, found by Mr. S. Wood in the Coralline
Crag at Sutton, mingled with _B. inclusus_.
16. BALANUS LÆVIS. Pl. 4, fig. 2-2 _g_.
BALANUS LÆVIS. _Bruguière._ Encyclop. Meth. (1789), Pl. 164, fig.
1.[91]
---- DISCORS. _Ranzani._ Mem. di Storia Nat., 1820, Tab. 3, figs. 9
to 13.
---- COQUIMBENSIS. _G. B. Sowerby_, in Darwin's Geology of South
America (1846), Tab. 11, fig. 7.
[91] M. Deshayes, in his descriptions of the plates, considers this
figure, I have no doubt erroneously, as that of _B. perforatus_, of
Bruguière. The _B. Coquimbensis_ of Sowerby, is a different species
from the _B. Coquimbensis_, of Chenu, 'Illust. Conch.,' tab. 6,
which latter is unknown to me.
_Shell covered by brown membrane, or naked and white or pale purple;
orifice small; radii very narrow. Scutum with one or two deep
longitudinal furrows._
_Var._ nitidus (fig. 2): _shell not covered by membrane, white or
pale purple: orifice but slightly toothed: scutum generally with two
furrows_. Hab.--Chile, as far south as Concepcion; Peru; California.
_Var._ Coquimbensis (fig. 2 _a_): _with the basal cup partly filled
up with thin, irregular, calcareous layers, making a cancellated
mass_. Fossil, and recent.
_Hab._--Strait of Magellan, ten to twenty fathoms, attached to
shells; often entirely surrounding pebbles, forming globular masses;
associated with _Verruca lævigata_. Chile and Peru, (generally _var.
nitidus_), often attached to _Balanus psittacus_. California. Very
common.
_Fossil_ in an ancient tertiary formation (middle bed) at Coquimbo,
Chile. In a recent deposit (_var. nitidus_) at the height of 1000
feet at Valparaiso; with Human remains at San Lorenzo, Callao, Peru.
I may premise that, having myself collected this species from the same
locality, the Strait of Magellan, where no allied species occurs,
attached to the same Mytilus and associated with the same Verruca, I
feel confident that it is the _B. lævis_ described by Bruguière; and
there can hardly be any doubt that it is the _B. discors_ of Ranzani.
With respect to the old tertiary specimens from Coquimbo, named _B.
Coquimbensis_ by Sowerby, they differ from the recent in no respect,
except in being considerably larger; and therefore I cannot consider
them specifically distinct. At first I was unwilling to believe that
the specimens with a single very broad longitudinal furrow, and those
with two rather broad, or with one narrow furrow, on their scuta,
could belong to the same species; but I soon found that all these
varieties occurred mingled together, and that they differed in no
other respect whatever. Generally, however, all the individuals in the
same cluster had the same variety of scutum,--thus adding one more to
the many instances amongst cirripedes of variations common to whole
groups of specimens. Still more unwilling was I to believe that _var.
nitidus_ and the common variety could belong to the same species. Their
general aspect is totally unlike: _var. nitidus_ has a smooth, clean,
naked shell, either white or pale purple, somewhat globulo-conical,
often with a nearly entire orifice; whereas the other common variety
generally has a more steeply conical shell, with a toothed orifice,
and is covered by a dirty brownish membrane. Moreover, though I have
seen hundreds of specimens from Tierra del Fuego, I have not seen one
specimen of _var. nitidus_, or even of an approach to it in appearance;
and, on the other hand, _var. nitidus_ is the common form in Chile and
Peru; though I have seen one or two specimens of the membrane-covered
variety from Valparaiso. Such facts strongly induced me to believe that
these forms were specifically distinct; but upon careful examination
I could find no other or more important differences than those just
specified. Some specimens from northern Chile are in an intermediate
condition; and from Concepcion, in the south of Chile, where the
climate approaches in character to that of the more southern parts of
the Continent, there are many specimens, in so intermediate a condition
that I know not whether or no to rank them under _var. nitidus_. Thus
I became convinced that these forms are only varieties. At Concepcion,
some few specimens are pale purple, and yet are wholly invested by
thick brown membrane, thus uniting the two extreme varieties. From
California I have seen both varieties, but I do not know which is
most common there. With respect to the great difference in aspect
between the specimens from northern Chile and Tierra del Fuego, we
shall hereafter see a strictly analogous case in _Balanus flosculus_.
Finally, I may add that _B. lævis_ seems to represent in the southern
hemisphere and on the west coast of North America, the _B. perforatus_
of Europe and Western Africa.
_General Appearance._--Shell conical, sometimes slightly globular;
surface smooth (that is, not folded), either naked, and in that case
white or pale purple, or covered by dirty yellowish-brown membrane.
Orifice small, more or less toothed, rarely exceeding one third of
the basal diameter. Radii very narrow, often not developed, the six
sutures forming in all cases deep and narrow clefts. The largest
recent, but much depressed, specimen which I have seen (from the
Strait of Magellan) was three fourths of an inch in basal diameter;
specimens growing congregated are often much elongated. I have seen
one with the basal cup between two and three times as deep as the
height of the compartments. Of the ancient tertiary specimens, the
largest had a diameter of three fourths of an inch, and a total
length of actually two inches (fig. 2 _a_); another of these fossils
had a basal cup in depth equalling four fifths of the entire length
of the shell and basis.
The _scutum_ has either one very broad and deep longitudinal furrow
(2 _b_), or two moderately broad and deep (2 _e_), or two narrow and
deep, or less frequently one narrow and inconspicuous longitudinal
furrow (2 _f_); rarely there is not one furrow; sometimes there are
none towards the apex, whilst furrows have been formed in the lower
part of the valve. In young specimens the furrows extend down to
the actual basal margin, but in old specimens they often fall short
of this, and, as a consequence, the furrows become crossed by one,
two, or three calcareous ridges, which ridges at successive periods
formed the basal margin of the valve. The external surface is covered
by yellow membrane; and fragments of several successive opercular
membranes are often attached to the zones of growth.
Internally the articular ridge is not very prominent, but is
remarkable (2 _c_, 2 _d_) from its lower point being produced into a
long, sharp, sub-cylindrical, free style (like the hinge of a common
gate), which is generally broken off in disarticulating the valve
from the tergum. The adductor ridge is either sharp and prominent
or blunt: it extends up the middle of the valve nearly to the apex,
and downwards it trends a little towards the occludent margin. The
pit for the lateral depressor muscle is minute but deep: the basal
margin is sometimes hollowed out under this pit. Sometimes there is
a distinct, but blunt ridge, caused by one of the furrows outside,
parallel to the adductor ridge, and placed between it and the little
pit for the lateral depressor; in this case, the basal margin, as
viewed internally, is rendered sinuous (2 _d_), as is best exhibited
in the great fossil specimens from Coquimbo.
_Tergum_ (2 _g_).--Spur of moderate length and breadth, with its
lower end obliquely truncated and rounded. The longitudinal furrow
has its edges somewhat folded in. The basal margin on the carinal
side of the spur is sometimes a little hollowed out. The crests for
the depressor muscles are well developed; but the corner of the valve
supporting them is extremely thin, and is often imperfectly calcified.
_Compartments._--The parietal tubes are not crossed by transverse
septa, but in their upper parts are filled up solidly. The radii are
always very narrow, with their summits oblique, though to a variable
degree: their sutural edges have fine and closely approximate septa,
with minute denticuli: the sutural edges are received in a furrow, on
the opposed compartment, of unusual depth; hence the lines of suture
run, in the lower part of the shell, almost exactly in the middle
between each two compartments. The alæ are added to above the level
of the opercular membrane.
The _Basis_ is often thick, with an underlying layer, largely
cancellated or honeycombed. When many specimens grow crowded
together, the basis is generally deeply cup-formed, or even
sub-cylindrical; and equals as much as four fifths of the length of
the entire shell. In such cases, in some few recent specimens, and
in all the large or even quarter-grown old tertiary specimens, but
not in the quite young fossil specimens, a structure is presented,
which I have not seen in any other Cirripede, namely, the basis (Pl.
4, fig. 2 _a_) is filled up for one third, or even for more than
half its depth, by successive, separate, calcareous, transverse
layers or septa. It would appear as if the basal cup had grown too
large for the animal's body, and so required filling up. The layers
are thin and fragile; a single layer never stretches across the
whole shell; each is irregularly mammillated or blistered, with the
convex surfaces generally directed upwards; the layers are furnished
on their under sides with little pillars and short ridges, resting
on the layers beneath; it rarely happens that the supports of one
layer lie directly over those of another, though this is sometimes
the case. In a vertical section, the mass formed by these irregular
layers has a coarsely cancellated structure. This structure, although
confined to this one Cirripede, is not so anomalous as might at first
be thought, for in most species of the genus, each time that the
circumference of the basis is added to, an excessively thin calcified
film is thrown down over its whole inner surface; and in any of these
species, if the films had been formed thicker and had rested only
on certain points, instead of over the whole underlying layer, the
cancellated structure above described would have been produced.
_Mouth_: the labrum is either destitute of teeth, or has two or three
very minute teeth. The palpi have a tuft of very long spines at their
ends. The third tooth of the mandibles is thicker and larger than
the two upper ones. The maxillæ have either a nearly straight edge,
or the inferior corner is obliquely truncated, and projects much
beyond the rest of the edge. In the _Cirri_, none of the segments are
very protuberant: in the first pair, one ramus is nearly twice as
long as the other: in the posterior pairs, the segments are not much
elongated, but each supports seven pairs of spines.
_Var. nitidus_: with respect to this variety I have little to add
to my preliminary remarks on its peculiar appearance, owing to its
smooth, naked condition, and pure white or pale purple colour. This
colour, when examined through a lens, is seen to consist of very
fine longitudinal stripes; and is produced by the calcareous matter
within the longitudinal parietal pores being thus coloured. Generally
the scuta have two longitudinal furrows; but I have seen a scutum
of one perfectly characterised specimen with only a single broad
furrow, like that which frequently occurs in the membrane-covered
variety. _Var. Coquimbensis_, as before stated, differs only in its
greater size: the scutum, in the one specimen examined, had two
broad longitudinal furrows; neither it, nor the tergum differed from
certain varieties now found on the coast of Chile.
17. BALANUS PERFORATUS. Pl. 5, fig. 1 _a_-1 _d_; Pl. 4, fig. 3 _a_-3
_c_.
BALANUS PERFORATUS. _Bruguière._ Encyclop. Meth., 1789, Tab. 164,
fig. 12 _infra_.
LEPAS ANGUSTA. _Gmelin._ Syst. Naturæ, 1789.
---- ORE ANGUSTIORE. _Chemnitz._ Vol. viii, Tab. 98, fig. 835.
BALANUS CORNUBIENSIS CONICO ORE MINORE. _Ellis._ Phil. Trans. vol.
50, 1758, Tab. 34, fig. 16.
LEPAS BALANUS ET FISTULOSUS. _Poli._ Test. Siciliæ (1795), Tab. 4,
fig. 5, Tab. 6, fig. 1.
BALANUS COMMUNIS. _Pulteney._ Dorset Catalogue, 1799.
---- ---- _Montagu._ Test. Brit., 1803.
LEPAS ANGUSTATA. _Wood._ General Conchology, 1815, Pl. 6, fig. 5.
BALANUS CRANCHII. _Leach_ (!). (_B. Blainvillii_ in Tab.) Encyclop.
Brit. Suppl., vol. iii, 1824.
---- ---- _Brown._ Illust. Conch., 1827, Pl. 7, fig. 9, 10, and 2d
Edit., Pl. 53, fig. 9-12.
---- PERFORATUS. _Chenu._ Illust. Conch., Tab. 3, fig. 9, Tab. 6,
fig. 15.[92]
[92] I have very little doubt regarding any of these references:
I have no means of ascertaining the priority, within the same
year, of Gmelin and Bruguière, but have given it to the latter,
as _perforatus_ is much the best known specific name. English
conchologists seem generally to suppose that the _B. communis_ of
Pulteney and Montagu is the _B. porcatus_ of this work; but I have
not the smallest doubt that I have given it rightly as a synonym
of the present species; the indistinctness of the compartments,
the multitude of fine ridges, the smallness of the orifice, the
longitudinal furrow on the terga, the colour, size, and habitat,
all given by Pulteney or Montagu, will agree with no other British
species. The _Lepas balanus_ of Poli, which is certainly a synonym
of our present species, has been erroneously considered by several
authors to be the same with the _L. balanoides_ of Poli, which
latter undoubtedly is the _B. amphitrite_ of this work.
_Shell pale purple, or white, or dirty ash-colour; smooth, or, from
being corroded, finely ribbed longitudinally; sheath purple; orifice
generally small; radii generally narrow or absent. Scutum, internally,
with a short minute ridge, parallel and close under the prominent
adductor ridge. Tergum with the apex somewhat produced._
_Var._ angustus (_Gmelin_) Pl. 5, fig. 1 _a_: _pale dull purple
or white; orifice small or of moderate size; radii very narrow or
moderately wide, white or pale purple, with oblique summits_.
_Var._ Cranchii (_Leach_) Pl. 5, fig. 1 _b_: _corroded, covered with
fine longitudinal ridges owing to the exposed, filled-up, parietal
tubes; dark dirty ash-colour, with a tinge of purple: radii not
developed, or very narrow with oblique summits; orifice small_.
_Var._ fistulosus (_Poli_) Pl. 5, fig. 1 _d_: _shell cylindrical,
white or dull purple; orifice of moderate size or small; basis deeply
cup-formed_.
_Var._ mirabilis, Pl. 5, fig. 1 _c_: _bright purple; radii white,
very broad, with their summits parallel to the basis; orifice entire,
large_.
_Hab._--Southern shores of England; South Wales; Mediterranean;
Western Africa, southward to Loanda, in 9° S.; West Indies (?).
Generally adhering to rocks at a low tidal level; in one case
attached to the floating _Lepas Hillii_, Mus. Jeffreys.
This is a well-marked species, and in its essential characters does
not vary much; but owing to the shell being almost as often white as
purple,--to its being remarkably subject to disintegration,--to its
often becoming cylindrical,--to the radii being either not at all, or
slightly, or moderately, or largely developed, and consequently to the
orifice of the shell varying in size, the general external appearance
of the different varieties is singularly diversified; but when a series
of specimens is examined, it is easy to see how one form passes into
another.
_General Appearance._--Shell conical, with the orifice oval,
unusually small, being generally only from one third to half of the
basal diameter; sometimes moderately large; in one single instance as
wide as the basis. Radii, often represented by mere lineal fissures,
or they are narrow, or sometimes moderately wide. Colour pale, dull
purple, sometimes lilac, often passing into a dead pure white: the
same individual will occasionally have one part of its shell white,
and another purple: the purple tint almost invariably is nearly
uniform, or not in stripes. The radii are generally white, when
the whole shell is purple, but sometimes they are pale purple: the
sheath is apparently always coloured of a fine claret-purple, with
the triangular portion of the alæ, added during diametric growth,
generally white, but sometimes purple. The surface is quite smooth,
but very often, especially on the shores of England, whole groups of
specimens (excepting the very young ones,) have had the outer lamina
of the parietes entirely corroded and removed; in this case the shell
assumes a dirty, more or less dark, ash-colour, feebly tinted with
purple, and the whole surface, owing to the exposure of the solidly
filled-up parietal tubes, becomes finely striated, or covered with
very narrow, longitudinal ridges. When specimens are crowded together
they often become cylindrical, and much elongated, owing to the
basis becoming deeply cup-formed: I have seen specimens, half an
inch in diameter in the widest part, one inch and a half in height,
the walls forming only a third of this. The largest specimen which I
have seen (from the southern shores of England) had a basal diameter
of 1.2 of an inch; some very steeply conical specimens were .9 of an
inch in height, and .8 in basal diameter.
_Scuta_, externally, slightly convex; growth-ridges approximate,
moderately prominent. Internally (Pl. 4, fig. 3 _a_) the articular
ridge is moderately developed, with the lower end produced downwards
into a freely depending, flattened style, somewhat variable in
size, but not so long as in _B. lævis_, and easily broken in
disarticulating the valves. The adductor ridge is very prominent,
running from almost the apex of the valve, close to the articular
ridge, to near the basal margin. The basi-tergal portion of the
valve is converted by the adductor ridge into a rather deep cavity,
within which there is a short, sharp, and minute ridge, close and
parallel to the adductor ridge, and bounding the impression left by
the lateral depressor muscle: this insignificant ridge was present
in every specimen; it occurs only in very few other species, as in
_B. nubilus_ and _cariosus_. The thickness of the valve sometimes
varies a little, and when thick the adductor ridge does not appear
quite so prominent. _Tergum_, with the apex moderately beaked and
produced; beak triangular in section, coloured dark purple, as is
the upper internal surface of the valve; the longitudinal furrow is
deep, and has its edges folded in, and even quite closed. The spur
is moderately long and narrow; but its width varies a little (Pl. 4,
fig. 3 _b_, 3 _c_), and consequently it stands at either rather above
or at twice its own breadth from the basi-scutal angle: its lower
end is either bluntly pointed or square, and generally is feebly
toothed on the under-side. The basal margin of the valve generally
slopes a little, on both sides, towards the spur. Internally, the
scutal margin is but slightly inflected: the articular ridge is but
slightly prominent, and but little curved; in the upper part of the
valve there are generally several very minute ridges, parallel to the
articular ridge, on the side towards the scutum. The internal surface
of the spur itself is sometimes concave. The crests for the carinal
depressor muscle are barely developed. It may here be mentioned that
on the opercular membrane many long spines stand rudely arranged in
rows.
_Parietes_: the parietal tubes have not transverse septa; but
are solidly filled up in their upper parts by dark-purple layers
of shell. The _radii_, as already stated, are either not at all
developed, or are extremely or only moderately narrow, with their
summits more or less oblique: in Mr. Cuming's collection, however,
there is an unique specimen, _var. mirabilis_ (Pl. 5, fig. 1
_c_) with the aperture of the shell as wide as the basis, with
bright purple parietes, and white, very broad radii, having their
summits parallel to the basis. The septa of the radii are finely
denticulated, and the interspaces are filled up solidly. The _alæ_
have very oblique summits, and their edges are finely crenated.
_Basis_, flat, or deeply cup-shaped; there is often an underlying,
coarsely-cancellated layer.
_Mouth_: labrum finely hairy, but without any teeth; mandibles, with
the 4th tooth small; the 5th confluent, with the sometimes smooth,
sometimes pectinated inferior angle. Maxillæ, rather broad, with a
slight notch under the upper pair of spines. _Cirri_, first pair,
with one ramus, having 29 segments, and above one third longer than
the shorter ramus, having 17 segments; these latter segments are
remarkable by the extent to which their upper front surfaces are
laterally produced into projections, twice as long as the breadth of
that portion of the segment which is articulated to the adjoining
segment. These projections have a double row of serrated spines on
their upper edge, and a beautiful radiating bundle at the end; the
projections decrease in length, both in the upper and lower segments.
The second cirrus (Pl. 29, fig. 4) has the segments (13 in number, in
the same individual with the segments above enumerated) of both rami
produced in the same singular manner as in the first pair. The third
pair have only inverted conical segments, coloured darker purple than
the other cirri. The sixth pair had in the same individual 31 or 32
segments, and therefore one or two more than in the longer ramus of
the first pair.[93] The segments in the posterior cirri have their
anterior faces shield-shaped, and bear 6 or 7 pairs of spines, with
some minute intermediate spines. There is the usual point at the
dorsal basis of the penis.
[93] Under the Genus (p. 190) I have given the numbers of the
segments in the cirri of this species at successive ages.
_Range._--This species is common on the southern shores of England
and in the Channel Islands: the largest specimens which I have seen
came from these quarters. The most northern point whence I have
seen specimens, is Tenby, in South Wales. This species is common
throughout the Mediterranean; I have seen specimens from Malaga,
Sicily, Algiers, and Smyrna; thence it ranges down the western
coast of Africa, as far south as the Gambia and Loanda, in 9° south
latitude. I believe British specimens are more often corroded than
those from further south. Amongst some old, ill-kept specimens in a
box in the British Museum, marked "Kingston, Jamaica," there were
some of this species: also I received some specimens, marked "S.
America," from Mr. G. B. Sowerby: again, Ellis, in Phil. Trans., vol.
50, part 11, gives a figure (Tab. 34, fig. 15) of some specimens from
the West Indies, which I believe to be _B. perforatus_: hence, it is
in some degree probable that this species, like _B. tintinnabulum_,
and _amphitrite_, and _improvisus_, may be found on both sides of the
equatorial Atlantic. _Balanus perforatus_ is attached, together with
_B. tulipiformis_, _trigonus_, _amphitrite_, _Chthamalus stellatus_,
and _Pollicipes cornucopia_, usually to rocks, near the lower limit
of the tidal level; but I believe, from specimens kindly sent me
by Mr. Mac Andrew, that it is frequently obtained by dredging;
one specimen was even marked 30 fathoms. According to Poli, it is
sometimes attached to the bottoms of vessels; and I have seen a
specimen adhering to the floating _Lepas Hillii_.
_Affinities._--This is a distinct species, closely allied to no other
species, but comes nearest to _B. lævis_, which is its representative
in Southern America, and on the whole west coast of that continent.
It is allied to that species, and differs from most other species,
in the general form of the shell, its small orifice, narrow radii,
and often deeply cup-formed basis. It agrees to a certain extent in
the colouring, though the purple here is much more prevalent, and
is not confined to the shelly matter filling up the parietal tubes.
It agrees with that species in the general structure of the scutum;
but the two or three deep, longitudinal furrows are here absent; and
the minute ridge, parallel to and almost under the adductor ridge,
is a peculiarity confined to this and very few species in the genus.
The terga differ from those of _B. lævis_, chiefly in the spur
being narrower, and in the apex being beaked. Lastly, the highly
protuberant segments of the one ramus in the first cirrus, and of
both rami in the second pair, are here remarkable. With regard to the
varieties, I have nothing to add to their short diagnostic characters
above given.
18. BALANUS CONCAVUS. Pl. 4, fig. 4 _a_-4 _e_.
BALANUS CONCAVUS. _Bronn._ Italiens Tertiär-Gebilde (1831) et Lethæa
Geognostica, b. ii, s. 1155 (1838), Tab. 36, fig. 12.[94]
---- CYLINDRACEUS, _var._ c. _Lamarck._ Animaux sans Vertèbres
(1818).
LEPAS TINTINNABULUM. _Brocchi._ Conchologia Sub-Appen., t. ii, p. 597
(1814).
[94] I suspect that _B. pustularis_, _miser_, and _zonarius_, all
figured by Münster, in his 'Beiträge,' b. iii, Tab. 6, may be this
species.
_Shell longitudinally striped with white and pink; or dull purple;
sometimes wholly white. Scutum finely striated longitudinally;
internally, adductor ridge very or moderately prominent._
_Hab._--Panama; Peru; S. Pedro in California; Philippine Archipelago;
Australia. Mus. Brit., Cuming, Stutchbury, Aug. Gould.
_Fossil_ in Coralline Crag, England; Mus. Brit., S. Wood, Bowerbank,
Lyell, J. de C. Sowerby, Tennant. Sub-Appennine formations, near
Turin, Asti, Colle in Tuscany, Mus. Greenough, &c. Tertiary beds,
near Lisbon, Mus. D. Sharpe and Smith. Bordeaux (?) Mus. Lyell.
Tertiary beds, Williamsburg; and Evergreen, Virginia, Mus. Lyell.
Maryland, Mus. Krantz. Recent formations[95] near Callao, Peru, Mus.
Darwin. Red Crag (Sutton) Mus. S. Wood.
[95] I procured this specimen from the Island of S. Lorenzo, off
Callao; it was imbedded, together with seventeen species of recent
shells and with human remains, at the height of eighty-five feet.
This species has caused me much trouble. Looking first to the recent
specimens, I examined several from Panama and California, which,
though differing greatly in colour, resembled each other in their scuta
having the adductor ridge extremely prominent, and in having (Pl. 4,
fig. 4 _a_), an almost tubular cavity for the attachment of the lateral
depressor muscle,--characters which at first appeared of high specific
value; but I soon found other specimens from Panama in which these
peculiarities were barely developed. I then examined a single specimen
from the Philippine Archipelago, resembling in external appearance one
of the Panama varieties, but differing in the scuta being externally
strongly denticulated in lines instead of being merely striated,--in
the adductor ridge being far less prominent,--and in the spur of the
tergum being broader and more truncated; I therefore considered this
as a distinct species. I then examined a single white rugged specimen
from the coast of Peru, which differed from the Philippine specimen
in the shape of the well-defined denticulations on the scuta, and in
some other trifling respects, and in the segments of the posterior
cirri bearing a greater number of spines; with considerable doubt, I
also named this as distinct. But when I came to examine a large series
of fossil specimens from the Coralline Crag of England, from northern
Italy, from Portugal, and from the southern United States, I at once
discovered that the form of the denticuli on the scuta was a quite
worthless character,--that in young specimens the scuta were only
striated,--that the prominence of the adductor scutorum ridge and the
depth of the cavity for the lateral depressor muscle varied much (as
in the case of the recent specimens), owing apparently to the varying
thickness of the valve,--that in the terga the spur varied considerably
in length and breadth, the latter character being in part determined
by the varying extent to which the edges of the longitudinal furrow
are folded in,--and lastly, that in young specimens the basal end of
the spur is much more abruptly truncated than in the old. Hence I have
been compelled to throw all these forms, originally considered by me
as specifically distinct, into one species. I must repeat that this
considerable variation in the prominence of the adductor ridge, and in
the depth of the pit for the lateral depressor muscle--the pit in some
cases becoming even tubular--is a very unusual circumstance.
With respect to the fossil specimens[96] from the above-stated several
distant localities, I consider them as certainly belonging to one
species, though varying considerably in several points of structure.
When compared with the recent specimens, they differ from them in often
attaining a considerably larger size; in the parietes being often, but
not always, longitudinally ribbed; and in the radii often having more
oblique summits. On the other hand, considering the many points of
identity between the fossil and the recent specimen, I have concluded,
without much doubt, that they ought all to be classed together. I may
remark that, in the Coralline Crag specimens, the spur of the tergum
(Pl. 4, fig. 4 _d_), is unusually long and narrow; it is broader and
shorter in the Italian specimens (4 _e_), and variable in this respect,
in the United States specimens; the scuta of the Lisbon specimens
are remarkable for the greater prominence of the adductor ridge, and
for the depth of the lateral depressor cavity. Some of the specimens
from all the several localities are identical with the recent ones
from the coast of Peru. The walls of the shell in the Coralline Crag
specimens, are generally ribbed longitudinally. I have entered into
the above particulars, on account of, in the first place, its offering
an excellent example how hopeless it is in most cases to make out the
species of this difficult genus without a large series of specimens;
secondly, as showing how the characters alter with age; and thirdly, as
a good instance of the amount of variation which seems especially to
occur in most of the species which have very extensive ranges.
[96] These will be fully illustrated in the monograph on the Fossil
Balanidæ, to be published by the Palæontographical Society.
Some of the pink-striped Panama varieties, though having a somewhat
different aspect, can be distinguished from certain varieties of _B.
amphitrite_ only by their scuta being longitudinally striated,--a
character in this species variable in degree, and in most cases of
very little value. Some of the other recent varieties are sufficiently
distinct from _B. amphitrite_; and the great fossil Coralline Crag
specimens, which stand at the opposite end of the series of varieties,
with their ribbed walls, very oblique radii, and coarsely striated
scuta, are extremely unlike _B. amphitrite_. With respect to the
nomenclature of the present species, I have little doubt that I have
properly identified the Italian fossil specimens with _B. concavus_ of
Bronn, who has given a very good figure of this species in his 'Lethæa
Geognostica;' it must, however, be confessed that the longitudinal
striæ on the scuta are not there represented. Considering the large
size and frequency of this species in Europe and in the United States,
it has probably received several other names, besides the two incorrect
synonyms, quoted at the head of this description. I should add that
the true _B. cylindraceus_ (not _var._ C) of Lamarck, according to the
plate given by Chenu in his 'Illust. Conch.,' is the _B. psittacus_ of
South America. I have seen in collections specimens of _B. concavus_
labelled as _B. tulipa_ of Poli (_B. tulipiformis_ of this work),--a
very natural mistake, without the opercular valves be carefully
examined.
_General Appearance._--Shell conical, often steeply conical; orifice
rather small, with the radii narrow, and generally in the fossil
specimens very oblique; surface generally smooth, sometimes rugged,
and in the coralline crag specimens generally ribbed longitudinally,
the ribs being narrow. Colour various, either dull reddish-purple
with narrow nearly white, or wider dark longitudinal bands; or,
again, pale rosy-pink with broad white bands; or lastly, wholly
white. The radii are either darker or paler than the parietes. The
opercular valves are either dark purple or nearly white. Pale pink
and white stripes are visible on some of the Italian and Portuguese
tertiary specimens; and in most of the fossils the sheath is tinged
dull red.
_Dimensions._--The largest actually recent specimen which I have
seen, from the Philippine Archipelago, had a basal diameter of 1.2
of an inch; the Peruvian pleistocene specimen is 1.7 in diameter;
specimens from the crag and from the Italian deposits, however,
sometimes slightly exceed two inches in basal diameter, and three in
height.
_Scuta_: these in young and moderately-sized specimens are striated,
sometimes very faintly, but generally plainly, causing the lines of
growth to be beaded; but often, in large and half-grown specimens,
the lines of growth are extremely prominent, and being intersected by
the radiating striæ, are converted into little teeth. As the striæ
often run in pairs, the little teeth frequently stand in pairs,
or broader teeth have a little notch on their summits, bearing
a minute tuft of spines. In very old and large specimens, the
prominent lines of growth are generally simply intersected by deep
and narrow radiating striæ. In one case, a single zone of growth in
one valve was quite smooth, whilst the zones above and below were
denticulated. The valve varies in thickness, which I think influences
the prominence of the lines of growth and the depth of the striæ.
These striæ often affect the internal surface of the basal margin,
making it bluntly toothed. The articular ridge is rather small,
and moderately reflexed: the adductor ridge (as already stated,)
varies remarkably; in most of the Panama specimens, it is extremely
prominent, and extends down to near the basal margin; in other
specimens it is but slightly prominent, especially in some of the
fossil specimens from Virginia. The cavity for the lateral depressor,
also, varies greatly; it is often bounded on the side towards the
occludent margin by a very slight straight ridge, which occasionally
folds a little over, making almost a tube; this, at first, I thought
an excellent specific character, but far from this being the case,
the cavity often becomes wide, quite open, and shallow.
_Terga_, very slightly beaked; the surface towards the carinal end of
the valve, in some of the fossil specimens, is very slightly striated
longitudinally. There is either a slight depression, or more commonly
a deep longitudinal furrow, with the edges folded in and touching
each other, extending down the valve to the spur, and causing the
latter to vary in width relatively to its length. When the furrow
is closed in, the spur is about one fourth of the entire width of
the valve, and has its lower end obliquely rounded, and stands at
about its own width from the basi-scutal angle: when there is only
a slight depression and no furrow (as is always the case with young
specimens), the spur is broader, equalling one third of the width
of the valve, with its lower end almost truncated, and standing at
about half its own width from the basi-scutal angle. But the absolute
length of the spur, also, varies considerably; it is often very long,
compared to the whole valve. The basal margin on the carinal side
is sometimes slightly hollowed out; when the furrow is closed, this
latter side slopes towards the spur. Internally, the articular ridge
and crests for the tergal depressor muscle are moderately prominent.
_Parietes_, the longitudinal septa sometimes stand near each
other, making the parietal pores small. The _radii_ have oblique
summits, but to a variable degree; their septa are unusually fine,
and are denticulated on their lower sides; the interspaces are
filled up solidly. The _alæ_ have their summits very oblique,
with their sutural edges nearly or quite smooth. In most of the
fossil specimens, and slightly in some of the recent specimens, the
surface of the sheath presents an unusual character, in a narrow,
longitudinal, slightly raised border, running along the sutures, on
the carinal side of each compartment.
_Basis_ thin, porose; sometimes with an underlaying cancellated layer.
_Mouth_: labrum with six teeth: mandibles with the fourth and fifth
teeth small, either sharp, or blunt: maxillæ with a straight edge,
or with the inferior part slightly prominent. _Cirri_ with the rami
of the first pair unequal by four or five segments: the segments in
the shorter ramus are extremely protuberant. The segments in the
second cirrus only moderately protuberant: but all the specimens were
in bad condition, and it appeared as if, in the Panama specimens,
the segments of the second cirrus were more protuberant than in the
Philippine Island specimens. In the posterior cirri there are from
three to five pairs of spines on each segment: even amongst the
Panama specimens some had three and some four pairs, and a white
Panama specimen had five pairs of spines.
All the recent specimens which I have seen, were, with one exception,
attached to various shells and crabs, and to each other. The Peruvian
specimen was associated with _B. flosculus_. The tertiary specimens are
often congregated together into great masses. Including the recent and
fossil specimens, this species encircles the globe. During the miocene
period it seems to have been the commonest existing sessile cirripede;
now, it does not appear to be common, excepting, perhaps, at Panama:
Mr. Cuming procured only one specimen from the Philippine archipelago.
19. BALANUS AMPHITRITE. Pl. 5, fig. 2 _a_-2 _o_.
LEPAS RADIATA. _Wood's_ General Conchology (1815), Pl. 7, fig. 7.
---- MINOR? _Wood's_ General Conchology (1815), Pl. 7, fig. 6.
---- BALANOIDES. _Poli._ Testacea utriusque Siciliæ (1795), Tab. 5.
BALANUS BALANOIDES. _Risso._ Hist. Nat. de l'Europe Merid., tom. iv,
1826.
_Shell longitudinally striped with purple or pink; sometimes with the
stripes confluent; sometimes wholly white. Scutum internally with a
prominent broad adductor ridge._
_Var._ (1) communis: (2 _e_, 2 _h_, 2 _l_,) _nearly white, with
pale or dark violet-coloured longitudinal stripes: epidermis rarely
persistent: shell either thin or thick: radii white or freckled
with reddish mahogany colour, with their summits either oblique,
sometimes in a high degree, or nearly parallel to the basis: basal
point of spur of the tergum either square or bluntly pointed_. Hab.
Mediterranean, W. Indies, S. Africa, Philippine Archipelago, New
South Wales.
_Var._ (2) venustus: (2 _a_,) _white or pale pink, with narrow bright
pink, or broad pinkish-purple stripes; orifice either much dentated
or nearly entire. Tergum with the carinal half of the basal margin
sometimes much hollowed out_. Hab. W. and S. Africa, Ceylon.
_Var._ (3) pallidus: (2 _c_, 2 _k_,) _white, with or without a
yellowish persistent epidermis; sometimes with the edges of the
compartments tinted purple: radii moderately oblique: tergum
generally narrow, with the spur sharp, and the basal margin on its
carinal side much hollowed out_. Hab. W. Africa, Madagascar, Red Sea.
_Var._ (4) niveus: (2 _f_,) _white, with longitudinal hyaline lines;
epidermis not persistent_. Hab. W. Indies, Florida, S. Africa, &c.
_Var._ (5) modestus: _upper part of shell white, lower part uniform
blueish-gray, opercular valves as in_ Var. (1). Hab. unknown.
_Var._ (6) Stutsburi: (2 _d_, 2 _i_, 2 _m_, 2 _n_, 2 _o_,) _white,
with or without pinkish-purple stripes, which are often confluent,
rendering the lower part of the shell of a uniform purplish tint;
epidermis persistent: radii very narrow: tergum narrow, spur sharp,
varying in form and in exact position; carinal margin sometimes
highly protuberant; basal margin on the carinal side of the spur
generally, but not invariably, much hollowed out_. Hab. West Africa.
_Var._ (7) obscurus: (Pl. 5, fig. 2 _g_,) _with narrow, approximate,
obscure and often almost confluent, slaty, or pale purplish-brown,
or dark slate-coloured stripes_. Hab. West Indies, Australia, and
unknown.
_Var._ (8) variegatus: _with narrow, approximate, dusky,
claret-coloured stripes, transversely freckled with white; shell
conical; walls very thin: scutum with the adductor ridge small_. Hab.
New Zealand.
_Var._ (9) (_an. spec._?) cirratus: (fig. 2 _b_,) _shell very pale
purplish-brown, with faint, more or less plain longitudinal stripes,
transversely freckled with white; walls thin: scuta with the lines of
growth beaded: basis, in specimens growing in groups, irregularly cup
formed: maxillæ with the inferior corner extremely prominent_. Hab.
Mouth of Indus, Australia, Philippine Archipelago.
_Hab._--Warmer temperate and tropical seas; extremely common;
Mediterranean, Smyrna, Sicily, Coast of Portugal; West Coast of
Africa, River Gambia, West Indies, Demerara, Natal, Madagascar, Red
Sea, Mouth of the Indus, Ceylon, Philippine Archipelago, East Indian
Archipelago, Pacific Ocean, east coast of Australia, New Zealand;
extremely common on ships' bottoms; often attached to floating
timber, canes, &c.; often associated with _B. tintinnabulum_;
attached to pebbles and various shells.
With respect to the nomenclature of this extremely common species,
which is widely distributed in all the warmer seas (excepting, as
far as I have seen, on the west coast of America), there is some
difficulty. I have no doubt that it is the _Lepas radiata_ of Wood
(1815), but Bruguière, in 1789, gave this same name to a Balanus which
he had not seen, but which is figured in Chemnitz, Tab. 59, fig. 842.
I should have thought that this also had been the present species,
but Spengler, in describing (Skrifter af Naturhist. Selskabet i, B.
1790) this individual specimen, which he calls _L. purpurea_, states
that it is 13 lines in basal diameter; now this is a size which is
never acquired by _B. amphitrite_; and the description, habits, and
size, would apply equally well to the species which I have called _B.
amaryllis_; but when no notice is taken of such points of importance,
as whether the walls are permeated by pores, whether the radii are
smooth-edged, whether the scuta are striated, it is impossible to
identify with any approach to certainty sessile Cirripedes; and
the names given ought, in my opinion, to carry little weight with
them. With respect to Lamarck's _Balanus radiatus_ (1818), the
synonyms quoted exhibit some great and inextricable confusion. The
_B. radiatus_, again, of Risso, is a fossil and apparently distinct
species. There can be no doubt that the present species is the _Lepas
balanoides_ of Poli, (and of several authors who have followed him),
and equally little doubt that the present species is not the true _L.
balanoides_ of Linnæus, which has a membranous basis, and which I have
not seen from the Mediterranean. Under these circumstances I have
concluded that less confusion would be caused by giving a new name to
this species than by taking that of Wood, which ought not to have been
used by him, considering Bruguière's previous adoption of it.
Under the head of _B. tintinnabulum_ I have alluded to the great
variation of _B. amphitrite_, which consists not only in a vast
diversity in the colouring and in the general aspect, but likewise
in the degree of obliquity of the summits of the radii, in the form
of the terga, and slightly in that of the scuta. In order to show
that it has not been from indolence that I have put so many forms
together, I may state that I had already named and fully described
in detail eight of the following forms as species, when I became
finally convinced that they were only varieties: it would require at
least thirty figures, which I have not the power to give, fully to
illustrate the transitional forms. As with _B. tintinnabulum_, the
deception is wonderfully enhanced by whole groups of specimens from the
same locality exactly resembling each other, and sometimes differing
from other groups attached to the very same object. If a person were
to get together only some fifty or sixty specimens from only half a
dozen different localities, he would almost certainly come to the same
conclusion, as I at first did, that several of the varieties are true
species; but when he gets several hundred specimens from all quarters
of the globe, he will find, to his trouble and vexation, that character
after character fails and blends away by insensible degrees, and he
will be led, as the more prudent course, to include, as I have done,
and I hope rightly, all under one specific name. I have experienced
more doubt regarding the last variety, _cirratus_, than on any other,
on account of its peculiar colouring, and from the basis being often
irregularly cup-formed. Under _B. concavus_ I have remarked how closely
some of its varieties approach to _B. amphitrite_, and it is to this
last variety that they approach; almost the only difference being that
the scuta in _B. concavus_ are longitudinally striated. Yet some of the
varieties of the two species are so distinct that it would be puerile
to class them together. I will only add, that after studying such
varying forms as _B. tintinnabulum_ and _amphitrite_ it is difficult to
avoid, in utter despair, doubting whether there be such a thing as a
distinct species, or at least more than half a dozen distinct species,
in the whole genus Balanus.
As with _B. tintinnabulum_, I will first give a full description of the
more common forms, alluding only to each less frequent variation, and
then separately describe briefly the more marked varieties.
_General Appearance._--Shape conical, either steep or considerably
depressed; sometimes tubular; orifice either nearly entire or deeply
toothed, not large, varying from rhomboidal to rounded-trigonal.
Surface of shell smooth, never ribbed, generally naked, but
occasionally the yellowish epidermis is persistent; in the same
individual, I have seen all the lower part of the shell thus covered
and the upper part naked, the line of separation being defined.
The _colour_ varies much, even sometimes considerably on the same
individual; generally white or pale gray, with dull violet-coloured,
longitudinal, moderately broad stripes; these stripes are sometimes
equidistant, but more usually they are arranged so as to leave
broad white spaces; the stripes fade away by endless variations,
the edges of the compartments and the carinal end of the shell
longest retaining any colour, until we have a uniformly white
shell, generally covered with a yellowish epidermis; or the white
is longitudinally marked with hyaline lines; this latter variety
has a very peculiar aspect, and I did not doubt it was specifically
distinct, until, in a number of specimens on a ship from the West
Indies, I got the most perfect series, and another scarcely less
perfect series from the Mediterranean, graduating into common
coloured varieties. Rarely the dull violet or purple stripes become
approximate and dark, so that the whole shell is tinted of a brownish
slate-colour, occasionally freckled with white. Again, we have
another set of very pretty varieties, with a white or very pale
pink ground, with either narrow bright pink or broad pinkish-purple
stripes. Again, I have seen numerous specimens of a variety, _var.
Stutsburi_, from the west coast of Africa, in which the upper part
of the shell is white, and the lower part shaded with pinkish or
dark purple approximate stripes, which often become confluent; in
one group, the whole shell being thus uniformly coloured, without
any vestige of stripes. I have seen another group from an unknown
locality, in which the lower part of the shell was uniformly
blueish-gray. A variety from Australia has narrow approximate dark
claret-coloured stripes, transversely freckled with white. Lastly, in
the variety _cirratus_, the whole shell is very pale purplish-brown,
with indistinct longitudinal brownish stripes, transversely freckled
with white lines. I considered this as a distinct species, until
quite lately finding forms which I could not possibly determine
whether to class as _B. cirratus_ or _amphitrite_.
The radii are generally snow-white, or freckled with a bright
mahogany tint, or rarely clouded with purple, or in the pink
varieties with pink. The scuta are dull purple or pink, generally
with a white band along their tergal margin; often, however, they
are white, with merely one or two purple fasciæ. The thickness or
strength of the shells varies much; some specimens attached to a
floating cane, from Natal and the Philippine Archipelago, were
extremely strong; others, from the Mediterranean and Australia,
and some tubular varieties from the West Indies, were very thin,
translucent, and fragile. _Size_: large specimens generally attain a
diameter of from half to three quarters of an inch in basal diameter;
and I have seen one or two specimens an inch in diameter.
_Scutum_; sometimes the surface is very smooth, but generally the
growth-ridges are moderately prominent; the latter are occasionally
very finely beaded, and this seems always the case with _var.
cirratus_. Internally, the articular ridge is prominent and reflexed:
the adductor ridge is sharp, very prominent, and straight; it runs
parallel to the occludent margin; close to its lower side there is
often a depression (Pl. 5, fig. 2 _i_), sometimes bounded by a slight
ridge, as if for the attachment of a muscle, but there certainly is
no muscle here: rarely the adductor ridge is only slightly prominent:
there is a small and shallow little pit of variable depth for the
lateral depressor muscle.
_Tergum_ (2 _k_-2 _o_); this valve is here far more variable than in
any other species: in the commonest purple-striped forms (2 _l_), the
valve is rather broad, the basal margin lies in nearly a straight
line on the opposite sides of the spur, which is placed at rather
less than its own width from the basi-scutal angle; the spur is
rather short, and in width about one fourth of the entire valve; its
lower end is either bluntly pointed or more commonly nearly square
(2 _k_) and parallel to the basal margin: in young specimens it is
generally sharper than in older ones. Externally, in the line of
the spur, there is either a slight longitudinal depression, or more
rarely a deep furrow. The carinal margin is more or less convex, and
is formed by upturned lines of growth: the scutal margin is broadly
inflected. Internally, the articular ridge in the upper part is very
prominent: the crests for the tergal depressor muscle are moderately
prominent, but very variable. Sometimes the carinal portion of the
basal margin is slightly hollowed out. In _var. Stutsburi_ (2 _m_, 2
_n_, 2 _o_), and in some white varieties, which differ most in the
shape of the tergum from the commoner varieties, the whole valve is
narrower, the spur is much sharper and narrower, the carinal half of
the basal margin is much hollowed out and slopes down towards the
spur, with the crests for the depressor muscles depending beneath the
basal margin, and with the carinal margin sometimes extremely convex
or protuberant. But the shape and position of the spur, and the
outline of the carinal half of the basal margin vary much in nearly
all the varieties.
_Compartments._--The upper parts of the parietal pores are either
filled up solidly with, generally coloured, shell, or they are
crossed by thin transverse calcerous septa: the longitudinal parietal
septa occasionally bifurcate at their bases close to the outer
lamina, making an irregular outer row of minute pores. The _Radii_
have their septa rather fine, and finely denticulated on both sides,
but sometimes only on the lower side; the thickness of the septa
varies a little; the interspaces are filled up solidly; the summits
of the radii are jagged and oblique, and usually form an angle of
about 45° with the basis, not being added to above the level of the
opercular membrane; but not rarely they reach up much higher, and
are very nearly parallel to the basis, extending from tip to tip of
the compartments. Again, in some ordinary varieties, and always in
_var. Stutsburi_, the summits of the radii are extremely oblique, the
radii themselves forming a mere border to the compartments to which
they belong. In no other species have I seen so great an amount of
variation in the form of the summits of the radii. The _alæ_, in like
manner, have their summits either very oblique, not being added to
above the opercular membrane, or they are only slightly oblique; it
often happens that in those specimens in which the summits of the
radii are nearly parallel to the basis, the alæ are very oblique, and
the converse: in other individuals, both radii and alæ have equally
oblique summits. The sutural edges of the alæ vary in thickness,
being either very thin and obscurely crenated, or moderately
thick and ribbed. The _basis_ is porose; but I have never seen an
underlying cancellated layer of shell, as is so common in several
species.
_Mouth_: labrum, with from four to eight, generally with six, little
teeth: mandibles with three teeth, and two minute lower teeth, or
mere knobs: maxillæ with the edge straight, or with the inferior
part forming a slightly step-formed projection. _Cirri_: the rami
of the first pair are unequal by three or four segments, but in
some specimens by five or six segments, with the front surfaces of
the segments in the shorter ramus extremely protuberant. The second
pair of cirri are short, with the front surfaces of the segments
moderately protuberant: the third pair have a tuft of bristles at
their bases on the thorax. The segments in the sixth pair have from
four to six pairs of spines on the segments; equal-sized specimens
seem to vary in this latter respect. There is a small sharp
projection on the dorsal base of the penis.
_Varieties._
With respect to _var._ 1, _communis_, I have nothing further to
remark, except that I have seen specimens identically similar from
the Mediterranean, Natal, the Philippine Archipelago, and Sydney;
at the latter place it is said to be rare, but in most places it is
the commonest variety, and is often attached to ships' bottoms. Of
_var._ 2, _venustus_, I have seen specimens from the west coast of
Africa, Natal, and Ceylon, in groups by themselves, and associated
with _var. communis_; it is much less common than _var._ 1. The third
variety, _pallidus_, is not uncommon; I have seen many specimens
from the bottoms of ships, from the West Indies, and the west coast
of Africa. Of the _var._ 4, _niveus_, I have seen the most perfect
graduated series passing into _var._ 1, both from the West Indies,
Florida, and the Mediterranean: I have seen other specimens from the
Red Sea and Madagascar. Of the _var._ 5, _modestus_, I have seen
only one group from an unknown locality; it is only remarkable from
its uniform colouring. The _var._ 6, _Stutsburi_, is more remarkable
than the foregoing; until quite lately I did not doubt that it was
specifically distinct; but as I have seen every character graduate
into other varieties, I am now convinced that it is not a true
species: all the specimens which I have seen have come on shells,
or on ships' bottoms, from West Africa. Of _var._ 7, _obscurus_, I
have seen three or four groups of specimens from unknown quarters,
both on pebbles, shells, and on cork (probably from the Atlantic
ocean); and likewise some specimens taken from the bottom of Her
Majesty's ship "Fly," on the east coast of Australia; these latter
are intermediate in character with the next _var. variegatus_;
from the Australian seas, which I at first ranked as an undoubted
species, but I have subsequently failed in discovering any sufficient
diagnostic character. Lastly, of _var. cirratus_, I have seen several
groups of specimens from India and the Philippine Archipelago, and
a group intermediate in character between this and the first and
third varieties, from Australia; I retained this variety owing to
its peculiar freckled, pale brown colouring and beaded scuta (of
which, however, I have seen decided traces in the common variety),
as a distinct species, after I had given up all the foregoing forms.
I entertain some doubts whether I have now acted right; but when I
found some specimens which, I found it impossible to decide, whether
to rank as _amphitrite_ or _cirratus_, I determined to take the more
prudent course, and sink the latter as a species. This variety, also,
seems to connect _B. amphitrite_ and _concavus_ very closely.
20. BALANUS PŒCILUS. Pl. 5, fig. 3 _a_, 3 _b_.
_Shell dull red, freckled with white. Scutum internally without an
adductor ridge; tergum with the spur, sharply truncated, almost one
third of width of valve._
_Hab._--West coast of South America, Mus. Cuming; attached to an
Avicula.
The appearance of the fragile shell, in the one group of specimens
which I have seen, leads me to suspect that they may have grown under
unfavorable circumstances. This species differs considerably in general
aspect, but not much in essential characters, from _B. amphitrite_; the
absence, however, of an adductor ridge to the scutum, and the sharply
truncated spur of the tergum, are sufficient to distinguish them. In
the opercular valves this species comes near to _B. vinaceus_, also
from the west coast of South America; but the striated scuta of that
species, the cancellated inner lamina of the parietes, the general
colouring, and square porose radii, are amply diagnostic characters.
_General Appearance._--Shell fragile, tubulo-conical, orifice large,
passing from diamond-shaped into oval. Colour fine dark rose,
freckled with transverse, sharply pointed, fine zig-zag white lines:
the pink is also so arranged as to obscurely give to the walls a
longitudinally striped appearance: radii generally rather whiter than
the walls, and similarly freckled: terga similarly freckled: scuta
dull red, with a white band along the scutal margin. Basal diameter
of largest specimen half an inch.
_Scutum_, externally smooth: internally, articular ridge moderately
developed, slightly reflexed: there is no adductor ridge: there is
a distinct pit for the lateral depressor muscle. _Tergum_, with the
scutal margin unusually prominent, toothed: longitudinal furrow
shallow, the edges apparently having no tendency to fold in: spur
short, barely one third of width of valve, with the lower end sharply
truncated, parallel to the basal margin: articular ridge and crests
for the depressores moderately prominent.
_Compartments._--Walls very fragile, with the outer lamina not
thicker than the inner lamina. _Radii_ fragile, broad, with their
summits moderately oblique; their sutural edges have the septa
plainly denticulated on both sides, with the interspaces filled up
solidly nearly to the tips of the septa. _Alæ_, with their summits
more oblique than those of the radii; their sutural edges smooth.
_Basis_ with an underlying cancellated layer. _Mouth_: labrum with
three unusually large teeth on each side of the notch: mandibles with
the fourth tooth tolerably well developed, the fifth being confluent
with the inferior angle. _Maxillæ_ simple. _Cirri_,--first pair with
one ramus longer by about four segments than the other ramus, which
has considerably protuberant segments: second pair with segments only
moderately protuberant: sixth pair with segments much elongated, but
bearing only four pairs of spines.
21. BALANUS EBURNEUS. Pl. 5, fig. 4 _a_-4 _d_.
BALANUS EBURNEUS. _Aug. Gould_ (!). Report on the Invertebrata of
Massachussetts, 1841, fig. 6.
_Shell yellowish white. Scutum striated longitudinally: tergum with the
spur truncated, the basi-carinal margin generally much hollowed out,
and the carinal margin protuberant in the upper part._
_Hab._--United States, from about lat. 42° to Charlestown; West
Indies; Honduras; Venezuela; attached to shells and floating wood.
Attached to ships' bottoms from Trinidad and Jamaica, associated with
_B. tintinnabulum_, _amphitrite_, and _improvisus_. Brackish water,
Salem, Massachussetts, according to Mr. Stimpson. Mus. Aug. Gould,
Agassiz, Stutchbury, Cuming, W. Dunker, &c.; very common.
_General Appearance._--Shell conical, or almost tubular; white, with
the surface very smooth, covered by thin yellowish epidermis, but
with the radii naked. Orifice large, passing from rhomboidal into
pentagonal, moderately toothed. Average full size, about one inch in
basal diameter; I have seen a specimen 1.3 in basal diameter, and the
same in height.
_Scutum_, plainly striated longitudinally: the teeth on the occludent
margin small. Internally, the upper surface is roughened: the
articular ridge is prominent, and either slightly or not at all
reflexed: the pit for the adductor muscle is distinct; the adductor
ridge is prominent in a variable degree, and is almost confluent
with the articular ridge. In one specimen from Beverly Bay, U. S.,
the scuta were extraordinarily disintegrated, and I could perceive
no trace of the external radiating striæ. _Tergum_, with the basal
margin on the carinal side of the spur sometimes deeply (Pl. 5, fig.
4 _b_), and sometimes only slightly (fig. 4 _d_), and rarely hardly
at all, hollowed out: when much hollowed out the valve may almost
be said to be two-pronged, with the carinal prong narrower than
the spur. There is no distinct longitudinal furrow, but the whole
scutal margin projects above the general surface of the valve. In the
carinal margin, in the upper part, there is a remarkable convexity
or protuberance in the same plane with the valve, from which it
is separated by a very slight and narrow ridge. The spur is about
one fourth of the width of the valve, with its lower end abruptly
truncated. Internally, the upper surface is much roughened with
finely crenated ridges: the distinct crests for the depressores cover
the whole of the so-called carinal prong.
_Compartments_; the radii and alæ have their summits oblique,
sometimes a little rounded, but not smooth. The septa on the sutural
edges of the radii are remarkably fine, and closely approximate; the
denticuli are excessively minute. The sutural edges of the alæ are
most delicately crenated; the alæ are largely added to during the
diametric growth of the shell, and above the level of the opercular
membrane. The parietal pores are square and rather large: they are
crossed by transverse septa almost close down to the basis: the
longitudinal septa have tolerably large denticuli at their bases.
The pores in the basis are crossed by numerous transverse septa.
When specimens grow in a group, the basis is sometimes irregularly
cup-formed.
_Mouth_: _labrum_ serrated with small teeth, decreasing in size
downwards, on each side of the central notch. _Mandibles_ with the
third tooth rather thick and blunt, and with the fourth and fifth
knob-like. _Maxillæ_, with the inferior part projecting much beyond
the rest of the edge, and bearing two long single spines: between
these two spines and the large upper pair, there are, in a full-sized
specimen, about seven pairs of moderately long spines, feathered on
their sides. _Outer maxillæ_ thickly clothed with very fine spines,
and remarkably prominent.
_Cirri_: first cirrus, with one ramus having twenty-six segments,
and longer by ten segments than the shorter ramus, which has sixteen
segments: the shorter ramus, and both ramii of the second pair, have
their segments remarkably protuberant in front; the protuberance,
in the upper segments, equalling in length the supporting part of
each segment: rami of the second cirrus unequal in length by five
segments. Third cirrus with the segments only slightly protuberant;
rami considerably longer than those of the second cirrus: at the
dorsal base of the pedicel of this third cirrus there is no tuft of
fine hairs, as is common in many other species. _Sixth pair_, with
the upper segments elongated, bearing from six to seven pairs of
spines; dorsal spines short, thin, and few.
_Affinities_: in external appearance of the shell, this species can
hardly be distinguished from some of the white varieties of _B.
amphitrite_; and there is a considerable resemblance, in some of the
varieties, in the opercular valves; but the longitudinally striated
scuta of _B. eburneus_ suffice to distinguish these certainly very
distinct species. Equally, or even more like _externally_, is this
species to the _B. Hameri_, so that I have received from an eminent
naturalist in the United States both species mingled in the same lot,
all bearing the same name of _B. eburneus_; but when the internal
structure of the shell is examined, the species are at once seen to
be far removed from each other. Still more close is the affinity
of this species to _B. improvisus_, both in internal and external
characters: it agrees with this species in the singular habit of
being able to live in brackish water: these two species are the only
ones which have the labrum serrated with teeth, graduated in size, on
each side of the central notch. In the case of _young_ specimens of
the _var. assimilis_ of _B. improvisus_, an inhabitant of the same
seas with _B. eburneus_, the diagnosis is most difficult without long
practice; for in the young of _eburneus_, the compartments are only
partially covered by yellow epidermis, and have a striped appearance,
the radii are sometimes very oblique, the scuta externally have not
acquired their longitudinal striæ, and internally the adductor ridge
lies not so close to the articular ridge as it does subsequently;
hence I for some time mistook the _var. assimilis_ of _B. improvisus_
for the young of _B. eburneus_. But I found in the latter, that the
rami of the first pair of cirri, are always, even in the earliest
youth, more unequal in length, and that each segment of the posterior
cirri bears a greater number of pairs of spines, there being, even in
very minute specimens, seven pairs. Moreover, after having examined
scores of specimens, I found I could almost always distinguish the
two species by the smoothness and curvature of the summits of the
radii of _B. improvisus_; I entertain no doubt whatever about the
distinctness of the two species; indeed, when both are mature,
besides the greater size, striated scuta, &c. of _B. eburneus_, their
general aspect is very different.
22. BALANUS IMPROVISUS. Pl. 6, fig. 1 _a_-1 _c_.
_Shell white: radii narrow, with their upper margins smooth, slightly
arched, very oblique. Tergum with a longitudinal furrow; spur with the
end rounded._
_Var._ assimilis, _with longitudinal white hyaline lines_.
_Hab._--England, Scotland, Belgium (?), Nova Scotia, United States,
West Indies, Rio Plata, Southern Patagonia, Guayaquil, West Colombia;
attached to wood, shells, rocks, ships' bottoms, from low tidal level
to twenty fathoms depth.
_General Appearance._--Shell conical, with a rather large
diamond-shaped orifice, moderately or but little toothed; very
smooth; walls never folded longitudinally; white, with an extremely
thin pale-yellow persistent epidermis. The radii are very narrow,
with their summits very oblique, rounded, and smooth; the epidermis
is generally more persistent on the radii than on other parts, and
this is exactly the reverse of what is common with _B. eburneus_. The
specimens from nearly fresh-water in the R. Plata (hereafter to be
mentioned), are brownish, and have undergone a remarkable degree of
corrosion, the outer lamina of the walls having been entirely removed
to near the base; hence the external aspect of these specimens is
wholly different from ordinary individuals. The _var. assimilis_
has also a very different appearance, owing to the dead white of
the walls being relieved by narrow approximate longitudinal hyaline
lines, corresponding with and caused by the longitudinal parietal
septa being externally visible through the outer lamina of the
parietes; the epidermis on the radii is also of a rather brighter
yellow. The largest specimens which I have seen are those from the
Plata, and those attached to a ship from the West Indies, and they
had a basal diameter of .6 of an inch: from .4 to .5 of an inch is
the more usual full average size.
_Scuta_, with the lines of growth but little prominent: articular
ridge prominent, but little reflexed: adductor ridge straight and
very prominent, varying a little in its distance from the articular
ridge; there is scarcely any depression for the lateral depressor
muscle; the upper internal surface of the valve is roughened with
ridges. _Terga_, with a moderately deep longitudinal furrow; spur
short, rather narrow, with the end rounded, placed at less than its
own width from the basi-scutal angle; in the Rio Plata specimens the
spur is close to this angle: the basal margin is generally straight
on opposite sides of the spur, but sometimes on the carinal side it
is a little hollowed out. The lines of growth are upturned along the
carinal margin, which consequently is a little protuberant, but to a
varying degree. The crests for the depressores are extremely distinct
and prominent. In the varieties having the basi-carinal margin
hollowed out, and the carinal margin protuberant, there is a marked
resemblance to the peculiar tergum of _B. eburneus_.
_Walls_: the parietal pores are tolerably large, and are crossed by
numerous transverse septa: the longitudinal septa are very finely
denticulated at their bases, but occasionally almost smooth. The
_radii_ are, as stated, extremely narrow, and very remarkable from
their smooth rounded edges; their septa are barely denticulated.
The _alæ_ are remarkably protuberant; they have their summits much
less oblique than those of the radii, and sometimes they are almost
parallel to the basis: their sutural edges are coarsely crenated.
_Basis_, flat, thin, permeated by pores, but the pores do not
generally run to the very centre; they are, as usual, crossed by
transverse septa.
_Mouth_: the _labrum_ is the most remarkable part; on each side of
the central notch there are generally two teeth; and on the two
sides of the notch itself nine or eleven smaller teeth, decreasing
regularly in size downwards till they become so minute as to be
hardly visible even under the compound microscope; thus, in the two
specimens closely examined, there were altogether twenty-two and
twenty-six teeth on the labrum. _Mandibles_ with the two inferior
teeth reduced to mere knobs: _maxillæ_ with the lower part of the
edge bearing two large spines, and generally, but not always,
forming a step-formed projection. _Cirri_: the ramii of the first
pair are but slightly unequal; in one specimen examined there were
fifteen segments in one ramus and twelve in the other: segments very
protuberant in front. Second cirrus with the segments only slightly
protuberant; segments thirteen. Third cirrus longer than the second
pair, with the rami rather unequal in length: there is a tuft of long
spines on the basal segment of the pedicel of this cirrus. Fourth
cirrus twenty-two segments. Sixth cirrus, in the same individual,
thirty-four segments: on each of these segments there are five or six
pairs of spines. I may specify that the longer ramus of the first
cirrus of a large Rio Plata specimen had twenty-four segments.
_Varieties, affinities._--When I first met with the _var. assimilis_,
misguided by its general aspect, I did not doubt that it was
specifically distinct; I was strengthened in this view by the
absolute identity of several hundred specimens attached to two
vessels from Jamaica and Trinidad, in the West Indies, with one
specimen from Charlestown, in the United States, sent me by Prof.
Agassiz, and with several in three lots from the western tropical
shores of South America: yet on close examination I can point out
no one distinguishing character, either in the shell or animal's
body, excepting the longitudinal hyaline lines on parietes, due to
the septa being externally visible. The presence of similar lines is
variable in white _vars._ of _B. amaryllis_ and _amphitrite_, and
they are seen in very young specimens of _B. eburneus_: hence it is
impossible to consider so trifling a character as specific; moreover,
lately I have seen a British specimen with hyaline lines, and some
few other specimens in an intermediate condition. Under the head
of _B. eburneus_, I have stated that although that species and _B.
improvisus_, which in the West Indies are associated together, are
most readily discriminated when old, yet when young, they so closely
resemble each other that the eye requires much practice to separate
them. On account of this species and _B. crenatus_ being sometimes
associated together on the shores of England, I have pointed out
under _B. crenatus_, the relative diagnostic characters of the two.
The chief affinity of _B. improvisus_ is certainly towards _B.
eburneus_; but in the narrow, oblique, rounded, and smooth-edged
radii, there is a relationship shown to the species in the last
section of the genus, such as _B. amaryllis_, and more especially to
the fossil _B. dolosus_: so close is the resemblance in the external
appearance of the shell, and in the structure of the opercular
valves, to the latter species, that I for some time did not discover
their distinctness. _Balanus improvisus_ has hitherto been overlooked
by naturalists, and has probably been confounded with _B. crenatus_
or _balanoides_.
_Range and habits._--This species, as far as my experience goes,
is commoner on the shores of Kent than on other parts of England:
the first specimens which I met with, I owed to the kindness of Mr.
Metcalf, they were attached to wooden stakes from Herne Bay, together
with a single specimen of _B. crenatus_: I have seen other specimens
from near Woolwich, from the Kentish oyster-beds, from Sandwich, and
from Ramsgate. The only other British specimens which I have seen
are from the River Itchen, in Hampshire, and from Loch Shieldaig,
in Ross-shire (Mus. Jeffreys), from a depth of twenty fathoms. This
species is often attached to wood. At Ramsgate, the specimens were
attached to a small coasting vessel, and they must have been immersed
five or six feet; they were associated with _B. crenatus_, and with
a few of _B. balanoides_. In the Brit. Mus. there are specimens
collected by Mr. Redman, from Nova Scotia, in North America. When
her Majesty's ship Beagle was beached at Santa Cruz, in Southern
Patagonia, numerous specimens were found adhering to her copper
bottom, some so small as to show that the species breeds in those
latitudes. Near Monte Video, in the estuary of La Plata, I found
many large, but much corroded specimens, adhering to some rocks in a
small _running_ stream of perfectly fresh water. The rise of the tide
is here small, but at high water the specimens apparently were for
a short time covered by the waters of the estuary, here itself only
brackish, and occasionally almost fresh. I took home some specimens,
and placing them in perfectly fresh water they continued for many
hours expanding and retracting their cirri with perfect regularity
and vigour. Here then we have a Balanus capable of living in fresh
water, and likewise in the saltest seas: even brackish water is a
deadly poison to several, probably to most, species of the genus; but
this, as we have seen, is not the case with the allied _B. eburneus_.
The water, I may add, at Woolwich, on the Thames, whence I have
received _B. improvisus_, must at times be very brackish. I have
already incidentally mentioned that the _var. assimilis_ was attached
in great numbers, associated with _B. eburneus_, _tintinnabulum_, and
_amphitrite_, on vessels from the West Indies: one specimen sent me
by Prof. Agassiz, from Charlestown, was attached to a specimen of _B.
eburneus_; and, lastly, I have seen three sets of the same identical
variety attached to shells from Guayaquil (in Mus. Brit. and Cuming),
and from West Colombia. Here, then, we have the same species with
an enormous range, from Nova Scotia and Great Britain to South
Patagonia; and, which is the case with scarcely a single mollusc, it
lives both on the eastern and western tropical shores of the South
American continent.
23. BALANUS NUBILUS. Pl. 6, fig. 2 _a_-2 _c_.
_Shell white, rugged: basis in parts imperfectly porose. Scutum with
the articular ridge minute; adductor ridge prominent, forming a deep
pit for the lateral depressor muscle: tergum with an internal patch of
purple; apex produced, purple._
_Hab._--California, Mus. Brit. and Aug. Gould; associated with _B.
glandula_, and attached to wood.
I have seen two specimens of this species, brought by Lady K. Douglas
from California; and two from Monterey, sent me by Dr. Aug. Gould. This
is a very distinct species, coming nearer to _B. porcatus_ than to any
other species: it is also allied to _B. cariosus_. In the basis being
in parts solid or not permeated by pores, it has claims to be placed in
the next section, in which I at one time included it.
_General Appearance._--Shell conical, rugged, sometimes furnished
with sharp longitudinal ribs; dirty white. Orifice not large, oval,
toothed. Radii rather narrow, with their summits oblique, much
jagged. Basal diameter of largest specimen 2.1; height only 1.3 of an
inch.
_Scuta_, broad, with the lines of growth prominent; internally,
articular ridge very little prominent, sometimes hardly developed,
but thick, ending downwards in a small free point. Adductor ridge
prominent, blunt, produced straight downwards, making a deep
longitudinal cavity for the lateral depressor muscle; in some
specimens this cavity is almost arched over, so as to tend to be
tubular, with a short ridge in the middle (Pl. 6, fig. 2 _a_): in
other specimens there is no trace of this tubular structure. _Terga_,
with the apex beaked, beak triangular, dull purple; the longitudinal
furrow is so shallow as hardly to exist. The basal margin slopes
down on both sides, with a nearly equable curvature towards the
spur; hence the spur is broad in its upper part, and narrow at its
obliquely truncated lower end. Internally, there is an elongated dark
purple patch: the shallow articular furrow is of quite remarkable
breadth; the articular ridge is medial, and the inflected scutal
margin is not wide. The internal surface of the spur is formed into
a ridge, which runs a little way up the valve, and is sometimes
partially separated from the spur itself (fig. 2 _c_), making the
basal extremity toothed or double. The crests for the depressores are
pretty well developed.
_Walls_, moderately strong: inner lamina slightly ribbed: the
denticuli on the bases of the parietal longitudinal septa are sharp:
I could not see any transverse septa in the parietal tubes. The
_radii_ are rather narrow; their summits are remarkably jagged and
very oblique; the septa are plainly denticulated on both sides, but
chiefly on the lower side; each septum itself, towards the inner
lamina of the radius, branches and divides: the interspaces are
filled up nearly solidly. The _alæ_ have apparently their summits
less oblique than those of the radii: their sutural edges are finely
crenated. The lower edge of the sheath is hollow underneath. The
_basis_ is flat; it is rather thin, and imperfectly porose; in parts
it is not at all porose, in others it is traversed only by very
minute pores: there is nevertheless, in some parts, even where the
upper layer is not porose, an underlying, cancellated layer.
_Animal's body_ unknown.
24. BALANUS CORRUGATUS. Pl. 6, fig. 3 _a_, 3 _b_.
_Shell white, longitudinally folded; radii narrow. Scutum internally
without an adductor ridge._
_Fossil_, Sub-Appennine formations; Colle in Tuscany; Mus. Greenough.
I have seen only two specimens of this species attached to rock,
collected by Mr. Greenough, at Colle, and kindly given by him to me.
The species comes near to the living _B. crenatus_, also found fossil
in deposits of this same age; it differs, however, distinctly from
that species, in having its basis permeated by pores, and, in a less
degree, in the sutural edges of the radii being more plainly crenated:
the opercular valves of the two species closely resemble each other.
This may be the _B. stellaris_ of Bronn, but it is futile attempting to
identify the species of this genus merely by external characters, even
when aided, as in this case, by an excellent drawing of the shell.
_General Appearance._--Shell conical, with broad rounded longitudinal
folds; orifice of moderate size, oval; radii narrow, with their upper
margins oblique; but the summits of both specimens had been much
broken. Colour, as it appears, originally white. Basal diameter of
largest specimen 3/4 of an inch.
_Scuta_, with the upper portion much reflexed; the articular
ridge is very prominent, and the articular furrow of great width;
when the valve is viewed from the outside the articular ridge is
very conspicuous: there is no adductor ridge. _Terga_, with the
longitudinal furrow very slight; the spur is from one third to
one fourth of the width of the valve, and its basal end is blunt
and almost truncated; it stands about half its own width from the
basi-scutal angle. Internally, the articular ridge is very prominent,
and the articular furrow narrow and deep, extending down the valve in
the line of the spur.
_Parietes_: the parietal tubes are remarkably large, and I think
this can hardly be an individual peculiarity: the tubes are crossed
by many transverse septa, close down to the basis. The _radii_ are
narrow, and have jagged, oblique summits: their sutural edges have
very distinct septa, barely denticulated, with the interspaces filled
up solidly. The _alæ_ have oblique summits; I was unable to make out
the structure of their sutural edges. The _Basis_ is very distinctly
permeated by pores, which are crossed by transverse septa.
The shell and opercular valves of _B. corrugatus_ so closely resemble
the same parts in _B. crenatus_, that I should not be much surprised at
seeing the two species graduating into each other, if a larger series
of specimens, from beds intermediate in age between the Sub-Appennine
formations and the present time, were obtained. If indeed the basis of
_B. crenatus_ were permeated by pores, the two species could hardly be
discriminated.
_Section_ D.
_Parietes permeated by pores. Basis and Radii not permeated by pores._
25. BALANUS PORCATUS. Pl. 6, fig. 4 _a_-4 _e_.
BALANUS PORCATUS. _Emanuel da Costa._ Hist. Nat. Test. Brit., p. 249
(1778).
LEPAS BALANUS. _Linn._ Syst. Naturæ (1767).
---- ---- _Born._ Testacea Mus. Cæs. Desc., Tab. 1, fig. 4, (1780).
---- ---- _Chemnitz._ Syst. Conch., 8 Band., Tab. 97, fig. 820,
(1785).
BALANUS ARCTICA PATELLIFORMIS. _Ellis._ Philosoph. Transact., vol.
50, Tab. 34, fig. 18 (1758).
---- SULCATUS. _Bruguière._ Encyclop. Method., Tab. 164, fig. 1
(1789).
LEPAS COSTATA and BALANUS. _Donovan._ British Shells, 1802-1804, Tab.
30, fig. 1, 2.
LEPAS SCOTICA. _W. Wood._ General Conchology, Pl. 6, fig. 3, sed non
_Lepas balanus_, Pl. 7, fig. 3, (1815).
BALANUS ANGULOSUS. _Lamarck_ (1818), in _Chenu_, Illust. Conch., Tab.
11, fig. 11.
---- TESSELATUS. _Sowerby_ (!). Mineral Conchology, Tab. 84 (1818).
---- SCOTICUS. _Brown._ Illust. Conch. Great Britain, Pl. 7, fig. 2,
sed non, Pl. 6, fig. 9 et 10 (1827); 2d edit., Pl. 53, fig.
1-3, 22, 23 et Pl. 54, fig. 1-3.
---- GENICULATUS. _Conrad._ Journal Acad. Philadelphia, vol. 6, part
2, p. 265 (1830), Tab. 11, fig. 16.
---- ---- _Aug. Gould_ (!). Report on the Invertebrata of
Massachussetts, fig. 9 (1841).
_Shell white, generally sharply ribbed longitudinally: radii with their
summits almost parallel to the basis. Scutum longitudinally striated:
tergum with the apex produced and purple._
_Var._ (_a_): _Walls without longitudinal ribs_. Mus. Brit., Cuming,
Stutchbury, Jeffreys.
_Hab._--South shores of England, Ireland, Scotland, Shetland Islands,
Iceland, Davis's Straits, 66° 30′ N.; Lancaster Sound, 74° 48′ N.
(Mr. Sutherland). Maine and Massachussetts, United States. China (?).
In deep water, common on shells, crustacea, and rocks, sometimes
imbedded in sponges.
_Fossil_ in the glacial deposits of Scotland, Uddevalla, and Canada;
in the mammaliferous and Red Crag of England; Mus. Lyell, Sowerby, S.
Wood, &c.
_General Appearance._--Shell conical, somewhat convex; white,
sometimes tinted yellowish, from the thin investing membrane;
the produced tips of the terga are purple: the parietes of each
compartment have from two to four strong, prominent, sharp, straight
longitudinal ribs; these are sometimes irregular, and rarely, as will
presently be described, they are absent. The radii are smooth and of
considerable breadth; their summits are nearly parallel to the basis
or only slightly oblique: hence the orifice is entire; it is rather
small and ovate, being broad at the rostral end, and very sharp and
narrow at the carinal end.
_Dimensions._--The largest specimens which I have seen from Great
Britain or Ireland, have been 1.3 of an inch in basal diameter:
in Mr. Cuming's collection, however, there was one much depressed
specimen from the Shetland Islands, 2.1 in basal diameter: a
regularly conical specimen from the coast of Massachussetts attained
a nearly equal diameter; out of the glacial deposits in the Isle of
Bute, Scotland, several specimens had this same diameter, namely, two
inches, and were even more steeply conical, being 1.85 in height;
some glacial specimens from Uddevalla and Canada, in Sir C. Lyell's
collection, were 1.7 in basal diameter. Hence, it appears, as we
shall presently see is likewise the case with _B. crenatus_ and
_Hameri_, that northern specimens, and those from the United States
and from the glacial deposits, often exceed in dimensions those from
Great Britain or Ireland.
_Scutum_: the lines or ridges of growth are broad and prominent;
they are divided into square beads by fine striæ, radiating from the
apex: and hence the valve is longitudinally striated. Internally,
the articular ridge is extremely little prominent; the adductor
ridge, or what must be called such, runs straight down under the
articular ridge, making a deep longitudinal pit for the lateral
depressor muscle. _Tergum_: the apex is a little produced, and
coloured purple, as well as the upper internal surface of the valve;
there is no longitudinal furrow, only a very slight depression: the
spur is placed close to the basi-scutal angle; it is rather long,
and measured across the upper part, is half as wide as the valve:
its lower end is truncated and rounded; the basal margin slopes
towards it. Internally, a very small portion of the scutal margin is
inflected: the articular furrow is shallow and broad: the crests for
the depressores are feeble. In young specimens the spur is bluntly
pointed.
The _Parietes_ (4 _e_) have large square parietal tubes: in the
upper part these are filled up solidly without transverse septa:
the longitudinal septa are finely denticulated at their bases, and
the denticuli extend unusually close to the outer lamina. In very
young specimens the inner lamina of the parietes is ribbed, in lines
corresponding with the longitudinal septa, as is the case with most
species of the genus; but in medium and large-sized specimens,
there are between the ribs, thus produced, from one to four smaller
ribs, which do not correspond with any longitudinal septa; they are
finely denticulated at their bases, and may be considered as the
representatives of longitudinal septa which have not been developed
and reached the outer lamina. I have seen no other instance of this
structure, namely, the presence of a greater number of ribs, on
the inner lamina of the walls, than there are longitudinal septa.
The _radii_ have their summits generally parallel to the surface
of attachment, as is usual in the first section of the genus, but
sometimes they are slightly oblique: the septa sometimes rudely
branch a little, but they exhibit scarcely a trace of denticuli: the
interspaces are filled up quite solidly. The _alæ_ have their summits
very oblique; their sutural edges are finely crenated.
_Basis_, rather thin, translucent, not permeated by pores; obscurely
furrowed in lines radiating from the centre: the circumference is
marked in a peculiar manner by the longitudinal septa, and by the
tips of those intermediate, denticulated ribs, which occur on the
inner lamina of the parietes.
_Mouth_: labrum with six teeth: mandibles with the fourth and fifth
teeth small and rudimentary: maxillæ, with a small notch under
the upper pair of spines; in the lower part there is a single
large spine. _Cirri_, dark brownish purple, making a singular
contrast with the white operculum and shell; first pair, with one
ramus, having twenty-six segments, and about twice as long as the
shorter ramus, having twelve or thirteen segments, with their front
surfaces protuberant. In the second pair the segments are but little
protuberant: third pair about one third longer than the second pair:
sixth pair, elongated, having in the same individual forty-six
segments; these segments have shield-shaped fronts, bearing five
pairs of spines, with some minute intermediate bristles. There is the
usual point at the dorsal base of the penis.
_Range: Geological History._--This species is common on the shores
of Scotland and Ireland; the most southern point of Europe whence I
have happened to see a specimen is Tenby, in South Wales: but I have
no doubt it is found further south; and Mr. Jeffreys, who knows this
species well, has found it common on the extreme southern shores of
England. In the United States, it is found on the shores of Maine and
Massachussetts: northward, I have seen specimens from Iceland, from
Davis's Straits, and from Lancaster Sound, in lat. 74° 48′ north;
these latter I owe to Sir J. Richardson. It is an inhabitant of deep
water; in Mr. Thompson's collection there are several specimens from
the Bay of Belfast, marked twenty-five fathoms, and one group said
to have come from "about fifty fathoms, on the coast of Antrim:" one
specimen from Cape St. Anne, Massachussetts, is marked as having come
from only five fathoms. This species is commonly associated, on both
sides of the Atlantic, with _B. crenatus_, and sometimes with _B.
Hameri_ and _Verruca Strömia_: mollusca, such as pectens, modioli,
and oysters, offer the most usual surfaces of attachment: I have,
however, seen many specimens on crustaceans, on rocks, and even on
the roots of the larger sea-weeds. This species is very common in the
glacial deposits of Uddevalla, of Skien in Norway, and of Canada, and
is associated with the same species as in the living state: I have
seen, also, specimens from the same formation in the Island of Bute,
Scotland. I have seen numerous specimens from the mammaliferous crag,
and a few from the Red Crag of England. I owe to the kindness of Mr.
J. de C. Sowerby an inspection of the original specimens of the _B.
tesselatus_ of the Mineral Conchology, which is certainly the present
species.
_Affinities._--This species is very distinct from every other; it
comes nearest, as shown in all the characters derived from its
opercular valves, to _B. nubilus_, and in this latter species we
have seen the basis plainly tending to lose its pores and thus
become solid. _B. porcatus_ is perhaps allied in some degree to _B.
trigonus_, and slightly to _B. crenatus_. The rather broad radii,
with their summits hardly oblique, give this species a very different
aspect from those species of the genus amongst which it must be
placed.
_Varieties._--A conical specimen, sent to me from the coast of
Massachussetts, is remarkable from the radii not having been at all
developed, being represented by mere fissures. I have seen a few
specimens of _var._ (_a_), (one collected by Sir E. Parry in the
arctic seas) which had a remarkably different aspect from the common
forms, but which, after a careful examination of the opercular valves
and of the animal's body, I feel convinced are not specifically
distinct: they are characterised by the walls being smooth and
absolutely destitute of the external longitudinal ribs; by the shell
being more cylindrical, with broader radii, and with the orifice
larger and more rhomboidal; the walls and radii are much thinner, and
the internal lamina is less plainly ribbed: the beak of the tergum is
not purple. As most of these specimens had grown in a group crowded
together, the difference of shape, and perhaps the thinness of the
walls, is thus explained. In a specimen from Davis's Straits, in
Mr. A. Hancock's collection, most of the above characters are in an
intermediate condition; there are only a few external longitudinal
ribs on the parietes; and the terga have not purple apices. In Mr.
Cuming's collection there are some fine, brilliantly white specimens
(_without opercula_) from the coast of China; these have thin walls
and radii, and the walls are not longitudinally ribbed, but they are
not smooth: the orifice is not large, nor the shape of the whole
shell cylindrical. It is just possible that these latter specimens
may be a distinct and representative species, but I do not think so.
26. BALANUS PATELLARIS. Pl. 6, fig. 5 _a_-5 _c_.
LEPAS PATELLARIS, (_Gmelin_). _Spengler._ Schriften der Berlin.
Gesellschaft, &c. b. i (1780), Tab. 5; Chemnitz, Neues Syst.
Couch., Tab. 98, fig. 839.
_Shell depressed; brown, generally with obscure longitudinal violet
stripes: radii (in full-grown specimens) with their summits rounded and
surfaces finely ribbed parallel to the basis: basis sometimes permeated
by imperfect pores. Scutum internally with an adductor ridge._
_Hab._--Bengal, on wood, Mus. Brit.; on a shell, Mus. Stutchbury;
Philippine Archipelago (young specimen), Mus. Cuming. According to
Spengler, on the Coromandel and Malabar coasts.
_General Appearance._--Shell depressed, sometimes much depressed:
orifice elongated, rhomboidal, but little toothed; surface smooth,
but in old specimens sometimes with the walls slightly folded
longitudinally. The radii are rather narrow, with their summits
oblique; in old specimens their summits are rounded, and their
whole surface finely ribbed parallel to the basis. Colour, in old
specimens dirty brown, tinged with violet, sometimes in longitudinal
bands, and with whiter irregular marks in the upper parts owing to
disintegration: in young specimens the walls are regularly banded
longitudinally, with violet-brown and dirty white; the radii being
generally of a paler dirty red or violet. Basal diameter of largest
specimen .9 of an inch.
_Scuta_, externally rather smooth; internally, articular ridge
prominent, reflexed, with the lower edge hollowed out so as to be
slightly hook-formed: adductor ridge small; there is a slight pit
for the lateral depressor. _Tergum_, with the spur bluntly pointed,
placed at about its own width from the basi-scutal angle; there is no
longitudinal furrow, only a slight depression; carinal margin arched
and protuberant: internally, articular ridge extremely prominent,
running down in the direction of the middle of the spur: crests for
the tergal depressores well developed.
_Parietes_, with the pores rather large; the internal lamina is very
strongly ribbed, the ribs being but slightly denticulated at their
bases: the parietal pores do not appear to be crossed by transverse
septa: sheath closely attached to the walls. The radii have jagged
oblique summits forming an angle of about 45° with the horizon; in
old specimens they become more oblique and narrow: and are then
very remarkable from their summits being arched and rounded, with
a crenated edge, and with their whole surface transversely ribbed
in horizontal lines; this is likewise the case with the recipient
furrow in the opposed compartments: in young specimens the radii are
externally quite smooth: the septa on the sutural edges are bluntly
denticulated; the interspaces being filled up solidly. The alæ have
their summits oblique, but much less oblique than the summits of the
radii; their sutural edges are very finely crenated.
_Basis_ thin, either quite solid, that is, not permeated by pores,
but only furrowed in lines radiating from the centre, or permeated
by pores towards the circumference, the pores being of very small
diameter;--so that we here have an important character variable
within the limits of the same species. Base flat, and this holds
good, as remarked by Spengler, even when the specimens are attached
to cylindrical pieces of wood.
_Animal's body_ unknown.
_Affinities._--In the basis being sometimes permeated towards the
circumference by pores, and by the colouring (the other species in
this and the next section being dirty white), _B. patellaris_ has
almost as strong a claim to be ranked in the last as in the present
section: in the rounded summits of the radii, and in the state of
the basis, it, perhaps, shows more affinity to _B. improvisus_ than
to any other species; it is, however, almost equally allied to _B.
glandula_.
27. BALANUS CRENATUS. Pl. 6, fig. 6 _a_-6 _g_.
B. CRENATUS. _Bruguière._ Encyclop. Method. (des Vers) 1789.
LEPAS FOLIACEA, _var. a_. _Spengler._ Skrifter af Naturhist.
Selskabet, b. i, 1790.
---- BOREALIS. _Donovan._ British Shells, Pl. 160 (1802-1804).
B. RUGOSUS. _Pulteney_ (?) Catalogue of Shells of Dorsetshire,
1799.
---- ---- _Montagu_ (?) Test. Brit. 1803.
---- ---- _Gould_ (!). Report on Invertebrata of Massachussetts
(1841), fig. 10.
B. GLACIALIS (?) _J. E. Gray._ Suppl. Parry's Voyage, 1819.
B. ELONGATUS (!), CLAVATUS (!), _Auctorum variorum_.
_Shell white: radii with their oblique summits rough and straight.
Scutum without an adductor ridge: tergum with the spur rounded._
_Hab._--Great Britain, Scandinavia, Arctic Regions as far as
Lancaster Sound, in 74° 48′ N. (Mr Sutherland); Behring's Straits
(Captain Kellett); United States; Mediterranean; West Indies, (Mus.
Brit.); Cape of Good Hope, (Mus. Krauss). Generally attached to
shells and crustacea in deep water; sometimes to ships' bottoms. Very
common.
_Fossil_ in glacial deposits of Scandinavia and Canada, Mus. Lyell;
in the mammaliferous, and Red, and Coralline Crags, Mus. S. Wood, J.
de C. Sowerby, Bowerbank; Miocene formation, Germany, Mus. Krantz.
I find, in most collections, this species confounded with _B.
balanoides_; I have even seen the two species, placed by Leach, on
the same tablet in the British Museum: _B. balanoides_ is, moreover,
generally confounded with _Chthamalus stellatus_; nor has any one
hitherto separated the present species from _B. improvisus_. On the
other hand, trifling varieties, both of _B. balanoides_ and _B.
crenatus_, have commonly been considered as specifically distinct. From
these facts it will be seen in what confusion our commonest British
species of Balanus have been left. After due deliberation, I have
little doubt that this is the _B. crenatus_ of Bruguière, and probably
the _B. rugosus_ of Montagu, but this latter author omits all reference
to the really important diagnostic characters between this species and
_B. balanoides_. The _B. crenatus_ is certainly the _B. rugosus_ of Dr.
Aug. Gould. In various collections, I find specimens of _B. crenatus_,
when coming from the arctic regions, called _B. glacialis_, _arcticus_,
and _borealis_; though I have not met with an authentic specimen of the
_B. glacialis_ of Gray ('Supp. Parry's Voyage,' 1819, p. ccxlvi), I
have little doubt that it would prove to be the present species.
_General Appearance._--White, usually of a dirty tint, from the
yellowish or brownish persistent epidermis: conical, generally
(fig. 6 _a_) with the parietes rugged and irregularly folded
longitudinally; but sometimes much depressed and extremely smooth (6
_b_); often cylindrical and very rugged; occasionally club-shaped (6
_c_), the upper part being much wider than the lower: specimens in
this latter condition sometimes have extremely narrow parietes, like
mere ribs, and wide radii. The orifice in the cylindrical varieties
is often most deeply toothed. The radii are generally narrow, and
have jagged oblique summits; but not infrequently they are so narrow
as to form mere linear borders to the compartments. The orifice is
rhomboidal, passing into oval, either very deeply or very slightly
toothed.
_Dimensions._--The largest British specimen which I have seen was
only .55 of an inch in basal diameter: specimens from Greenland
and the northern United States frequently attain a diameter of
three-quarters of an inch, and I have seen one single somewhat
distorted specimen actually 1.6 of an inch in basal diameter. The
specimens from the glacial deposits of Uddevalla and Canada appear,
on an average, to attain as large or larger dimensions than those
from the United States: on the other hand, the specimens from the
mammaliferous and Red Crag are smaller, the largest being only .35 in
basal diameter. When individuals have grown crowded together, their
length is often twice, and even occasionally thrice, as great as
their greatest diameter; thus I have seen a Greenland specimen 1.6 of
an inch in length, and only .75 in diameter. In the British Museum
there are some arctic specimens, one and a half inch in length, only
half an inch in diameter at the summit (fig. 6 _c_), thence tapering
downwards to a blunt point.
_Scuta_; the lines of growth are but little prominent: the surface
is generally covered by disintegrating membrane. The upper ends
are usually a little reflexed, so that the tips project freely as
small flattened points. Internally, the articular ridge is highly
prominent and somewhat reflexed: there is no adductor ridge, but a
very distinct impression for the adductor muscle: the depression for
the lateral depressor muscle is small, but variable. The _terga_
are rather small: the spur is short, and placed at rather less than
its own width from the basi-scutal angle; the basal margin slopes a
little towards the spur, of which the lower end is rounded or bluntly
pointed in a variable degree. There is no longitudinal furrow, hardly
even a depression. Internally, the articular ridge is very prominent
in the upper part; the crests for the tergal depressores are well
developed, but variable.
_Compartments._--The internal carinal margin of each compartment,
from the sheath to the basis, generally, but not invariably, projects
a little inwards beyond the general internal surface of the shell,
in a manner not common with the other species of the genus: the
basal edge of this projecting margin rests on the calcareous basis,
and is crenated like the basal edges of the longitudinal parietal
septa. The whole internal surface of the shell is ribbed, but the
ribs are not very prominent. The parietal tubes are large, and are
crossed in the upper part, and often low down, by transverse thin
septa: the longitudinal parietal septa are only slightly denticulated
at their bases; occasionally they divide at the basis close to the
outer lamina of the parietes, making some short outer subordinate
pores. In the circular furrow beneath the lower edge of the sheath,
there are sometimes little ridges, dividing it into small cells:
sometimes, however, this furrow is filled up by irregular knobs of
calcareous matter. The _radii_ are always rather narrow, and often
they form mere linear ribbons of nearly uniform width along the
edges of the compartments. Their summits or edges are always more or
less irregular and jagged: they form an angle with the horizon of
generally above 40°. Their septa are fine, and barely or not at all
denticulated. The alæ have oblique summits: their sutural edges are
rather thick and distinctly crenated. _Basis_ flat, calcareous, very
thin, with the surface slightly marked by radiating furrows, which
furrows answer to the radiating pores that occur in the bases of most
species. In a club-shaped arctic specimen, one inch and a half in
length, the summit being half an inch and the base only one fifth of
an inch in diameter, the basis was still calcareous, thick, and not
permeated by pores.
_Mouth_: labrum with six teeth: mandibles with the fourth tooth
minute or rudimentary, and the fifth generally confluent with the
inferior angle. Maxillæ with generally, but not invariably, a small
notch under the upper pair of great spines. _Cirri_, first pair with
the rami very unequal in length, one ramus being nearly twice the
length of the other; in a large specimen having a cylindrical shell
the proportional numbers of the segments in the two rami of the first
cirrus were ten to twenty-three; in a small conical specimen the
numbers were only eight to thirteen. The second cirrus has only two
or three more segments than the shorter ramus of the first pair: the
third cirrus has one or two more segments than the second; but it is
nevertheless decidedly longer than the second. On the dorsal surfaces
of both segments of the pedicel of the third cirrus, there is a
tuft of fine spines. The segments of these three pairs of cirri are
not much protuberant in front. The segments of the posterior cirri
have, each, four, or five, or six pairs of spines. _Penis_, with a
straight, sharp, short point on the dorsal basis.
_Range, habits, &c._--I have received specimens from all parts of the
coast of Great Britain and Ireland, generally attached to crustacea
and mollusca, and never hitherto from rocks uncovered by the tide.
This species is also attached to floating timber, sticks, fuci, and
occasionally to pebbles at the bottom of the sea. Mr. Thompson has
sent me specimens from twenty-five fathoms depth in Belfast Bay:
others on a Pinna from _about_ fifty fathoms on the coast of Antrim;
others from between three and six fathoms attached to Laminaria
digitata: there is a specimen in Mr. Jeffreys' collection marked
forty-five fathoms. It is often associated, both on the coasts of
America and Britain, with _B. porcatus_, and though these species are
so distinct, yet when both have their surfaces similarly affected by
being attached, as is often the case, to large Pectens, it is not at
first easy, by external characters, to distinguish them, except by
close inspection of the terga, which in _B. porcatus_ are beaked and
purple. The _B. crenatus_ is sometimes associated in deep water with
_B. Hameri_. At Ramsgate, in Kent, I saw a rudder of a ship, in which
the two or three upper feet were thickly coated with _B. balanoides_,
and the two or three lower feet with _B. crenatus_ and _improvisus_
mingled, together with a few of _B. balanoides_: occasionally
vessels are thickly encrusted with this species, but I have never
seen an instance of its concurrence with _B. tintinnabulum_ and
_amphitrite_--the commonest species on ships coming from the south.
I have seen specimens from Greenland, Baffin's Bay, the coast of
Labrador, and other specimens marked simply, "Arctic regions," and,
again, others from the shores of Maine and Massachussetts. The arctic
specimens, and those from the northern United States, are larger
than the British. I have seen one single minute specimen on a crab,
marked as having come from the Mediterranean. In the British Museum,
amongst some specimens of _B. eburneus_, ticketed as having been sent
from Jamaica, there was a small group of specimens, differing in no
one essential respect from the common varieties of _B. crenatus_:
at first I concluded that this was an erroneous habitat, and that
the specimens had really come from the United States, where _B.
eburneus_, is found as well as in the West Indies: for it appeared
to me exceedingly improbable that an animal which can exist in lat.
75° N. should inhabit the hot shores of Jamaica: but subsequently I
have received a specimen from Prof. Krauss, collected by himself in
Algoa Bay, which is perfectly characterised, and even has the little
cells in the furrow under the sheath: so that I am compelled to admit
this enormous range and capability of resisting the most extreme
climates. That this species should live in the tropical seas is the
more surprising, as the large size of the specimens in the northern
seas and in the glacial deposits, might fairly have been supposed
to have indicated special adaptation for a cold climate. The great
geographical range of this species accords with its range in time
from the present day to the Coralline Crag period.
The specimens from the glacial deposits which I have examined,
chiefly in Sir C. Lyell's collection, are very fine and large; they
are often associated, like the now living individuals, with _B.
porcatus_ and _Hameri_: they come from the well-known formation of
Uddevalla and from Canada. There are well-characterised specimens
in the mammaliferous Crag, at Bramerton and near Norwich, in Sir C.
Lyell's collection, and from Sutton and other places in the Red Crag
of the eastern shores of England: these specimens are decidedly not
only smaller than the glacial, but than the recent English specimens;
for the largest Crag specimens which I have seen had a basal diameter
of only .35 of an inch. The specimens which I have seen from the
Coralline Crag, and some others sent me by Krantz from the miocene
formation of Flonheim bei Abzei, in Germany, had not their opercular
valves, yet I cannot doubt, considering how few species there are in
the present section of the genus, that I have rightly identified them.
_Diagnosis._--Under the head of _B. balanoides_ I shall make a few
remarks on the diagnosis between that and the present species; as _B.
improvisus_ is found on the British shores, sometimes mingled with
_B. crenatus_, I may observe that, externally, the only difference
consists in the edges of the radii in _B. improvisus_ being much
smoother and rounded, and in the whole shell being less rugged.
Internally, in _B. improvisus_ the porose basis, the presence of an
adductor ridge on the under side of the scutum, the graduated teeth
on each side of the central notch in the labrum, and the little
inequality in length of the rami of the first pair of cirri, are
clearly and amply diagnostic.
28. BALANUS GLANDULA. Pl. 7, fig. 1 _a_, 1 _b_.
_Shell white; parietes with the internal lamina generally strongly
ribbed longitudinally, with the pores imperfect and small, sometimes in
part absent; radii narrow, with their summits rounded. Scutum with an
adductor ridge; tergum with the spur truncated and rounded._
_Habitat._--California, Mus. Cuming, Aug. Gould; attached to shells
and wood, together with _B. nubilus_. Southern Pacific ocean,
attached to _Pollicipes polymerus_; Mus. Brit.
_General Appearance._--Shell steeply conical, or cylindrical and
elongated; dirty white; walls rugged, longitudinally folded; radii
narrow, with their summits very oblique and rounded; orifice toothed.
Basal diameter of largest specimen half an inch.
_Scutum_, resembling externally that of _B. crenatus_; rather broad,
surface smooth; articular ridge very prominent, and articular furrow
very wide; hence, when the summits of the opercular valves are worn
down, the two scuta together form a square projection indenting
the two terga, as in _B. balanoides_. Internally, there is a small
adductor ridge, on the lower side of which there is a pit, as if
for a muscle. The depression for the lateral depressor muscle is
small, but variable. _Tergum_ without any longitudinal furrow, and
hardly a depression: spur broad, with its lower end truncated and
rounded; internally, articular ridge very prominent; crests for the
depressores well developed.
_Compartments_:--The internal surface of the parietes is smooth in
the upper part beneath the sheath, but generally very strongly ribbed
in the lower part, the ribs being plainly denticulated at their
bases; in other specimens, the ribs are very small, and even in parts
quite obsolete. The parietal pores are short and imperfect, sometimes
reduced to an extremely minute size, to be detected only when the
walls are broken across near the basal edge, and most carefully
examined; occasionally not even a trace of a pore exists. Hence in
this respect, this species offers a singular case of variation. The
_radii_ are narrow, and of nearly the same width from top to bottom;
their very oblique summits, when well preserved, are smooth and
rounded; their sutural edges are ribbed or crenated with extremely
fine, smooth septa; the recipient furrow is plainly marked by these
septa. The sutural edges of the alæ are crenated; their summits are
less oblique than those of the radii.
_Basis_, thin, finely furrowed in lines radiating from the centre;
margin sometimes deeply sinuous.
_Mouth_: labrum with the central notch rather widely open, with four
teeth on each side of it: palpi with very short spines along their
inner margins: mandibles with the fourth and fifth teeth forming
mere knobs: maxillæ small, with a mere trace of a notch under the
two great upper spines. _Cirri_; first pair with the rami unequal by
three or four segments, the longer ramus being only one quarter of
its own length longer than the other ramus. Second pair short, with
the segments (and those of the shorter ramus of first pair) somewhat
protuberant. Third pair with the rami one third longer than those of
the second pair. Sixth pair with the upper segments elongated, and
bearing six or seven pairs of spines.
_Affinities._--This species in general appearance closely approaches
_B. crenatus_ and _balanoides_, and it is related to them in many
essential parts, such as in the opercular valves. It agrees with _B.
balanoides_, and differs from _B. crenatus_, in the smallness and
imperfection of the parietal pores, and in the radii having rounded
summits; it agrees with _B. crenatus_ in the structure of its basis,
and in the prominent longitudinal ribs on the internal surface of the
parietes, and differs from that species in the spur of the tergum
being squarer, and in the scutum having an adductor ridge.
_Range._--From the appearance of the Californian specimens, I suspect
that they had adhered to tidal shells and to wood. The specimens in
the British Museum, adhering to _Pollicipes polymerus_, consist of
two lots, one of unknown origin, and the other certainly brought from
the southern half of the Pacific Ocean by Sir James Ross: it deserves
notice, that the _Pollicipes polymerus_, the supporting object,
ranges from California to the southern Pacific Ocean.
_Section_ E.
_Basis membranous._
29. BALANUS BALANOIDES. Pl. 7, fig. 2 _a_-2 _d_.
LEPAS BALANOIDES. _Linn._ Fauna Succica, 1746, et Syst. Naturæ, 1767.
---- ---- _O. Fabricius._ Fauna Groen., p. 424, 1780.
---- ---- ET CLIVATUS. _Montagu_ (!). Test. Brit., 1803.
BALANUS VULGARIS (?) _Da Costa._ Hist. Nat. Testacea, Pl. 17, fig. 7,
1778.
---- OVULARIS ET ELONGATUS. _Aug. Gould_ (!). Report, Invertebrata
of Massachussetts, figs. 7 and 8, (1841).
---- PUNCTATUS, CYLINDRICUS, ELONGATUS, FISTULOSUS CLAVATUS.
_Auctorum variorum_. Sed non B. punctatus, _Bruguière_,
Encyclop. Method., et non B. punctatus, _Montagu_, Test. Brit.
_Parietes either solid, or cancellated, or rarely formed by a single
row of pores. Tergum, with the spur bluntly or sharply pointed._
_Var._ (_a_) _with the parietes permeated by tubes; spur of tergum
sharply pointed; segments in the posterior pairs of cirri, bearing
from eight to ten pairs of spines_.
_Habitat._--Great Britain, France, Norway, Shetland Islands;
Greenland, according to O. Fabricius; North America, in lat. 66°
34′ N.; Labrador; Nova Scotia; Massachussetts, Delaware. Extremely
common, attached to rocks, shells, and wood, within the tidal limits.
I have no doubt that the present species is the _Lepas balanoides_
of Linnæus; though O. Fabricius is the only author who gives, in his
"Fauna Groenlandica," a sufficient description for the species to be
recognised with certainty. I believe this also is the _B. balanoides_
of Bruguière, though he is in error, as far as my experience goes,
in stating that the basis is ever calcareous. I have little doubt,
also, that this is the _B. vulgaris_ of Da Costa. The _B. balanoides_,
in its corroded and therefore punctured state, is certainly the _B.
punctatus_ of most British collections; but I do not believe it is
the _B. punctatus_ of Montagu, which I have scarcely any doubt is
the _Chthamalus stellatus_, so often found in the southern shores of
England, and even in some of the best arranged collections, mingled
with our present species.
_General Appearance._--The shell, in middle-sized and old specimens,
is almost invariably folded longitudinally and irregularly; it is
either dirty white or very often pale brown, and punctured from the
outer lamina having been corroded, to which action it is extremely
subject. In very young specimens, the surface is usually quite white
and smooth. The shell is sometimes much depressed; generally conical,
but when crowded together, cylindrical or club-shaped, one specimen
being even more than five-and-a-half times as long as wide. In Mr.
Jeffreys' collection there is a specimen 2.5 of an inch long, .45 in
diameter at the summit, only .2 in the middle, and rather more than
.2 near the base. Another specimen was 1.8 in length, its greatest
diameter being .35 of an inch at the summit. On the other hand, I
have seen a very depressed variety, with deeply folded walls, in
Mr. Thompson's collection from near Dublin, which was no less than
four times as wide as high; so that the difference in proportion of
height and greatest width, in the two extreme specimens, was nearly
as 10 to 1. Occasionally, from some unknown cause, isolated specimens
become cylindrical. The orifice of the shell, in the much elongated
specimens, is generally deeply toothed. The radii are always narrow,
sometimes extremely narrow, and have their summits smooth and rounded.
English specimens do not usually attain half an inch in basal
diameter; I have, however, seen one from near Yarmouth .9 of an inch
in diameter. Specimens from Massachussetts seem rather larger than
the average size of British specimens, many being .6 of an inch, and
one specimen a whole inch in basal diameter.
The _opercular_ valves so closely resemble those of _B. crenatus_,
that the description is necessarily comparative; in some cases they
could hardly be discriminated; generally, owing to the disintegration
to which this species is subject, the tips of the scuta are worn off,
and hence the articular ridges together form (Pl. 7, fig. 2 _a_) a
square projection, indenting the two terga; but I have examined young
specimens and others when not disintegrated, in which the opercular
valves, viewed externally, presented no difference whatever from
those of _B. crenatus_. The _scuta_, however, are, I think, generally
rather thicker, with the growth-ridges more prominent, and with
the tips certainly less reflexed than is usual with _B. crenatus_.
Internally, the articular ridge is rather less prominent: there is
no distinct adductor ridge. The _terga_ are often rather narrower
in proportion, and this especially holds good in the elongated
varieties; in these latter, there is occasionally a moderately deep
longitudinal furrow: the spur is often exactly the same shape as in
_B. crenatus_, but it is apt to be rather longer (Pl. 7, fig. 2 _c_)
and more pointed: in _var._ (_a_) it is pointed (fig. 2 _d_) in a
very remarkable manner. Internally, the articular ridge is decidedly
more prominent than in _B. crenatus_; the crests for the tergal
depressor muscles are either well developed or almost absent. From
this description it will be seen, how singularly the opercular valves
of the common varieties of these two species resemble each other. I
may mention that in some of the much elongated specimens, the muscles
going to the opercular valves partially lose their transverse striæ,
and become ligamentous.
The _Parietes_ are either quite solid, or more commonly are permeated
by minute pores, or by small irregular square tubes (Pl. 7, fig. 2
_b_), which only run up each successive zone of growth, for very
short distances, giving to the shell a cancellated structure, which
from corrosion is often externally visible. In the rather rare
variety (_a_) the parietes are permeated by regular tubes, extending
up to the apices of the compartments, but crossed by transverse
septa. The longitudinal septa, when such can be said to occur, in
no case are denticulated at their bases. The internal surface of
the parietes is either quite smooth or is traversed (Pl. 7, fig. 2
_b_) by very slight anastomosing ridges, but never, even in _var._
(_a_), by regular longitudinal ribs, as in most other species. The
carinal margin of each compartment, on the inside, projects, as in
_B. crenatus_, inwards, beyond the general surface of the shell, and
running down, rests on the basal membrane. The lower edge of the
sheath is rarely hollow beneath. The walls are lined by purplish,
or pale brown, or sometimes by almost black corium; numerous tubuli
penetrate the under sides of the walls and opercular valves; and
it is the intersection of these tubuli that gives the punctured
appearance to the often corroded surface of the shell. The _radii_
are narrow, generally very narrow; they have their upper and outer
margins, as seen externally, very oblique, rounded and (when well
preserved) smooth; their sutural edges are either quite smooth,
or sometimes just perceptibly pitted, like the basal margin of
the walls, or occasionally furnished with globular or arborescent
little ridges. The _alæ_ are also very oblique, but to a variable
degree, sometimes only slightly oblique: their sutural edges are
either smooth or obscurely crenated. _Basis_, membranous; in some
much elongated specimens, during continued growth, the basal edges
of the compartments approach each other so closely as almost to
touch, so that the whole shell becomes pointed at the bottom; but
on careful inspection I have never failed to find, even in the most
pointed specimens, a minute basal membrane; in other much elongated
varieties, in which the shell has apparently become too large for the
animal's body, the basal membrane, instead of being flat, becomes
drawn up deeply inwards, so as to touch the surface of attachment
only close round the basal edges of the shell.
_Mouth_: labrum with the teeth on each side of the central notch
unusually variable in number; I have seen specimens with only two
on each side, with four on each side, with five on one side and
four on the other, with five on one side and none on the other,
and with six on both sides; hence the total number ranges from four
to twelve. Mandibles, with the fourth and fifth teeth small, or
quite rudimentary. Maxillæ, with scarcely even a trace of a notch
under the upper pair of spines. _Cirri_; first pair, with one ramus
one third or one fourth longer than the other; in one specimen the
number of segments were nine and sixteen in the two rami: second
and third cirri short, very nearly equal in length, having in the
just-mentioned specimen respectively ten and eleven segments; the
sixth cirrus in this same specimen had twenty-five segments, each
segment being about as long as broad, and supporting six pairs of
spines. In the singular variety (_a_) the posterior cirri are more
elongated, and each segment supports seven or eight, and in one case
even ten pairs of spines! the third pair is also in this variety
proportionally rather longer. At the base of the third pair there
is a tuft of fine spines. The penis has not, as in _B. crenatus_, a
point at its dorsal basis. The branchiæ are very little plicated.
_Varieties._
Of the varieties having much elongated, club-shaped, hour-glass
shaped, and depressed shells, there is no necessity to say anything
in particular. With respect to the remarkable variety (_a_), I at
first named and described it as a distinct species: I have received
two lots, both from North America, one being sent me by Professor
Agassiz from Cape Cod. These agreed in having the parietes permeated
by regular tubes; in having the spur of the tergum most sharply
pointed; in the third pair of cirri being proportionally longer
compared with the second pair; in the sixth pair having more numerous
segments, namely, three times as many as in the third pair; in the
segments of the posterior cirri being more elongated, and especially
in the number of pairs of spines on each segment--amounting in one
case even to ten, a number unparalleled in other cirripedes. It may
naturally be asked why I have not retained so well marked a form as
a distinct species? In the first place, I found the most remarkable
character in _var._ (_a_), namely, the number of pairs of spines
on the posterior cirri variable, there being in one lot seven or
eight pairs, and in the other lot nine or ten pairs on each segment.
Secondly, all the characters by which this variety differs from the
common _B. balanoides_, are those which are variable in the latter;
this is especially the case with the structure of the parietes, and
in a lesser degree with the spur of the tergum. Thirdly, I found a
specimen in Mr. Cuming's collection, from Sweden (so that this _var._
(_a_) is not confined to North America), in which the cirri quite
resembled those of the American specimens, but the spur of the tergum
was in an intermediate condition as compared to that of ordinary
varieties; and the parietal tubes were of unequal sizes, and scarcely
more regular than sometimes in the true _B. balanoides_. And lastly,
I have seen specimens from Ayrshire, with the parietes permeated by
regular tubes, but with the tergum in an intermediate condition, and
with the segments of the posterior cirri not more numerous or more
elongated than in _B. balanoides_, supporting only six or seven pairs
of spines, that is only one more than is common with _B. balanoides_;
so that it was impossible to decide whether to rank the Ayrshire
specimen under _var._ (_a_) or under the common form, so that I was
compelled to give up _var._ (_a_) as a species.
_Monstrous individuals, with the male organs aborted:
Parasite._--Amongst some specimens, chiefly elongated ones, sent to
me from Tenby, in South Wales, I found no less than seven individuals
with some of the posterior cirri distorted, unequal on the opposite
sides, and in an almost rudimentary condition, and in each case
with the penis truncated, without any muscle entering the stump,
which was _absolutely imperforate_: the vesiculæ seminales were
much shrunk; in one case without any zoosperms; in another case
with headless zoosperms cohering in an unusual manner; hence it is
certain that these individuals were functionally only female, and
could not impregnate their own ova; yet in two instances the ova
had been impregnated, no doubt by neighbouring perfect individuals,
for they contained well-developed larvæ. Several of these monstrous
individuals were infested by one, two, or three curious crustaceans,
which have been described by Mr. Goodsir,[97] as the male of the
Balanus; but these supposed males are females, and were distended
with ova containing almost mature larvæ; I believe that they are the
females of the unnamed genus, belonging to the family of Ioniens,
described by Mr. Goodsir, which live parasitic within the sack (as I
likewise found) of the same individual Balani.
[97] 'Edinburgh New Philosophical Journal,' July, 1843.
_Diagnosis._--I have seen several specimens of this species and of
_B. crenatus_, absolutely undistinguishable in external appearance. I
may specify one of _B. balanoides_, imbedded in an alcyonidium, and
one of _B. crenatus_, imbedded in a sponge, and therefore neither at
all abraded. Generally, the tips of the scuta in _B. crenatus_ are a
little reflexed, whereas in _B. balanoides_, when the shell has been
at all disintegrated, the tips form a square projection locked into
the terga. _Bal. crenatus_ never assumes the punctured appearance so
common in _B. balanoides_. Very young specimens of the latter can be
distinguished by their dead white colour and smoothness. The edges
of the radii are almost always smoother than in _B. crenatus_, and
they are never so wide as is sometimes the case with _B. crenatus_.
When a specimen is disarticulated, our present species can at once
be distinguished from _B. crenatus_ (and from _B. improvisus_), by
its membranous basis, and by the solid or cancellated walls, which
are rarely permeated by regular tubes or pores; and the walls when
porose are not internally ribbed. I have already pointed out the
few very trifling points, in which the opercula of the two species
differ. The mouth and cirri offer likewise very few differences: in
_B. balanoides_ there are often more teeth on the labrum than in _B.
crenatus_; the rami of the first cirri are perhaps here rather less
unequal; the second and third pairs of cirri are certainly in most
cases more equal in length; and lastly, the segments of the sixth
cirri, even in the common varieties, bear, in equal-sized specimens,
more pairs of spines than in _B. crenatus_. We shall see that in
habits, with regard to depth, the two species differ, _B. balanoides_
inhabiting much shallower water than _B. crenatus_.
_Range, Habitats, &c._--This species is extraordinarily abundant
within the tidal limits round the shores of Great Britain, and
apparently of the northern United States. Besides numerous specimens
sent to me from very many English localities, the late Mr. W.
Thompson, of Belfast, kindly placed in my hands his very large
collection; from these materials it appears that _B. balanoides_
is the only tidal species in the northern parts of our island; but
in the south and south-west, it is associated with the _Chthamalus
stellatus_ and _Balanus perforatus_. I doubt whether this species
ever lives below the lowest tides; the case of a few specimens being
mingled with _B. improvisus_ and _crenatus_, (mentioned under the
latter species,) at the bottom of a rudder of a small vessel, about
six feet deep, is hardly an exception, for the water would there
be troubled and aërated almost as in a breaker; and on this very
rudder the upper two or three feet were coated exclusively by the
_B. balanoides_. This species lives on rocks at both the uppermost
and lowest limit of the tides; I am informed by Mr. Thompson, that
he has seen specimens attached to a spot not covered by water
during neap-tides. As a proof of its tenacity of life, Mr. Thompson
informs me that he accidentally kept some specimens in a box, in a
warm sitting-room, and found them alive seven days afterwards. This
same most accurate observer finds, however, that _B. balanoides_
is very susceptible to brackish water; he says, "that having kept
some specimens alive for a week in excellent health, the water being
changed once in thirty-six hours, they were one day killed instantly
by some water, though brought from the same part of the estuary as
usual, having been rendered brackish by much rain having lately
fallen." I may recall the fact, that _B. improvisus_ lives daily for
hours in absolutely fresh running water.
The _B. balanoides_ lives attached, often continuously coating
many square feet of the surface, to rocks, pebbles, wooden-piers,
littoral shells and ulvæ. The most northern point whence I have
received specimens, is in lat. 66° 34′ in North America, collected
by Mr. Sutherland; and the most southern point is Delaware Bay, in
the United States, in lat. 39°; I do not believe that this species
extends into the Mediterranean, for Ranzani (Mem. di Storia Nat.),
who particularly attended to the nature of the basis, was not
acquainted with any _Balanus_ having a membranous basis; and Poli
(Test. Ut. Siciliæ,) describes only two species thus characterised,
and these are manifestly _Chthamali_.
With respect to the rate of growth of this species, I am indebted to
Mr. W. Thompson for the following note:--
"Sept. 29, 1848.--I examined a great number of Balani, in reference
to the growth made by them during the present season, and found it
to average three lines in diameter, and at most four lines. I saw a
few minute specimens, only one line in diameter, showing that the
species continued to breed until lately: these latter were probably
not more than four weeks old. The young of the present year are
plainly distinguished from the older ones, by their pure white colour
and fresh appearance. Judging from the size of this year's specimens,
and of the older ones on the same stones, I am of opinion that the
term of life of the species is two years. Of the older shells, which
I examined and found living in the spring, nine tenths are now dead,
the walls only remaining, the opercular valves having been washed
away."
Mr. Thompson goes on to say, that the individuals which had, on July 3,
a basal diameter of from two and a half to three lines, had attained,
by the 30th of September, a diameter of four and a half lines, this
being here the maximum size of the species.
30. BALANUS CARIOSUS. Pl. 7, fig. 3 _a_-3 _e_.
LEPAS CARIOSA. _Pallas._ Nova Acta Acad. Scient. Petrop. tom. ii
(1788), p. 240, Tab. 6, fig. 24.
_Parietes thick, formed by several rows of unequal-sized pores. Tergum
narrow, with the apex beaked, and spur sharply pointed._
_Hab._--Columbia River, west coast of North America, Mus. Brit.
and Cuming; Behring Straits (Capt. Kellett); the Kurile Islands,
according to Pallas. Attached to shells, and to each other in groups.
_General Appearance._--Shell steeply conical, with a rather small
oval orifice; or cylindrical, with a large rhomboidal and little
toothed orifice. Colour dirty white. Surface either simply rugged,
or more commonly covered by numerous, narrow, extremely prominent,
longitudinal plaits; from the manner in which these overlap each
other, the shell almost appears as if thatched with straw. The
upper corroded part of the shell usually exhibits a cancellated
and finely punctured surface. The radii are generally very narrow,
forming towards the base of the shell a mere narrow ribbon to each
compartment, and often hardly distinguishable; but in one specimen
they were of considerable width: in the former case, the alæ are
often widely exposed. The largest specimen which I have seen was 1.5
of an inch in basal diameter, but Pallas gives 2.2 as the measurement
of a specimen from the Kurile Islands.
The opercular valves are united to each other and to the shell by
unusually strong membrane; and the upper parts of both valves, in all
the specimens seen by me, have been much disintegrated. The _Scutum_,
in old specimens, is faintly striated longitudinally, but in some
there is hardly a trace of this: the occludent margin is furnished
with a few large knobs, not corresponding with every alternate line
of growth (as is usual with other species), but with every fourth or
fifth line. Internally, the articular ridge is moderately prominent
(in young specimens more prominent) and reflexed. The adductor
ridge is sharp and prominent; in the upper part it is confluent
with the articular ridge, but in young specimens can be seen to be
distinct; in the lower part it borders a large deep cavity for the
lateral depressor muscle, in the middle of which there is a very
slight longitudinal ridge; this cavity sometimes is almost closed or
arched over in its upper part. In one specimen, the basal margin of
the scutum was deeply hollowed out in the middle. The _Tergum_ is
remarkably narrow, with its apex produced into a triangular beak,
hollow within, and sometimes faintly tinged purple. A deep, closed,
longitudinal furrow runs down the valve. The spur is long, remarkably
narrow, and pointed. Internally, the spur is produced up the valve
as a ridge: the inflected scutal margin, and the prominent articular
ridge, are both nearly straight, and parallel to the spur. The crests
for the depressores are sharp and very prominent.
The _Parietes_ are very thick and strong: unlike every other
species of the genus, they consist of several very irregular rows,
of unequally sized, round or angular tubes (3 _b_). These tubes or
pores are generally short, and are at frequent intervals crossed by
transverse septa; they often rather deserve to be called cells than
tubes. New tubes are formed along the inner as well as along the
outer lamina. They are lined by dusky purple corium. The internal
surface of the parietes is smooth in the upper part, and in the
lower, it is reticulated by slight, irregularly branching ridges.
The carinal internal margin of each compartment projects a little,
as in the case of _B. crenatus_ and _balanoides_. The lower edge of
the sheath is either hollow beneath, or is united to the walls. The
_radii_ in one specimen were broad, with slightly oblique, jagged
summits; generally they are extremely narrow, forming more ribbons
along the lower edges of the compartments, barely extending up as
high as the sheath. They can sometimes hardly, or not at all, be
seen, until the shell is disarticulated: in rather young specimens
the sutural edge is sometimes quite smooth; in old specimens the
lower part of the edge has coarse arborescent septa, with the
interspaces filled up solidly, whilst the upper part is smooth.
The _alæ_ are conspicuous from the outside, owing to the little
development of the radii; but owing to the diametric growth not
having been great, the part added during such growth is narrow;
the summits of the alæ are only slightly oblique: the sutural edge
is coarsely crenated, with the teeth denticulated or slightly
arborescent.
_Basis_ membranous.
_Mouth_: labrum with only four very minute teeth: mandibles with four
teeth; the third tooth broader than the first; the fourth small.
Maxillæ, with the two upper spines placed on a slight prominence,
beneath which there is a small notch. _Cirri_ of a very dark colour
(much injured): the segments of the first, second, and third pairs
very broad and short, protuberant in front, and most thickly clothed
with spines; the third pair is very little longer than the second
pair: the sixth pair (in a large specimen above one inch in basal
diameter) had the segments broader than long, each furnished with
seven pairs of spines.
_Affinities._--This species, though very distinct, evidently comes
near to _B. balanoides_, especially to _var._ (_a_). By merely
doubling or trebling the irregular rows of short tubes in the
walls of _B. balanoides_, with their reticulated inner lamina and
longitudinally folded outer lamina, we should have the structure
exhibited in _B. cariosus_. We have seen, also, that in _var._
(_a_) of _B. balanoides_, the spur of the tergum is remarkably
sharp, as in _B. cariosus_. This species, also, in a very marked
manner approaches in many characters, especially in the opercular
valves, in the cirri, and to a certain extent in the shell and
basis, to _B. flosculus_, and even in external appearance to _var.
sordidus_ of the latter--an inhabitant of the opposite extremity
of the continent, namely, of Tierra del Fuego. Again, the tergum
to a certain extent, and the scutum in a singular manner, resemble
these valves in _B. nubilus_, showing an unequivocal affinity to
that species. With respect to the most remarkable character of the
species, namely, the several irregular rows of tubes or pores in the
walls, it deserves notice that in _B. crenatus_, which is certainly
closely allied to _B. balanoides_, the longitudinal septa sometimes
divide near the outer lamina, thus giving rise to a few additional
tubes. Of the above several species, to which our present species is
allied, _B. flosculus_ stands in the next section, and _B. nubilus_
and _crenatus_ in the last: hence we see that _B. cariosus_ has
singularly divergent affinities. The peculiar structure of the
parietes, together with the general appearance of the shell, made me
at the first moment suppose I was examining a Tetraclita (or Conia of
Leach); hence, also, it has arisen, that _Lepas cariosa_ of Pallas
has often been quite erroneously given as a synonym of _Tetraclita
porosa_.
31. BALANUS DECLIVIS. Pl. 7, fig. 4 _a_-4 _d_.
_Parietes solid; rostrum nearly twice as long as the carina or
carino-lateral compartments, hence the basis is oblique. Tergum with
the spur truncated, half as wide as the valve._
_Hab._--West Indies; Mus. Brit.--Jamaica, imbedded in a sponge; Mus.
Cuming.
This is a remarkable species; when first seeing it imbedded in numbers
in a sponge, I did not in the least doubt but that it was an Acasta:
on examination, however, it is found to have a membranous basis, and
therefore cannot by the definition enter into that sub-genus, to which,
however, it is very closely allied. It differs from other sessile
cirripedes very remarkably in the rostrum being nearly twice as long as
the carinal compartments, so that the basis is always very oblique, or
placed almost on one side; in this elongation of the rostrum, although
in a different direction, we are reminded of _B. calceolus_ and its
allies; and these latter we know can hardly be separated from certain
species of Acasta. Hence the position of our present species in this
section, is not natural; but I am unable to place it elsewhere, without
breaking down every definition: it should stand somewhat isolated,
on one side of a line of affinity connecting _Balanus calceolus_ and
_navicula_ with _Acasta purpurata_.
_General Appearance._--The shell is thin, fragile, smooth, and
white, but covered to a considerable extent by a brown membrane,
which on the sheath and opercular valve is of a bright tint, and
clothed with bristles. Viewed laterally, the rostrum is seen to
be considerably bowed, and from its being nearly twice as long as
the other compartments, with its lower end bluntly pointed, the
basal margin of the whole shell is rendered very oblique, forming a
slightly concave line. The lateral compartments are rather longer,
and about one third broader than the carino-lateral compartments.
The rostrum, from terminating downwards in a blunt point, instead
of being square or truncated, as in all other Cirripedes, and from
the upper end being, as is usual, pointed, has, when disarticulated
from the other compartments, the shape of a boat. The parietes are
not at all porose: their internal surface sometimes shows traces
of longitudinal ribs, but sometimes there are none. The radii are
narrow, with their summits very oblique, and their sutural edges
smooth. The sutural edges of the alæ are likewise smooth. The largest
of Mr. Cuming's specimens was .2 of an inch in diameter; but a
disarticulated specimen in the British Museum must have been larger,
having a rostrum .3 in length. The _Basis_ is membranous.
The _Scuta_ are rather convex; they have their lines of growth
approximate, most finely crenated, so as to be very feebly striated
longitudinally. Internally, the articular ridge is pretty well
developed, its lower edge being very oblique; there is barely an
adductor ridge: the pit for the lateral depressor muscle is deep.
The spur of the _Tergum_ is placed close to the basi-scutal angle
of the valve; it is about half as wide as the valve, with the
lower end truncated: sometimes it may be rather said to be bluntly
pointed, owing to its carinal side sloping up to the basal margin.
The articular ridge is pretty well developed. The crests for the
depressor muscles are barely discernible.
_Animal's body_ unknown.
_Section_ F.
_Parietes and radii not permeated by pores; basis sometimes permeated
by pores, sometimes not permeated, sometimes excessively thin and
hardly distinguishable._
32. BALANUS HAMERI. Pl. 7, fig. 5 _a_-5 _c_.
LEPAS HAMERI. _Ascanius._ Icones rerum naturalium, Tab. 10, 1767.
---- TULIPA. _O. F. Müller._ Prodromus. Zoolog. Dan. 1776; sed non
_L. tulipa_, in Poli, Test. ut. Siciliæ; neenon _B. tulipa_, in
Bruguière, Encyclop. Method; neenon _B. tulipa_, in Sowerby,
Genera of Shells.
---- TULIPA ALBA. _Chemnitz._ Syst. Conch. Tab. 93, fig. 832.
---- FOLIACEA. _Spengler._ Skrivter of Naturhist. Selskabet, 1 B.
1790.
BALANUS CANDIDUS. (Tab. emendata) _Brown._ Conch. Great Britain
(1827), Tab. 6, figs. 9 and 10, et 2d edit. Tab. 54, figs. 9-12.
---- TULIPA. _Lyell._[98] In Phil. Transact., 1835, p. 37, Tab. 2,
figs. 34-39.
[98] Sir C. Lyell remarks that this is apparently the _B.
Uddevallensis_ (Linn.) of Swedish lists of fossils. Prof. E. Forbes
has shown (Mem. Geolog. Survey of England, vol. 1, p. 364) how this
name arose, from a short description, prior to the introduction of
the binomial system, "Lepas quæ Balanus Uddevallensis," given by
Linnæus in his Wast-Gotha Resa, in 1747.
For the reference to Ascanius' work, which is on the binomial
system, and subsequent to the 10th edit. of Linnæus, in 1758, I am
greatly indebted to Mr. Sylvanus Hanley. Had it not been for this
gentleman, I should have used Müller's name of _B. tulipa_ as the
first name.
_Shell white: radii with their oblique summits smooth and arched;
sutural edges smooth: basis solid. Scutum feebly striated
longitudinally: tergum with the spur narrow._
_Hab._--Coast of Yorkshire; Scotland; Galway, Ireland; Isle of Man,
and Anglesey, twelve fathoms. Generally in deep water; not very
common. George's Bank, Massachussetts, United States; Mus. Aug.
Gould. Iceland, Finmark, and the Faroe Island, according to Spengler.
Attached to crustacea, mollusca, stems of fuci, and stones; often
associated with _B. porcatus_ and _crenatus_.
_Fossil._--In glacial deposits at Uddevalla in Sweden, and Beaufort
in Canada; Mus. Lyell. Banks of the Dwina, Russia; Mus. Murchison.
Greenland, "in blue clay," according to Spengler. Red Crag (Sutton)
Mus. S. Wood.
_General appearance._--Shell tubulo-conical, very smooth, white,
generally more or less covered by yellow thin membrane: orifice
large, sub-triangular: radii moderately broad, with their more
or less oblique summits slightly rounded and smooth; from this
circumstance the shell has been justly compared to the half-opened
flower of a white tulip. Specimens often exceed an inch in basal
diameter; I have seen one from Scarborough two inches in diameter
and one and three quarters in height: another specimen was 1.6 in
diameter and 3 in height. The specimens in the glacial deposits seem
even to have acquired larger dimensions, one from Uddevalla being
nearly four inches in height.
_Scuta_, elongated, flat, feebly striated longitudinally:
internally, articular ridge short, moderately prominent: adductor
ridge, confluent in the upper part with the articular ridge,
running straight down and forming a rather large cavity for the
lateral depressor. _Terga_ feebly striated longitudinally, with a
longitudinal furrow, having the sides, in old specimens, partly
closed in: the basal margin slopes much towards the spur, which is
rather long and narrow, with its end rounded: it is placed at about
its own width from the basi-scutal angle. Internally, articular
furrow narrow; crests for the depressores moderately prominent, but
in a variable degree.
_Compartments_: these are unusually thin, and separate easily: the
parietes are finely ribbed longitudinally on their insides; the
bases of these ribs being just perceptibly denticulated. _Radii_,
with their summits oblique (usually at about an angle of 45°),
slightly arched and quite smooth: the smoothness is produced by the
edge being a little inflected: sutural edge quite smooth, without
even a trace of septa or denticuli. _Alæ_ oblique, generally rather
less oblique than the summits of the radii: sutural edges smooth,
with an excessively fine linear furrow running along the edge, a
little towards the inner side, and filled with a yellow ligamentous
substance: a furrow of this kind I have seen in no other species.
_Basis_, solid, not permeated by pores; either smooth, or slightly
furrowed in lines radiating from the centre.
_Mouth_: labrum with scarcely perceptible minute bead-like teeth
thinly scattered along the edge. _Palpi_ and _outer maxillæ_ rather
sparingly clothed with hairs. _Mandibles_ with teeth rather sharp;
the fourth and fifth teeth small, but well developed; inferior angle
pointed with fine spines. _Maxillæ_ with a deep notch under the two
upper great spines. _Cirri_, the first pair is short, with rami of
nearly equal length: the segments are not protuberant in front either
in the first or second pairs. In the posterior cirri, the segments
bear four pairs of spines, with a tuft of rather long intermediate
spines: in young specimens there are only three pairs: the spines in
the dorsal tufts are short and thin.
When the shell is disarticulated, this species cannot be confounded
with any other; but judging by external characters alone, it may
sometimes be very easily confounded with _B. eburneus_, and I have
received the two species under this one name from Massachussetts:
generally _B. Hameri_ may be distinguished from _B. eburneus_ by
the smoothness of the summits of its radii, and by the so-called
epidermis being of a darker yellow.
With respect to the fossil specimens from the glacial deposits,
I have little to add; I have seen one from Uddevalla, as already
remarked, four inches in height, and a lateral compartment broader by
one fourth than the same compartment in any recent specimen. As Sir
C. Lyell remarks (Phil. Transact.), the compartments are always found
separated, which is accounted for by their weak union in a recent
state. This species, when fossil, is usually associated with its
deep-water congeners _B. porcatus_ and _crenatus_, as at the present
day.
I must here mention that I have examined a considerable number of
_separated_ compartments, without opercular valves, brought from
Barbados, in the West Indies, showing the existence there of a
closely allied or possibly identical species. The only difference
which I can point out in these compartments is, that the parietes
are rather thicker, and the radii rather narrower, with more oblique
summits: some of the compartments are two inches in length. It seems
very improbable that the true _B. Hameri_ should extend to the West
Indies, but after what has been seen in the case of _B. crenatus_,
this is possible.
33. BALANUS AMARYLLIS. Pl. 7, fig. 6 _a_-6 _c_.
_Shell striped or clouded with pinkish-purple, or quite white; radii
narrow, with their oblique summits smooth or arched: basis porose.
Scutum plainly striated longitudinally: tergum with the spur narrow._
_Var._ (_a_):[99] _bright rosy pink, not distinctly banded
longitudinally_. Hab. North-east coast of Australia.
_Var._ (_b_): _snow white, glossy; orifice deeply toothed_.
[99] This variety perhaps is the _B. roseus_ of Lamarck, as
figured in Chenu, 'Illust. Conch.' Tab. 2, fig. 9; but as Lamarck
does not even notice such conspicuous external characters as the
longitudinal striæ on the scuta, and the smooth rounded edges of
the radii, it is impossible to identify his species.
_Hab._--Mouth of the Indus; East Indian Archipelago; Philippine
Archipelago; Moreton Bay, and the north-east coast of Australia.
Attached frequently on ships' bottoms, associated with _B.
tintinnabulum_ and _amphitrite_. Sometimes attached to _Gorgoniæ_
with _B. calceolus_.
_General Appearance._--Shell steeply conical, with the orifice
sub-rhomboidal, moderately large, very slightly, or deeply notched:
surface very smooth: white, longitudinally banded with pinkish or
leaden purple, with sometimes a purplish, sometimes a yellowish tint,
the latter owing to the persistent epidermis; the bands are pale,
and often fade away in the lower, and sometimes in other parts of
the shell; the epidermis is generally more persistent on the narrow
rounded radii than on the parietes, and hence the radii are generally
yellowish. The opercular valves are pale dull purple: the sheath is
darker purple, with the exception of the portions of the alæ added
during the diametric growth, which are of a dead white, and are
externally conspicuous. The scuta are striated longitudinally. I
may remark, that, excepting the narrowness of the radii, with their
quite smooth, rounded and very oblique summits, some specimens are
hardly distinguishable, in external aspect, from varieties of _B.
amphitrite_. If the specimens from the north-east coast of Australia,
of which I have seen many (but unfortunately only one small one had
its opercular valves), form, as I fully believe, merely a variety; it
is characterised by its nearly uniform beautiful rosy pink, without
any _distinct_ longitudinal bands: of these specimens I have seen
one two inches in basal diameter, and three in height: of ordinary
duller-coloured striped specimens, the largest was 1.7 in basal
diameter. Of the perfectly white _var._ (_b_), I have seen several
specimens, the largest being .6 of an inch in diameter: these have
a somewhat peculiar aspect, but I have met with only one specimen
with opercular valves, and that was extremely young: I at first
considered this form as specifically distinct; but I can point out,
after careful examination of the whole shell, operculum, and internal
animal of the young specimen, no sufficient diagnostic characters.
_Scutum_, plainly striated longitudinally, with the striæ dividing
the prominent lines of growth into squarish beads: internally, the
upper part of the valve is roughened: the articular ridge is short,
remarkably little prominent, and not reflexed; the adductor ridge
is blunt and little prominent; sometimes it is almost confluent
with the articular ridge: there is a deep but variable depression
for the lateral depressor muscle; and in young specimens of _var._
(_a_) it was almost absent. _Tergum_: the surface exhibits traces of
longitudinal striæ: there is a deep longitudinal furrow, with the
sides folded in and quite closed in full grown specimens: the scutal
margin is considerably curved towards the scutum. The spur is long
and narrow, with the end bluntly pointed, placed at rather above its
own width from the basi-scutal angle; the basal margin slopes but
little towards the spur: the crests for the depressores are feebly
developed.
_Parietes_: their internal surfaces are strongly ribbed
longitudinally, with the basal ends of the ribs coarsely
denticulated, and with the denticuli extending close to the outer
lamina. The radii are generally narrow, but their width varies;
their summits are very oblique, smooth, rounded, and inflected, with
the lines of growth, in the uppermost part, curving inwards; their
sutural edges, in the upper inflected portion, are quite smooth,
without septa; in the lower and larger portion, the edge is crenated
with excessively fine teeth or septa, not denticulated: the radii,
like the parietes, have no inner lamina: the recipient grooves in
the opposed compartments are smooth, and are in the lower part of
the shell of unusual depth. The _alæ_, differently from the radii,
generally have their summits very slightly oblique, but sometimes
they are highly oblique: their sutural edges are most finely
crenated. The _basis_ is generally flat, sometimes cup-formed; it is
permeated by pores, crossed by transverse septa; and sometimes there
is an underlying cancellated layer.
_Mouth_: labrum with either six very small teeth, or with none.
_Mandibles_ (Pl. 26, fig. 5), with the third tooth a little thicker
than the first; fourth and fifth teeth small, but quite distinct.
_Maxillæ_ (Pl. 26, fig. 7), with the inferior part forming a square
step-formed projection, bearing, one behind the other, two spines
as long as the upper pair; in a young specimen of _var._ (_a_) this
step-formed projection was absent.
_Cirri_: first pair with the rami unequal by about four segments: the
shorter ramus has the segments very protuberant in front, thickly
clothed with strongly serrated spines; the second cirrus has segments
moderately produced; the third has them produced only in a slight
degree. The pedicels of the second and third cirri have dorsal tufts
of spines, but not a hairy plate prolonged over the thorax. The
posterior cirri have segments broader than long, bearing only two
pairs of nearly equally long spines; and between each pair there is a
small intermediate tuft. The penis has the usual basi-dorsal point.
_B. amaryllis_ is a distinct and well-defined species, more nearly
related to _B. Hameri_ than to any other form.
34. BALANUS ALLIUM. Pl. 7, fig. 7 _a_-7 _d_.
_Shell faintly tinged with purple: radii broad, with their summits not
oblique: basis not porose. Scutum with the lines of growth crenated:
tergum with the spur extremely short, truncated, broad as half the
valve._
_Hab._--Raine's Islet, Barrier Reef, Australia, Mus. Stutchbury. Hab.
unknown, attached to and coated by Porites. Mus. Brit.
_General Appearance._--Shell conical, smooth, but with the lower part
sometimes narrowly ribbed in lines corresponding with the internal
longitudinal ribs; tinted pale peach-blossom purple, owing to the
sheath being finely so coloured; or wholly white. Radii broad, white,
square on the summit, hence orifice entire, ovate passing into
rhomboidal. The parietal portion of the carino-lateral compartments
extremely narrow, about one eighth of the width of the parietes of
the lateral compartments. Basis concave, partially imbedded in the
coral. Largest specimen .35 of an inch in diameter.
There are some specimens in Mr. Cuming's collection which appear to
belong to this species, and are certainly very closely allied to it,
but not having the opercula, cannot be identified positively; the
shell is flatter, with the walls strongly ribbed up to the orifice,
which is more rhomboidal: the basis is much more cup-formed and more
deeply imbedded in the coral; but these differences by themselves are
by no means sufficiently diagnostic.
_Scutum_: the lines of growth are crenated, causing the surface to
be very obscurely striated longitudinally: the articular ridge is
very prominent, as can be best seen from the outside, and runs down
the whole length of the tergal margin with a very regular curve, and
hence differs from the articular ridge in the foregoing species. The
adductor ridge is either absent, or very indistinct, and parallel
to the articular ridge: there is a deep little pit for the lateral
depressor muscle. _Tergum_ (7 _d_), with the apex somewhat produced
or beaked, and tinged purple: external surface almost flat, without
any longitudinal furrow: scutal margin curved. Spur very short,
placed quite close to the basi-scutal angle of the valve; broad as
half the valve; lower end square. Internally, the articular ridge is
prominent only in the uppermost part of the valve: crests for the
depressores very feeble.
_Parietes_: their internal surface is very strongly ribbed
longitudinally, the ribs being coarsely denticulated at their bases,
and finely fluted along their sides. The sheath is transversely
ribbed, and clothed with an epidermis furnished with transverse
rows of fine hairs. The _radii_ are of a dead white, whereas the
parietes are translucent; the summits are parallel to the basis;
they are broad; the radii of the carino-lateral compartments appear
extraordinarily broad, owing to the narrowness of the parietal
portion: the sutural edges are furnished with coarse septa, which
are sinuous, irregular, and obtusely denticulated; the interspaces
are filled up solidly. The _alæ_ are thin, with their sutural edges
almost smooth, and their summits oblique: in some specimens, during
the diametric growth, a mere, almost thread-like ribbon is added
to their sutural edges. _Basis_ slightly cup or saucer-shaped;
moderately thick, permeated by fine pores, and generally ribbed in
lines radiating from the centre. The walls and basis adhere together
very firmly.
_Mouth_: labrum with six teeth: mandibles with five teeth; the three
upper teeth being sharp, narrow, and unusually prominent; the two
lower teeth minute and sharp; maxillæ without a notch. _Cirri_ much
injured: first pair with one ramus apparently one third longer than
the other: segments not very protuberant: the posterior cirri have
elongated segments with five pairs of spines.
_Affinities._--This species is very distinct from all the foregoing:
in the carino-lateral compartments being so narrow, and tending, as
we may suppose, to become aborted; in the form and structure of the
whole shell, and in its habits, this species shows an affinity and
passage to the coral-inhabiting sub-genus Creusia, which has only
four compartments. There is also a close affinity to the sub-genus
Acasta. This species is so closely allied to the following, that
I at one time felt some doubts whether they ought to have been
specifically separated; it is also probably closely allied to _B.
terebratus_, but the materials hardly suffice for judgment: it is
also related, though less obviously, to _B. vestitus_.
35. BALANUS CEPA. Pl. 7 fig. 8 _a_-8 _c_.
_Shell dirty reddish-purple, steeply conical: radii narrow: basis
obscurely porose. Scutum with the lines of growth crenated: tergum with
the spur truncated, broad as half the valve, and depending beneath the
basi-scutal angle as much as half its own breadth._
_Hab._--Japan, attached to an Isis, Mus. Cuming. Attached to an
oyster, Mus. Stutchbury.
As already stated, this species comes in all essential respects very
near to the last, though differing much in appearance; I have seen
two sets of specimens, and two sets of _B. allium_, and there was no
variability or passage in the points in which they differed; hence I
must consider them as specifically distinct.
_Shell_, steeply conical, strongly but bluntly ribbed longitudinally;
coloured either all over dull reddish purple, or with the upper part
only pinkish purple: in one set of specimens, the yellow epidermis
was partially persistent. Radii narrow. Orifice small, ovate. The
wall of the carino-lateral compartment is very narrow. The internal
surface of the parietes is ribbed, but finely, and only in the lower
part. The septa, on the sutural edges of the radii, are finer than in
_B. allium_. Basis flat, obscurely permeated by pores. The largest
specimen is .25 of an inch in basal diameter.
_Scuta_: these are longitudinally and finely striated; the
basi-tergal corner is more rounded off than in _B. allium_, and the
articular ridge is not nearly so prominent: internally, the adductor
ridge is rather more prominent. The _Tergum_ is rather broader: its
apex is produced into a minute sharp point: the scutal margin is
straight; the spur is broader, and measured from the basi-scutal
angle of the valve, considerably longer; namely, as long as half
the width of the basal margin of the spur, whereas in _B. allium_
it is only about a quarter as long as the basal margin of the spur:
the lower edge of the spur is not here so directly transverse to
the longitudinal axis of the valve as in _B. allium_: the external
surface is not so flat as in that species, and a depression runs down
to the basi-scutal angle of the spur.
Considering the difference in the shape and appearance of the shell,
with its narrow radii and small orifice; considering the less strongly
ribbed internal lamina of the parietes, the finer septa on the sutural
edges of the radii, the slight difference in outline in the scuta and
terga, more especially the greater length of the spur, I conceive I am
right in ranking this form as a distinct species, though assuredly it
is very closely allied to _B. allium_, and even still closer to the
following _B. quadrivittatus_.
36. BALANUS QUADRIVITTATUS. Pl. 8, fig. 1.
_Shell steeply conical, having four longitudinal gray bands placed
crosswise: radii with their summits oblique: basis thin, solid. Scutum,
with the lines of growth smooth; no distinct pit for the lateral
depressor muscle: tergum as in B. cepa._
_Hab._--East Indian Archipelago, attached to lamelliferous corals,
and associated with _Pyrgoma grande_ and _Creusia spinulosa_, Mus.
Brit. and Stutchbury and Darwin. Philippine Archipelago, attached to
a Tetraclita, Mus. Cuming.
I have seen four sets of specimens of this species, taken in four
different places, one set containing above twenty individuals, and
all resembled each other exactly: nevertheless, this species comes so
close to _B. cepa_, that I am somewhat doubtful about its specific
distinctness.
_General Appearance._--Shell smooth, or slightly folded, steeply
conical; white, with four longitudinal bands of pale brownish-gray
colour, namely, on the rostrum, the carina, and the two lateral
compartments: the carino-lateral compartments are very narrow and
almost white: the four brownish-gray bands are darkest in the upper
part of the shell, though always rather faint, and die out towards
the base: they can sometimes be seen to be formed of several narrow
longitudinal stripes; the tint shows a trace of containing purple.
The orifice of the shell is small, rhomboidal, and not quite entire,
owing to the obliquity of the summits of the moderately broad radii.
In structure, the shell, radii, and alæ resemble those in the last
species. The basis, however, does not appear to be permeated by
pores. Basal diameter of largest specimen .25 of an inch.
The _Scuta_ most closely resemble those of _B. cepa_, but the lines
of growth are not crenated, and internally there is only a very
minute pit for the lateral depressor muscle, placed almost on the
edge itself of the valve. The _Tergum_ hardly differs at all from
that of _B. cepa_, but is perhaps of rather greater breadth.
The _Mouth_ does not differ from that in the last two species. In the
_Cirri_, the three posterior pairs have elongated segments, bearing
only three pairs of spines, of which the lowest pair is minute: in
_B. allium_, and I believe in _B. cepa_, there are five pairs of
spines on each segment.
This species differs from the last only in the peculiar colouring,
smoother walls, more oblique radii, solid basis, and more especially
in the scuta having the lines of growth not crenated, and internally,
in the pit for the lateral depressor muscle being so very minute and
placed on the basi-tergal edge of the valve. The posterior cirri,
also, I believe, differ in the number of the spines which the segments
support; nevertheless, I cannot feel confident about the specific
distinctness of _B. quadrivittatus_.
37. BALANUS TEREBRATUS. Pl. 8, fig. 2 _a_-2 _b_.
_Shell white, strongly ribbed longitudinally, with the basal margin
produced into long points: basis concave, not permeated by pores, but
strongly ribbed externally in radiating lines; the interspaces between
the ribs being riddled by minute rounded apertures, often placed in
double rows._
_Hab._--Unknown, Brit. Mus., attached to a lamelliferous coral.
I have in this instance broken through my rule of not describing a
Cirripede without examining the opercular valves; but the species here
named is so peculiar, that it would have been a fault to have passed it
over. There is but a single specimen in the British Museum, without, as
just stated, the operculum, and of course without the animal's body.
_Shell_, white, depressed, conical, somewhat elongated in its
rostro-carinal axis; orifice rather small, pentagonal, toothed,
elongated. Parietes rather thin, with extremely prominent
longitudinal ribs, produced at the basal edge into long spikes: the
internal surface is also ribbed, but less strongly than the outside.
Radii rather narrow, with oblique, not smooth summits; sutural edges
very finely and obscurely crenated. Alæ with their summits extremely
oblique. Lower edge of sheath closely attached to the walls. The
carino-lateral compartments are rather narrow.
_Basis_, slightly concave or saucer-shaped; the circumference
is produced into long spikes, corresponding with those on the
basal margin of the parietes: these projections equal half the
semi-diameter of the shell. The internal surface of the basis has
slightly prominent, rounded ridges; and the external surface has
extraordinarily prominent, sharp ridges, radiating from the centre;
the edges of the external ridges are irregular, notched, and knobbed.
I have seen in no other species external ridges on the basis or
surface of attachment; and what is more remarkable, the interspaces
between the ridges are penetrated by small rounded apertures, of
irregular shape and unequal sizes; and these are generally arranged
in an irregular double row, and externally are closed by the
membrane, which clothes the basis. In the sub-genus Acasta, the basal
cup is sometimes penetrated by similar holes, but these seem never
to extend over the whole basis, and are very variable; nevertheless,
in some specimens of _Acasta spongites_ from the Cape of Good Hope,
portions of the basis closely resembled, except in the absence of
the radiating ridges, the structure here described, but the holes
were not arranged in any definite order. The internal surface of
the parietes in _Acasta sporillus_ presents a somewhat analogous
appearance, but the pits do not penetrate through the walls. This
species, I have no doubt, is closely allied to the sub-genus Acasta,
and to _Balanus navicula_ with its allies, and, but much less
closely, to _B. allium_ with its allies. Indeed, had _B. terebratus_
inhabited a sponge, I should have been compelled to have ranked it in
the sub-genus Acasta.
38. BALANUS VESTITUS. Pl. 8, fig. 3 _a_-3 _b_.
_Shell pinkish-purple or white, clothed by an orange-coloured membrane;
radii represented by mere fissures; basis solid. Scutum, with a sharp,
curved adductor ridge; with crests for the lateral depressor muscle:
tergum, with the spur short, truncated, one third of width of valve._
_Hab._--New Zealand, New South Wales, Mus. Brit. and Stutchbury;
attached to shells.
_General Appearance._--Shape conical, often steeply conical; orifice
small; radii not developed, represented by mere fissures. The walls
are smooth, or slightly, or strongly ribbed longitudinally. The shell
itself is of a fine peach-blossom pink, or nearly white, but it is
generally covered by a thick yellow or brownish-orange epidermis.
Opercular valves pinkish, but similarly covered. Basal diameter of
largest specimen .7 of an inch.
_Scuta_, with the lines of growth closely approximate; surface
somewhat convex. Internally, the articular ridge is very little
prominent, but runs far down the tergal margin; in some specimens,
however, it is shorter and more prominent. The adductor ridge is
strongly prominent, is curved towards the rostral angle, and runs
down nearly to the basal margin. The rostral depressor muscle is
lodged in a small cavity, formed, as usual, by the overlapping of
the occludent margin; within this cavity there are either tolerably
distinct little crests, or merely traces of them, for the attachment
of the muscle. The lateral depressor muscle is attached to several
quite distinct crests, seated in a concavity beneath the adductor
ridge. _Tergum_ rather narrow, with the apex produced or beaked; the
beak is purplish and flat. There is a slight rounded longitudinal
furrow, or depression. The spur is fully one third of the width of
the valve: it is short, with the end truncated, and placed close to
the basi-scutal angle; the basal margin on the carinal side slopes
gently towards the spur. Internally, the scutal margin is scarcely at
all inflected, and the articular ridge is very little prominent: the
crests for the tergal depressores are pretty well developed.
From the points here enumerated, it is clear that the opercular
valves are articulated together much less strongly than is usual with
most species, excepting _B. allium_ and its allies. It is remarkable
that in this species the terga are united to the sheath, not, as is
usual, by a single opercular membrane, but by five or six, one above
the other, the upper membranes not having been exuviated as each new
lower one was formed. The minute spines on the membrane lining the
sheath are rather larger and more numerous than is usual, and to the
base of each spine a tubulus of unusual diameter runs, imbedded in
the shell.
The _Walls_, internally, have unusually numerous, narrow,
approximate, strongly prominent, longitudinal ribs, denticulated at
their bases, and inserted into the furrows on the borders of the
basis: in old specimens these internal ribs die out in the upper
part of the walls. The _Radii_ are not developed in any of the many
specimens which I have seen, and the edges of the compartments on
both sides of each suture are equally marked by slight, irregular
ridges or knobs, answering to the septa and their recipient furrows,
in the species with ordinarily developed radii. There is very little
diametric growth, the orifice being gradually enlarged by the
disintegration of the upper ends of the walls; the _alæ_, however,
in some specimens, do grow a little along their lateral or sutural
edges, so that some little diametric growth must be effected. The
summits of the alæ are very oblique; their sutural edges are plainly
crenated. The sheath descends about half way down the walls. The
_Basis_ is flat, not permeated by pores, but deeply furrowed in lines
radiating from the centre.
_Mouth_: labrum sometimes with no teeth, sometimes with four minute
teeth; mandibles with four teeth, of which the third is blunt and
rather large; the fourth is a mere knob. Maxillæ; there is, as usual,
an upper pair of large spines (beneath which there is sometimes
a small notch), but all the lower spines, instead of standing as
usual in pairs, form a single row. _Cirri_; first pair with the rami
remarkably unequal in length, one ramus having twenty-two segments,
and being more than twice the length of the other, having only nine
segments: these segments, and likewise those of the second and third
pairs, have an inverted conical shape; and they are all less thickly
clothed with spines than is usual. The second pair is short, about as
long as the shorter ramus of the first pair, and has ten or twelve
segments. The third pair is above twice as long as the second pair,
and contains twenty-four segments: this very unusual length is owing
to the presence of numerous thin tapering upper segments, unlike
those generally found in _Balanus_, in the third pair of cirri, and
apparently serving as feelers. These upper tapering segments are
of an inverted conical shape, and support on their upper margins
two very small tufts of spines, one behind and one in front: on the
segments lower down these tufts increase in size, and the spines are
more spread out, so that in the basal segments, the tufts in front
form on the upper margin two or three crowded rows of bristles. The
three posterior pairs of cirri have elongated segments, which bear
on their upper half three pairs of spines; of these the lowest pair
is minute, and the middle pair is only one third of the length of
the upper pair. The sixth cirrus, in the same individual as before,
contained twenty-seven segments in each ramus, that is only three
more than in the third cirrus! I must observe, that the cirri in
all the specimens were irregular, often distorted and monstrous;
and therefore, probably, there is considerable variation in the
proportional numbers of the segments in the cirri.
At the base of the penis there is a minute, knife-edged, triangular
projection. The branchiæ are rather narrow, pointed, and not very
large.
_Affinities._--This is a very distinct species, as shown by the
peculiarities in the cirri, by the absence of radii to the shell,
and by the presence of crests for the attachment of the lateral
scutal depressores. With the exception of this latter character, the
opercular valves clearly show, that _B. vestitus_ is allied to _B.
allium_, _cepa_, and _quadrivittatus_. In some respects this species
manifests an affinity to _B. imperator_, which latter has its third
pair of cirri nearly similar to those of _B. vestitus_.
39. BALANUS IMPERATOR. Pl. 8, fig. 4 _a_-4 _c_.
_Shell internally imperial purple; parietes thick, with their internal
basal edges rough with irregular points and ridges; radii narrow; basis
very thin, solid. Scutum, with crests for the rostral and lateral
depressor muscles; tergum, with the end of spur rounded._
_Hab._--New South Wales, Sydney, Port Stephens, Moreton Bay; attached
to sandstone-rocks and shells, at low-water line; Mus. Brit., College
of Surgeons, Cuming, Stutchbury.
_Shell_ conical, very thick and very strong; longitudinally sulcated
more or less strongly; whole thickness of shell beautifully coloured
rich violet, or more strictly "imperial purple;" externally the
surface, from disintegration, is generally whitish; internally
the colour is best developed: the narrow radii and the thin basis
are white. The largest specimen which I have seen was one and
three-quarters of an inch in basal diameter, the walls close to the
basis being, in this instance, actually .3 of an inch in thickness.
_Operculum_ thick and strong, covered by yellowish-brown epidermis;
internally, the shelly substance is either all of the richest
purple or yellowish-white, tinged, especially in the upper part,
with purple. _Scuta_, with the apex beaked and somewhat reflexed;
articular ridge very thick, little prominent; articular furrow very
narrow, the impression made by the adductor muscle is seated very
high up the valve; there is hardly an adductor ridge, but the surface
of the valve is angularly prominent in a curved line, running from
the articular ridge to near the rostral angle of the valve. At the
rostral angle, the occludent margin is not folded inwards, as is
generally the case, but the surface is flat, and is marked by four
or five crests for the attachment of the rostral depressor muscle.
There are other crests for the lateral depressor muscle. _Tergum_,
with the apex somewhat produced and beaked, but blunt; longitudinal
furrow shallow; spur of moderate breadth, with its lower end rounded;
the basal margin on the carinal side of the spur slopes towards it.
Internally, articular ridge moderately prominent. Crests for the
tergal depressor well developed.
_Parietes_, solid, thick, with the basal internal edge (4 _c_) formed
of short ridges, or flattened and irregular points, which in very old
specimens are but little prominent; inner surface, finely, closely,
and irregularly ribbed longitudinally, but in some specimens nearly
smooth. The _radii_ are nearly white; they are narrow, sometimes
hardly at all developed, and have their summits very oblique and
jagged; exteriorly, they are sulcated in a transverse direction, and
sometimes form oblique steps, from having been formed layer over
layer: their sutural edges are formed of irregularly branching crests
or septa. The _alæ_ have their summits very oblique; their sutural
edges are thick and crenated: the part added during diametric growth
on the inner surface is smooth, and has a different appearance from
the transversely ribbed portions of the sheath, of which the alæ
form a portion. The lower edge of the sheath is hollow beneath. The
carino-lateral compartments are very narrow.
_Basis_ calcareous, thin, white, sometimes opalescent, apparently
formed by an aggregation of very minute calcareous beads, with no
trace of furrows radiating from the centre.
_Mouth_: labrum hairy, with apparently some very minute teeth;
mandibles, with the fourth and fifth teeth small and rudimentary;
maxillæ rather broad, with a narrow and rather deep notch under the
two great upper spines: outer maxillæ with the lower lobe very large.
_Cirri_: first pair, with the rami unequal by several segments:
second pair, with the rami unequal in length by about six segments:
third pair elongated, with the segments very numerous, almost
equalling those in the sixth cirrus; upper segments of both rami much
elongated, each with only a circle of spines; segments in the above
first three pairs of cirri only slightly protuberant. Posterior cirri
elongated, with the upper segments bearing three pairs, and the lower
segments four pairs of main spines, between which there is a small
intermediate tuft.
_Affinities._--This noble Balanus, in all the characters derived
from its opercular valves, and from its cirri, is closely allied to
the last species: in the structure, however, of the shell and of
the basis, it comes closer to the following, _B. flosculus_. The
crests on the under side of the scutum, for the lateral depressores,
are confined to these three species; and the crests for the rostral
depressores are confined to _B. imperator_ and _vestitus_, but they
are generally rudimentary in the latter. The internal basal structure
of the parietes is singularly like that of _Chelonobia caretta_,
though there is no other special affinity to that genus. In the
nature of basis; in the structure, to a certain limited extent, of
the walls of the shell; in the narrowness of the carino-lateral
compartments; in the elongation of the third pair of cirri; in the
crests for the rostral and lateral scutal depressores, _B. imperator_
comes nearer to the genus _Tetraclita_ than does any other species of
Balanus.
40. BALANUS FLOSCULUS. Pl. 8, fig. 5 _a_-5 _f_.
_Shell purple or dirty white, with the internal basal edges of the
parietes rough with irregular points and ridges; radii narrow or
absent; basis excessively thin, in appearance absent. Scutum with
crests for the lateral depressor muscle; tergum very narrow, with the
spur pointed._
_Var._ sordidus. (Pl. 8, fig. 5 _b_) _shell globulo-conical, dirty
white, with numerous sharp, narrow, longitudinal folds or ridges_.
_Hab._--Peru and Chile; generally attached to the _Concholepas
Peruviana_, or to _Balanus psittacus_, and associated with
_Chthamalus scabrosus_. _Var. sordidus_, inhabits Tierra del Fuego,
attached to littoral shells, wood, and rock, associated with _Ch.
scabrosus_.
_General Appearance._--Shell either extremely much depressed and
irregular, or globulo-conical, or more rarely cylindrical and
elongated. Walls, either with a few rather broad, smooth, irregular,
longitudinal folds, or, in _var. sordidus_, with numerous sharper
and more prominent longitudinal ridges; basal margin very sinuous.
Colour, either fine rich peach-blossom purple, or so pale as to be
almost white; or in _var. sordidus_ dirty white, generally stained
greenish from confervoid matter. Orifice small, oval, entire. Radii
very narrow, white, or not at all developed, and with even the
sutures not distinguishable. The purple coloured varieties, with
the narrow white radii, the small oval orifice, and folded walls,
have a very pretty appearance, which is far from the case with _var.
sordidus_. The largest specimens attained a basal diameter of .6
of an inch, but this is an unusual size: I have seen a cylindrical
specimen of _var. sordidus_ one inch in length.
The opercular valves are united to the sheath by unusually strong
membrane: internally, their upper parts are stained purple. _Scuta_,
these vary considerably in breadth, some being even broader than in
Pl. 8, fig. 5 _c_, and others as much elongated as in fig. 5 _d_:
these latter come from an elongated cylindrical shell. The valve
externally is unusually convex: the apex, also, projects freely to
an unusual degree. Internally, the articular ridge is moderately
prominent: the adductor ridge is prominent and much curved: in the
upper part it either lies close to, or at some little distance from
the articular ridge. The lateral depressores are attached to several
little crests, occupying a cavity, (often bordered above by a little
ridge) close beneath the adductor ridge. _Tergum_, extraordinarily
narrow and elongated; beak triangular, purple: a longitudinal furrow
runs down the valve: spur narrow, long, bluntly pointed, lying close
to the basi-scutal angle of the valve: the scutal margin is nearly
straight and parallel to the spur. Internally, the articular ridge
is prominent: the crests for the depressores are moderately well
developed: the upper part of the valve is marked by a purple patch,
bounded on one side by the articular ridge, and on the other side
by a very slight special ridge. There is some variability in the
narrowness of the whole valve, and in the sharpness of the spur.
_Parietes._--The under surface, in the more depressed varieties, is
roughened in a remarkable manner nearly or quite up to the sheath,
with very irregular, projecting, and branching ridges, and sometimes
with depending points. These ridges and points are granulated on
their under surfaces. The roughened surface in the more conical
varieties is confined to the basal inner margin. This structure is
nearly the same as that in _B. imperator_, represented (Pl. 8, fig.
4 _c_), but the little ridges are here more apt to be concentric,
instead of radiating. The lateral edges of the compartments on the
inside, especially the carinal edges, project inwards beyond the
inner surface of the shell. The sheath is but little hollow on its
under side. The diametric growth of the shell seems to be quite
capricious; in the same group, some individuals thus increasing, and
others not at all. When the radii are developed they are narrow and
white, with their upper margins nearly if not quite parallel to the
basis: their sutural edges are formed by large, irregular, branching
teeth or septa. The _alæ_, also, have their sutural edges coarsely
crenated; and when the shell increases by diametric growth, they are
added to above the opercular membrane, so as to be nearly square at
top.
_Basis._--When a shell is removed from the surface of attachment,
and inspected even under a lens, there appears to be no basis
whatever, either adhering to the shell, or to the supporting surface:
but when a more careful examination is made, with a higher power,
an excessively thin, translucent, calcareous, irregular layer, or
rather film, can almost always be discovered. This would be more
easily distinguished if the specimens had adhered to rock instead of
to the rugged shells of molluscs. At one time I thought the basis
was partially membranous, for I have certainly scraped off small
fragments of membrane from the supporting surface; but these, when
examined under the compound microscope, seemed always to consist of a
thin sheet of the yellow cementing tissue; and in some instances, a
brittle film of shell, representing the true basis of the cirripede,
still adhered to the upper surface of the membranous bits of cement.
Nevertheless, so imperfect is the calcareous basis, that I should not
be surprised if portions of a true membranous basis did really in
some cases exist.
_Mouth_: labrum with the notch wide, generally with a few little
teeth; mandibles with three teeth, and some inferior knobs; maxillæ
notched. _Cirri_, first pair with one ramus shorter by three segments
than the other ramus. Second and third pairs short, of nearly
equal length, thickly clothed with spines; segments very little
protuberant. Posterior cirri, having elongated segments, supporting
seven pairs of spines.
_Var. sordidus._--This form is very common on the tidal shores of
the Strait of Magellan, and of the southernmost parts of Tierra del
Fuego, near Cape Horn: it lives attached to rocks, mytili, and logs
of wood, and is associated with _Chthamalus scabrosus_. It almost
certainly is the most antarctic form of the genus Balanus. If I were
guided by external appearance alone, I should certainly separate
this form specifically from _B. flosculus_, but, as will be seen in
the following description, the differences consist only in _var.
sordidus_ being much duller and rather differently coloured, in the
longitudinal folds being sharper and more prominent, and in the whole
shell being rather more globular, and on an average rather larger;
but in the true _B. flosculus_ there is considerable variation in
all these respects, as there likewise is in _var. sordidus_; thus
some of the cylindrical varieties of the latter have less prominent
ridges than even _B. flosculus_. In general appearance I have seen
some _nearly_, but not exactly, intermediate forms; therefore, I do
not feel positive that these forms may not be specifically distinct,
but have failed, after careful examination, to find any sufficient
diagnostic _characters_. Moreover, in the case of _Balanus lævis_,
I was led to believe that there is an equal and somewhat analogous
amount of variation in the specimens inhabiting Tierra del Fuego and
northern Chile; and in this case I was enabled to show the existence
of strictly intermediate forms in the intermediate districts.
The shell in _var. sordidus_ is generally globulo-conical, dirty
white, frequently with a green tinge, from the growth of confervoid
matter. Orifice small. The exterior surface is covered with numerous
prominent, narrow, sharp ribs or folds, the basal margin being
serrated with projecting points where the folds terminate. When
the radii are not developed, the sutures are very often obscure.
Internally, the shell is faintly tinted of a port-wine purple. In
all points of structure this form is identical with the true _B.
flosculus_. In some few specimens the whole exterior surface was
disintegrated and smooth; and this is generally the case with the
upper parts of the shell. Some other specimens, which had grown
crowded together on wood, had become cylindrical, and consequently
the orifice was as large in diameter as the shell, namely, half an
inch: in some of these cylindrical varieties the sheath was entirely
soldered to the walls. The largest specimens which I have seen were
.6 of an inch in diameter; and above one inch in height.
_Affinities._--This species, in its opercular valves, even in such
trifling characters as the strength of the opercular membrane,
and in its cirri, approaches closely to _B. cariosus_. We even
see on the under side of the scutum, in that species, a single
little ridge, foreshadowing, as it were, the crests for the lateral
scutal depressores, so remarkable in our present species. In the
structure of the shell and of the basis, _B. flosculus_ is much
more closely related to the last species, or _B. imperator_. If it
had been possible to have arranged the species in a single line,
_B. flosculus_ ought undoubtedly to have been placed between _B.
cariosus_ and _imperator_.
41. BALANUS BISULCATUS. Pl. 8, fig. 6 _a_-6 _c_.
BALANUS SULCATINUS (?) _Nyst_, apud D'Omalius (sine descript. aut
tabulâ), Geologie de Belgique, 1853.[100]
[100] I am indebted to M. Bosquet for a specimen, bearing this
name and reference, found in the 'Systeme Bolderien' of Dumont,
(miocene according to Sir C. Lyell) at Bolderberg. The specimen
consists of a rostrum, with a portion of the base attached; and as
these parts are in some degree characteristic, I fully believe this
specimen to be the _B. bisulcatus_. I hope hereafter to give in
the Palæontographical Series fuller illustrations of this and the
following fossil species.
_Radii with their upper margins oblique and smooth; sutural edges
smooth: basis permeated by large pores. Scutum narrow, with from two to
four longitudinal furrows: tergum with the spur very short, broad as
half the valve._
_Var._ plicatus, _with the walls deeply folded; radii narrow, with
their upper margins very oblique_.
_Fossil_ in Coralline Crag, Ramsholt, Gedgrave, Sutton; Mus. S. Wood,
Bowerbank, J. de C. Sowerby. Rauville, dans le Cotentin, Mus. G.
B. Sowerby. _Var. plicatus_, Coralline Crag, Sutton, Mus. S. Wood.
Bolderberg, near Hasselt, Belgium, Mus. Bosquet.
_General Appearance._--Shell conical or tubulo-conical, often rather
globose; walls frequently thin, either very smooth, or deeply
plicated longitudinally: occasionally the same specimen is smooth
in the upper part, and strongly plicated in the lower. The Radii in
the larger specimens are wide, and with their upper margins only
slightly oblique; in the smaller, they are narrower and much more
oblique, but in each case their upper margins are smooth and slightly
bowed. Colour apparently originally nearly white, but with the alæ
generally, in the smaller specimens, clouded with a dark tint:
the radii are usually striped feebly in longitudinal lines. Basal
diameter of largest specimen .8 of an inch; but this seems to have
been an unusual size.
_Scuta_: narrow, with the basal margin forming an unusually small
angle with the occludent margin; surface slightly convex, with lines
of growth approximate, moderately prominent; on the tergal half
of the valve, two distinct rather broad furrows, with sometimes a
third, and even a fourth, nearer to the occludent margin (Pl. 8,
fig. 6 _a_), extend from the apex down the valve, and give it a very
peculiar appearance: the furrows near the tergal margin are the
deepest. Internally, the upper part of the valve is roughened with
small points: the articular furrow is unusually wide: the articular
ridge is very prominent and but little reflexed, with the lower end
almost abruptly cut off: the adductor ridge is prominent, but short:
there are small deepish pits for the rostral and lateral depressores.
_Terga_ (6 _b_), broad, flat, with a slight narrow prominent rim
along the scutal margin, which margin is slightly bowed. The basal
margin on the carinal side of the spur slopes so gradually towards
the spur, that the latter is barely distinct, and is very short, not
depending nearly half its own width beneath the basi-scutal angle: it
is broad, namely, measured across the upper part, as broad as half
the valve; its basal end is obliquely rounded off on the carinal
side; it is placed close to the basi-scutal angle. The carinal
margin of the valve is just perceptibly bowed, and is formed by
rectangularly upturned lines of growth. Internally, the upper part of
the valve is rough; the articular ridge is prominent; the crests for
the tergal depressor muscles are moderately well-developed.
_Parietes_, not porose; internally, the ribs are smooth, with their
basal edges very finely or barely denticulated. The _radii_ (as
already stated) are of variable breadth; they have their upper
margins either very slightly or highly oblique, but always smooth and
rounded: their sutural edges are quite smooth, or sometimes, with
a strong lens, traces of transverse striæ, representing septa, can
just be discovered. The _alæ_ have their upper margins very oblique;
their sutural edges are, in the large specimens, quite smooth; in the
younger ones, plainly crenated; the recipient furrow being clearly
marked with these teeth. _Basis_ plainly porose.
_Varieties._--It is certain that there are longitudinally plicated
specimens of this species, and that the obliquity of the upper
margins of the radii also varies a little: nevertheless some of the
deeply plicated specimens undoubtedly have a very different aspect
from the ordinary varieties, and do really differ in the sutural
edges of the alæ being crenated, and in the greater narrowness and
obliquity of the radii; but these points are all commonly variable.
I have not seen any large specimens of the variety, _plicatus_, so
as to compare them with the large specimens of the normal form, yet
I can hardly entertain any doubt, considering their agreement in
so many important points, that I have rightly treated these forms
as mere varieties; it is unfortunate that none of the specimens of
the _var. plicatus_ seen by me have had opercular valves, as their
presence would have removed all shadow of doubt.
_Affinities_: this is a strongly characterised species, and nearly
allied only to the following species, _B. dolosus_. The furrows on
the scuta in some degree resemble those on _B. lævis_, but there is
no alliance with that species. It is certain that amongst recent
species, the chief affinity is with _B. Hameri_ and _amaryllis_.
42. BALANUS DOLOSUS. Pl. 8, fig. 7.
_Radii with their upper margins oblique and smooth; sutural edges
smooth: basis permeated by large pores. Tergum with the spur not very
short, broad as one third of valve._
_Fossil_ in Red and Mammaliferous Crag, England; Mus. S. Wood,
Bowerbank, Lyell, J. de C. Sowerby, Henslow, &c. Mammaliferous Crag,
Postwick, near Norwich, Mus. Lyell.
This species so closely resembles _B. bisulcatus_, both externally and
in all the essential characters of the parietes, radii, and basis,
that it is quite superfluous to describe over again these parts. The
specific characters are derived from the opercular valves, which
present well defined distinctions, found by me constant in several
specimens of both species. _B. dolosus_, like _B. bisulcatus_, has
quite smooth and deeply plicated varieties, often adhering to the same
univalve. The ribs on the inner surfaces of the parietes are remarkably
prominent. I think the upper margins of the radii are in this species
rather more oblique than in _B. bisulcatus_. The sutural edges of the
radii are marked by the finest striæ, representing septa. The sutural
edges of the alæ are generally distinctly crenated. The basis is often
slightly cup-formed, and very plainly porose: its surface is marked by
radiating ridges. The orifice of the shell is large, and elongated,
especially in young specimens. The basal diameter of the largest
specimen is .4 of an inch.
The _Scuta_ have no trace of the two or three longitudinal furrows so
conspicuous on these valves in _B. bisulcatus_, and which, in that
species, run down from the apex of the valve; this fact showing that
the furrows occur in quite young individuals. The whole valve is not
quite so narrow as in _B. bisulcatus_, but otherwise agrees with it
in shape: internally, there is hardly any difference: the articular
furrow is not so wide: the articular ridge is very prominent, and
abruptly truncated at its lower end: the adductor ridge is also
prominent; it here runs a little higher up the valve than in _B.
bisulcatus_. The _Tergum_ differs more in the two species: the spur
is not so broad; measured in its upper part, it is only about one
third of the entire width of the valve, instead of being half as
wide as the valve: it is considerably longer, depending beneath the
basi-scutal angle more than half its own width: the basal margin of
the valve on the carinal side, does not slope so gradually into the
spur: the occludent and carinal margins are slightly arched, as in
_B. bisulcatus_. Internally, the surface is rough, the articular
ridge is prominent, and the crests for the tergal depressores are
well developed,--all as in _B. bisulcatus_. It is remarkable, how
generally the opercular valves have been preserved in this species
in its fossil condition, as compared with most other species of the
genus.
It is not easy to distinguish by external characters the rugged
varieties of this species from _B. crenatus_; indeed, the only
difference is that the furrows receiving the edges of the radii,
generally, exhibit in _B. crenatus_ a slight impression of the septa,
which are entirely absent in _B. dolosus_. By internal characters,
such as the non-porose parietes, and porose basis, our present species
widely differs from _B. crenatus_.
43. BALANUS UNGUIFORMIS. Pl. 8, fig. 8 _a_-8 _b_.
BALANUS UNGUIFORMIS. _J. de C. Sowerby_ (!). Mineral Conchology (sine
descriptione) Tab. 648, fig. 1, (Jan. 1846).
---- ERISMA. _J. de C. Sowerby_ (!). Ib., fig. 2.
---- PERPLEXUS. _Nyst_, apud D'Omalius (Sine descript. vel Tab.),
Geologie de la Belgique, 1853.[101]
[101] I am much indebted to M. Bosquet for specimens bearing this
title, from Klein Spauwen, which certainly appear to me, as far as
can be judged by the separated compartments, without the opercular
valves, to belong to our present species.
_Parietes thin, sometimes permeated by pores; radii with their upper
margins oblique; sutural edges very finely crenated: basis solid.
Tergum with the spur narrow, bluntly pointed._
_Var._ erisma, _with the walls longitudinally folded or ribbed_.
_Fossil_ in the Eocene formation, Isle of Wight, Colwell Bay;
Hordwell; Barton, (Chama Bed); Headon; Bembridge. Bergh, near Klein
Spauwen, Belgium (?). Attached to various shells and wood. Mus. J. de
C. Sowerby, E. Forbes, F. Edwards, Charlsworth, T. Wright, Bowerbank,
Tennant, Bosquet.
This species, the most ancient one as yet well known in the genus,
presents to the systematist a most unfortunate peculiarity, in the
parietes being almost as often as not permeated by small pores: I have
seen no other instance, except to a limited degree in _B. glandula_, of
this character being variable, and hence it must be still considered
of high classificatory value, in so varying genus as Balanus. Owing
to this varying condition of the parietes, together with the basis
being quite solid, our present species has as good a claim to be
ranked in the last as in the present section; indeed, I think it has
more affinity to _B. crenatus_ and _glandula_ in the last section,
than to any other recent forms: I have placed it in its present place,
owing to its intimate affinity to _B. varians_, in which the parietes
seem always to be solid; and partly, I believe, because all the first
specimens examined by me exhibited no traces of parietal pores.
Owing to the kindness of Mr. F. Edwards, I have seen the original
specimens, excellently figured by Mr. J. de C. Sowerby in the 'Mineral
Conchology:' I can perceive no difference between _B. unguiformis_ and
_erisma_, excepting that the walls in the latter are longitudinally
folded,--a character we know to be variable in so many species. In both
varieties, the parietes are sometimes porose and sometimes solid. The
smaller specimens, however, figured in the 'Mineral Conchology' to the
right hand, may possibly be a distinct species, as I infer from the
narrowness of their radii.
_General Appearance._--Shell, tubulo-conical, sometimes considerably
elongated and sub-cylindrical: surface either very smooth, or
slightly folded, or deeply folded so as to be strongly ribbed
longitudinally: orifice rather large, rhomboidal, narrow at the
carinal end, toothed, but not deeply: walls rather thin and fragile:
radii of moderate width, with their summits oblique, not quite
smooth. Basal diameter of largest specimen, about three quarters of
an inch.
_Scuta_, with the external surface smooth: there is a trace of a
furrow running down the valve from the apex, near to the occludent
margin, and this is only worth mentioning from the analogous furrows
in _B. bisulcatus_. Internally, the upper surface of the valve is
roughened: the articular ridge is very prominent, and slightly
reflexed: there is no distinct adductor ridge; there is a slight but
variable depression for the lateral depressor. _Tergum_, with the
longitudinal furrow shallow; spur moderately long, about one fourth
or one fifth of the width of the valve; placed at about its own width
from the basi-scutal angle; basal end bluntly pointed; the basal
margin on the opposite sides of the spur forms a nearly straight
line; the carinal margin has an extremely narrow border formed by
upturned lines of growth. Internally, the surface is roughened with
little points: the articular ridge is prominent: the crests for the
tergal depressores moderately prominent.
_Parietes_: the longitudinal ribs on the internal surface are either
feebly, or, in the lower part, strongly developed; their basal ends
are only just perceptibly denticulated. As already stated, in about
half the specimens, there were no traces of parietal pores; in the
other half there were either distinct or obscure pores; the pores
are circular, generally of unequal sizes, and never large; in the
same individual they would sometimes be almost wholly absent in some
of the compartments, and quite plain in the other compartments. The
Radii are either moderately wide or rather narrow, and have their
upper margins very oblique, and not distinctly arched, and not quite
smooth: their sutural edges are very finely crenated, the teeth or
septa not being denticulated. The upper margins of the _alæ_ are
rather less oblique than those of the radii: their sutural edges are
barely crenated. The _basis_ is thin, and without any trace of pores;
the upper surface is sometimes furrowed in radiating lines.
44. BALANUS VARIANS. Pl. 8, fig. 9.
B. VARIANS. _G. B. Sowerby_, in Darwin's Geolog. Observ. on South
America, (Sept. 1846), Tab. 2, fig. 4, 5, 6.
_Parietes moderately thick: radii with their upper margins very
oblique; sutural edges almost smooth, or finely crenated: basis finely
porose. Tergum with the spur small, narrow, bluntly pointed._
_Hab._--Port St. Julians, Patagonia; ancient Tertiary formation.
Eastern plain of Tierra del Fuego (?).
This species comes so close to _B. unguiformis_, that I have some
doubt whether they ought to be specifically separated: the whole shell
is stronger, and the basis can be seen to be porose when a polished
section is made: the spur of the tergum is smaller, more pointed and
more medial, but these latter differences may be due to mere variation.
Should _B. varians_ and _unguiformis_ prove to be the same species, the
latter name has the priority.
_General Appearance._--Shell moderately strong and thick; shape
conical or tubular, or even inverted conical; orifice moderately
toothed, large, sub-trigonal; walls either smooth or longitudinally
folded; the elongated specimens are most apt to be smooth. The
Radii are narrow and oblique. Diameter of largest specimen above
three-quarters of an inch.
_Scuta_, with the lines of growth moderately prominent; the internal
surface of the valve has been ill preserved; but a very prominent,
hardly reflexed, articular ridge, can be distinguished, as well
as the absence of an adductor ridge. _Terga_, with no distinct
longitudinal furrow running down the valve: spur short, bluntly
pointed, narrow, about one fifth or one sixth of width of valve;
placed at above its own with from the basi-scutal angle; the
basal margin, on each side close to the spur, curves towards it.
Internally, all that can be distinguished, is that the articular
ridge is prominent.
_Parietes_; their inner surfaces appear to have been nearly smooth;
the absence of parietal pores could be made out only by polishing
a transverse section. The Radii are narrow, and have their upper
margins very oblique and rather smooth: in the elongated varieties
the sutural edge appears to be almost absolutely smooth; in the
conical specimens it is slightly crenated, the septa being apparently
not denticulated. In living species we have a similar variation in
the state of the sutural edges of the radii, in _B. balanoides_ and
_crenatus_ the edges being much smoother in much elongated specimens
than in other varieties. The _alæ_ have their upper margins less
oblique than those of the radii, with their sutural edges barely
crenated. The _basis_ is either flat, or, in the elongated specimens,
deeply cup-formed; in section it can be seen to be finely and
irregularly porose.
45. BALANUS INCLUSUS. Pl. 8, fig. 10 _a_-10 _c_.
_Shell reddish-brown: radii broad, with their upper margins not
oblique, or only moderately oblique; sutural edges with plainly
denticulated septa: basis porose. Scutum without an adductor ridge;
tergum with the spur rather narrow._
_Var._ (_a_) (Pl. 8, fig. 10 _b_, 10 _c_), _with the shell elongated
in its rostro-carinal axis; basis narrow, clasping the stem of a
zoophyte; lateral compartments much broader than the almost linear
rostrum, carina, and carino-lateral compartments_.
_Var._ (_b_), _with rough longitudinally folded walls, and with the
summits of the radii forming an angle of about 45° with the basis_.
_Fossil_ in Coralline Crag, Sutton and Gedgrave; attached to
foliaceous Bryozoa; Mus. S. Wood, Bowerbank. _Var. a_, Coralline
Crag, Sutton, attached to cylindrical branches of corals; Mus. S.
Wood. _Var. b_, attached to shells, Osnabruck, Hanover, Mus. Lyell;
Bünde, Westphalia, Mus. Krantz.
My materials consist of a beautiful series of specimens in Messrs.
Wood and Bowerbank's collections; but unfortunately only a single
young specimen had its opercular valves preserved. Not one specimen
of the very curious variety (_a_) had opercular valves, yet I
cannot feel any doubt about its being only a variety, caused by its
attachment to a thin cylindrical branch of a coral, instead of to a
foliaceous Bryozoon; it will, however, be convenient to give a separate
description of this very remarkable form. With respect to var. (_b_),
both sets of specimens came to me with the name _B. stellaris_, of
Bronn; but as Bronn distinctly states, that in his species the parietes
are porose, and as such is not here the case, this cannot possibly be
that species: these specimens did not possess their opercular valves,
and therefore cannot be identified with certainty.
_General Appearance._--Shell conical, with the orifice rather large,
and rhomboidal. The surface is very smooth, except in var. (_b._)
from the Continent, in which it is rugged and longitudinally folded.
The colour is ochreous-brown (chiefly no doubt derived from the
imbedding substance) tinged with red. The radii often have a much
darker and more distinct red tint; they are sometimes longitudinally
striped with dirty white. The radii are broad, with their summits
straight, and very slightly oblique; in _var. b_, however, they slope
at an angle of about 45°. Basal diameter of largest specimens .6 of
an inch; but this is an unusual size.
_Scuta_ (from a young individual), with the growth ridges little
prominent. Internally the articular ridge is moderately prominent,
with its lower end very obliquely rounded off; there is no adductor
ridge; there is a minute pit for the lateral depressor muscle.
_Terga_, with a slight longitudinal depression extending down to the
spur: spur short, with its lower end almost square or truncated,
about one fourth of width of valve, and placed at about half its own
width from the basi-scutal angle. Internally, the articular ridge is
prominent; the crests for the tergal depressores are feebly developed.
_Parietes_, moderately thick and generally strongly ribbed
internally, without parietal pores. _Radii_, wide, with their upper
margins straight, not smooth or rounded, and very slightly (or, in
_var. b_ moderately) oblique; their sutural edges have well-developed
septa, which are denticulated: the interspaces between the septa
are filled up solidly. The _alæ_ have their upper margins oblique:
they are only slightly, and sometimes not at all, added to above the
level of the opercular membrane: their sutural edges are smooth. The
_basis_ is thin, but plainly porose.
_Var._ (_a_).--With respect to this remarkable variety, any one
would at first think it specifically distinct. The shell is much
compressed, or elongated in the rostro-carinal axis, sometimes to
a great degree; I have seen a specimen .25 of an inch in this axis
and only .1 in its broadest part; but this is a very unusual degree
of elongation. The most remarkable character is the extraordinary
narrowness of the carina, the carino-lateral compartments, and of the
rostrum, compared with the great breadth, especially along the basal
margin (Pl. 8, fig. 10 _b_, 10 _c_), of the lateral compartments.
The radii are of unusual breadth. The tips of the rostrum and of the
lateral compartments are a little arched in, tending to make the
shell somewhat globular. The true basis is extremely narrow (fig.
10 _c_): it is deeply grooved, from clasping the thin, cylindrical
stem of the coral to which it had adhered; and I have seen specimens
in which the opposite edges of the groove had met, a tube having
been thus actually formed. From the grooved basis, and from the
elongation of the shell in the rostro-carinal axis, this species
presents so close a general resemblance to _Balanus calceolus_, and
its allies, that I have seen it in a collection arranged on the same
tablet with a fossil specimen of _B. calceolus_. Notwithstanding
the above several strongly-marked characters, by which this variety
differs from the ordinary form, there is a resemblance in colour and
aspect, which though difficult to be described, made me from the
first suspect that the two were specifically identical. In no point
of real structure is there any difference, excepting that, perhaps,
the pores in the basis are here rather smaller; but this might arise
from the little development of the peculiar basis. Having come to
this conclusion, I was interested by finding a specimen in Mr. Wood's
collection, which had originally fixed itself (judging from the form
of the basis) on a cylindrical stem, but which had subsequently grown
on to an adjoining flat surface; consequently, one side of the shell
presented all the peculiar characters of the present variety, whereas
the other side, at the rostral end, was undistinguishable from the
ordinary form. The unequal development of the rostrum on the two
sides was very striking, and clearly showed how great an effect could
be produced by the nature of the surface of attachment.
This singular variety cannot be considered accidental, in the sense
in which this term may be applied to some varieties: the larva
evidently fixes itself intentionally, in a certain definite position,
on the branch of the coral (when a branch is chosen), exactly as in
the case of _Balanus calceolus_, or _Scalpellum vulgare_. But when
other species of Balani occasionally fix themselves on branched
corals, their position seems to be accidental and unsymmetrical; thus
among the symmetrically elongated specimens of the present species, I
found one specimen of _Balanus bisulcatus_, which had evidently been
attached in an almost transverse position to a branch, and had thus
become much distorted; so, again, I have seen specimens of the recent
_B. amaryllis_ attached irregularly to a Gorgonia, in the midst of
the symmetrically elongated shells of _Balanus navicula_, an ally of
_B. calceolus_.
This variety does not seem to attain so large a size as the ordinary
form.
_Affinities._--This species is allied to the two last described
fossils, namely, _B. varians_ and _unguiformis_, but is perhaps
more nearly related to the recent _B. allium_, an inhabitant of
the Barrier Reef of Australia. The longitudinally folded variety
(_b_) can hardly be distinguished by external aspect, or even
by the opercular valves, from _B. crenatus_; but when the shell
is disarticulated, the porose walls and non-porose basis of _B.
crenatus_, allow of no mistake in the diagnosis of the two species.
2. _Sub-Genus_--ACASTA. Pl. 9.
ACASTA. _Leach._ Journal de Physique, tom. lxxxv, 1817.
_Compartments six; parietes and basis non-porose: basis calcareous,
cup-formed, not elongated, attached to sponges, or rarely to the bark
of Isis._
Distribution mundane; imbedded in sponges and the sponge-like bark of
Isis.
This sub-genus, in one sense, is a very natural one, inasmuch as all
the species are closely allied in essential structure, in general
appearance, and in habit. On the other hand, in the structure of the
shell, in all the characters derived from the opercular valves and
animal's body, Acasta cannot properly be distinguished generically from
some species of Balanus; thus _B. navicula_ and _cymbiformis_ agree
in the parietes and basis not being porose and in all other essential
respects, differing only in the shell being more elongated in the
rostro-carinal axis and in being attached to Gorgoniæ instead of to
sponges; yet we shall see that _Acasta purpurata_ lives imbedded in
the bark of Isis, so that even the habit of being imbedded in sponges
fails. _Balanus terebratus_ would have been ranked as an Acasta had it
inhabited sponges. On the other hand, some species of Balanus inhabit
sponges, as is often the case with _B. spongicola_, and always with
_B. declivis_: but both these species are distinguished easily from
Acasta, the former by its porose walls and basis, and the latter by
its membranous basis; it may, however, be reasonably doubted whether
such differences ought to be considered as even sub-generic. The most
important character of Acasta probably consists in the anterior ramus
of the fourth pair of cirri, differing slightly in the arrangement
of its spines, and in some other points, from the rami of the two
posterior pairs of cirri,--a character not as yet observed in any
other cirripede. Had not the genus Acasta been already founded and
extensively admitted, certainly I should not have formed it; but
considering the close similarity in habits, aspect, and structure, of
the nine species of Acasta, and considering the already large size of
the genus Balanus, I hope I may stand excused for admitting Acasta as a
sub-genus.
_General Appearance._--The shape varies from nearly globular to that
of a somewhat flattened acorn, the orifice being often a little
contracted from the inward curvature of the tips of the parietes.
In _A. spongites_, however, the orifice is generally widely open;
and, on the other hand, in _A. sporillus_, the orifice is reduced
to a mere pore. The usual tint is pale reddish, but _A. purpurata_
is purple, and _A. sporillus_ purplish-brown. The surface is either
smooth, or is shagreened with minute points, as in _A. sporillus_,
and _fenestrata_, and in some specimens of _A. sulcata_; and in all
the species, except _A. sporillus_ and _fenestrata_, many individuals
are furnished with elongated, curved, sharp, shelly points, like
those in _var. spinosus_ of _Balanus tintinnabulum_. The summits of
the radii, which are generally of moderate breadth, are more or less
oblique; their surface is often marked by lines parallel to the basis,
instead of by vertical lines corresponding with the lines of growth,
as in most species of Balanus. The carino-lateral compartments vary in
proportional breadth in the different species: in _A. sporillus_, they
tend to become rudimentary, and in this species (Pl. 9, fig. 9 _b_)
their basal margins, or rather points, do not reach down to the basis.
The species are all small, _A. glans_ and _undulata_ are the largest,
being sometimes .55 of an inch in basal diameter.
_Opercular Valves._--These differ in no generic respect from those
of Balanus. The _Scuta_ are striated longitudinally in several of
the species: the adductor ridge is barely developed in any, being
most prominent in _A. cyathus_. The articular ridge is prominent in
_A. fenestrata_ and _purpurata_. In the _Terga_, the spur is either
truncated and very broad, or moderately narrow and bluntly pointed:
the surface of the valve is often depressed, and in _A. spongites_
and _fenestrata_ it is furrowed in the line of the spur. The articular
ridge and furrow are well developed in _A. fenestrata_ and _purpurata_.
The crests for the tergal depressor muscles are either absent or very
feebly developed.
_Structure of the Parietes and Radii._--The parietes are not porose;
internally, they are either smooth, or slightly, or strongly ribbed in
longitudinal lines; the presence of these ribs, which are homologous
with the parietal septa in Balanus, is variable even in the same
species. In _A. sporillus_ the inner surface is curiously reticulated.
The sutural edges of the radii are either smooth, or very slightly
crenated by the septa, in lines parallel to the basis. The upper
margins of both radii and alæ are always more or less oblique. The
radii sometimes do not extend down to the basis; and in this case, as
will presently be described, apertures are left in the lower half,
between the compartments. In _A. glans_ and _lævigata_ the internal
margin of the wall of each compartment, from the sheath to the basis,
projects inwards, forming inside the shell as many double ridges (Pl.
9, fig. 5 _b_), as there are compartments, namely, six: a nearly
analogous structure occurs in certain species of Balanus. The basal
edge of the sheath, in most of the species (5 _b_, 9 _b_), depends
freely, and is hollow beneath, but this is always a variable point.
_Basis._--The base is either saucer or cup-shaped, but in _A. cyathus_
it is almost flat; it is generally symmetrical and smooth, with the
lines of growth closely approximate. In _A. fenestrata_ the basis is
commonly as deep, as the shell is high. The edge, in several of the
species, is crenated with minute teeth or notches; and these are so
large in some specimens of _A. sulcata_ and _cyathus_, as to make the
edge almost pectinated. In _A. glans_, and in a lesser degree in _A.
lævigata_, there are six knob-like teeth (fig. 5 _a_), corresponding
with the points of junction, between the basal edges of the inwardly
prominent margins of the six compartments, and the basis: in those
specimens, in which the six teeth are largely developed, six ridges
produced by their successive development, extend down towards the
centre of the basal cup. When the basal cup is dissolved in acid,
there is but a little animalised tissue and an external membrane,
formed as usual in slips, and furnished with blunt little external
points (apparently representing spines), each of which has a short
tubulus extending to the corium. Although I dissolved the basis of
three specimens, I could not distinctly make out any cement; nor did I
see any cement-ducts; yet these are readily distinguished, after the
dissolution of the basis in acid, in Balanus, Elminius and Tetraclita.
There can be no doubt that the young shell must at first be cemented
to a fibre of the sponge; but I suspect that the cementing-tissue is
not subsequently formed, owing to the support afforded by the growth
of the enveloping sponge. As some species of Balanus are habitually or
occasionally imbedded in sponges, it is important to observe, that the
species of Acasta are not only imbedded, but are attached to the fibres
of the sponge: but even this character, as we have already seen, is
not sufficient to distinguish the genus Acasta from Balanus, for _B.
declivis_ is attached exclusively to sponge.
_Perforations in the Shell._--Calcification seems often to fail to a
certain extent in this genus: the basal cup in most specimens of _A.
spongites_, and in some of _A. glans_ and other species, is irregularly
perforated by numerous minute orifices, closed only by the external
membrane, and filled up inside by pulpy corium. In some specimens of
_A. spongites_, from the Cape of Good Hope, parts of the basis were
riddled like a sieve. I have seen similar perforations in the parietes
of a few specimens of _A. glans_. In some specimens of _A. sulcata_,
the radii do not extend quite down to the basal edge of the walls
(Pl. 9, fig. 2 _a_), and in consequence a small cleft, closed only
by membrane, is left between the compartments, for a little space
above the basal cup. In _A. fenestrata_ (fig. 7 _a_), and in a lesser
degree in _A. purpurata_ (8 _a_), not only do the radii not extend
to the basal cup, but the parietes either on one or both sides of
each suture are hollowed out, so that six, large or small, elongated,
membrane-covered openings are formed, which extend from beneath the
sheath down to the basal cup. These openings, which I have not seen
in any other genus, will be more fully described under the respective
species.
_Mouth._--The parts of the mouth are identical in the several species,
and present no generic differences from those in Balanus. The outer
maxillæ and palpi appear unusually prominent: the labrum is deeply
notched, with no teeth, or very obscure teeth on each side. The
mandibles have five teeth, but the fifth is sometimes confluent with
the inferior angle. The maxillæ are not notched; and carry one or two
spines, near their inferior angle, nearly as large as the upper pair.
The outer maxillæ are bilobed.
_Cirri_: in the first pair, the rami are very unequal in length, the
one ramus being from half to one-third of the length of the other.
The segments in the second and third pairs, are not so much broader,
or so much more crowded with bristles, in comparison with the three
posterior pairs, as is the case with most species of Balanus. The
three posterior pairs, except in _A. purpurata_, are much elongated,
and the long thin segments bear four, and sometimes only three, pairs
of spines, which are generally doubly and finely serrated, or even
feathered. The most remarkable fact respecting the cirri, is, that in
_A. spongites_, _sulcata_, _cyathus_, and _glans_, the fourth pair,
instead of being identical in structure, as in all other genera, with
the fifth and sixth pairs, has, on its anterior ramus, the pairs of
spines more crowded together, with the little intermediate spines, and
those in the dorsal tufts, a little longer than in the sixth cirrus;
and between the pairs of spines, there occur some straight, upwardly
pointed, very minute, and very thick spines or teeth. And, what is
still more remarkable (as will hereafter be described in detail), in
certain specimens, but not in all, of _A. sulcata_, the front surfaces
of the lower segments on the anterior ramus, are developed into thick,
small, downwardly curved, hook-like teeth; this likewise is the case
with the upper segment of the pedicel (Pl. 29, fig. 2),--a most
elegant, mandible-like organ for the prehension of prey being thus
formed. The variability of such beautifully contrived teeth is very
surprising. Some similar teeth occur on the segments of the anterior
ramus of the fourth cirrus, but not on the pedicel, in _A. cyathus_. A
few teeth resembling the above, but thicker, occur on the segments of
the anterior ramus of the same cirrus, in _A. purpurata_.
_Branchiæ, &c._--In _A. spongites_, I found the branchiæ rather small,
with transverse plications. The muscles of the sack, which run to the
opercular valves, seemed rather feeble in most of the species. The
penis in several species was remarkably long, and in _A. spongites_ I
noticed the straight projecting point at its dorsal base, as is common
in Balanus.
_Affinities, &c._--At the commencement of the description of the genus,
I gave my reasons for keeping Acasta distinct as a sub-genus from
Balanus. The species are particularly troublesome to identify, not only
from the great variability of the most obvious characters, but from
the very close general external appearance of most of the species, and
the consequent necessity for cleaning and disarticulating at least one
specimen in every group. The shape, however, of the shell and basis,
and the state of their disarticulated edges, whether smooth, crenated,
or toothed, here offer more serviceable, though still very variable,
characters for the identification of the species, than is usual with
sessile cirripedes; and this, probably, is in part due to the almost
free or unattached condition of the whole shell, suspended, as it were,
in the midst of sponges, which they inhabit. The opercular valves, on
the other hand, are less serviceable than usual.
_Range, Habitats, &c._--The species are found all over the world,
excepting in the very cold latitudes.[102] _Acasta lævigata_ ranges
from the Red Sea to the Philippines; _A. spongites_ from the south
of England and Wales to the Cape of Good Hope; and _A. cyathus_ from
Madeira to the West Indies; most of the other species seem to have
rather confined ranges. The East Indian Archipelago seems to be
the metropolis of the genus, for here _A. lævigata_, _fenestrata_,
_purpurata_, and _sporillus_, are all found. Of these four species,
_A. purpurata_ lives imbedded, not in sponges, but in the sponge-like
bark of an Isis; and I think it probable, that _A. sporillus_ may
have somewhat analogous habits. The same species often lives imbedded
in different kinds of sponge; thus, I have seen _A. lævigata_ and
_cyathus_ in apparently three kinds, and _A. spongites_ in, as I
believe, four kinds. The genus existed during the miocene period, in
the Coralline Crag, under a form closely allied to _A. spongites_.
[102] I am greatly indebted to Mr. Bowerbank for his great
kindness, in looking over his immense collection of sponges from
all parts of the world, and sending to me all the specimens of
Acasta he could find imbedded in them.
1. ACASTA SPONGITES. Pl. 9, fig. 1 _a_-1 _d_.
LEPAS SPONGITES. _Poli._ Testacea utriusque Siciliæ, 1795, Tab. 6,
fig. 3-6.
BALANUS SPONGEOSUS. _Montagu._ Test. Brit. Suppl., 1808.
---- SPONGITES. _De Blainville._ Dict. Sc. Nat., Pl. 116, fig. 3.
LEPAS SPONGIOSA. _Wood's_ General Conchology, p. 47.
ACASTA MONTAGUI. _Leach_, in Lamarck, Animaux sans Vertèbres, 1818,
et in Encyclop. Brit., Supplement, 1824, vol. 3, Pl. 57.[103]
---- ---- _J. E. Gray._ Annals of Philosophy, New Series, vol. 10,
Aug. 1825.
---- SPONGITES. _Philippi._ Enum. Mollus. Siciliæ, vol. 2, p. 211,
1844.
BALANUS MONTAGUI. _Brown's_ Illustrations of Conchology (2d edit.
1844), Pl. 53, fig. 24-26.
[103] As the plate to the Supplement to the 'Encyclop. Brit.' is
marked as engraved in 1817, I presume Dr. Leach gave a proof to
Lamarck, thus enabling him to publish this species four years
before the Supplement itself appeared.
_Carino-lateral parietes about one sixth of width of lateral parietes:
inner surface of the parietes generally ribbed feebly: scutum with the
articular ridge abruptly cut off at its lower end: tergum with the spur
rounded-truncated, about one third of width of valve._
_Hab._--South coast of England; South Wales, (Tenby); Portugal;
Naples; Sicily; Cape of Good Hope.
This species and the three following, have caused me much doubt and
trouble. At first I took the view here adopted, namely, that they
were distinct: more careful examination made me run them altogether
under one name. Finally, after still further deliberation, and the
examination of a few additional specimens, I concluded there was the
least chance of error in classing them separately. I may here mention,
that in some sponge from the Cape of Good Hope, this species was
mingled with _Balanus spongicola_.
_General Appearance._--The shape is usually that of a cup, the
orifice being, in most cases, rather large, and deeply notched, owing
to the great obliquity of the radii and alæ. The surface is generally
smooth, but furnished with sharp calcareous projections. The colour
is pinkish, and chiefly in the upper part of the shell; the lower
part is often yellowish from the preserved epidermis. The parietes
in the carino-lateral compartments are always narrow, being only one
sixth or one seventh of the width of the parietes in the lateral
compartments. The radii are not very wide, never equalling in width
the parietes. The basis is moderately deep, and sometimes very deep,
being even occasionally curled like a horn on one side. The specimens
from Lisbon and Naples are a little larger than any British specimen
which I have seen; the former being .3 of an inch in basal diameter.
_Scuta_: these are striated longitudinally in close lines, generally
plainly, but to a variable degree. The whole valve is flat, thin,
and rather elongated, with barely a trace of an adductor ridge:
the articular ridge is short and rather prominent: it terminates
downwards abruptly, and this does not appear to be the case in the
two following species. _Terga_: these are small compared with the
scuta, they are slightly beaked: the spur is truncated but rounded,
more especially on the carinal side; it is rather more than one third
of the width of the whole valve. The articular ridge and crests for
the depressor muscles are feebly developed.
_Structure of the Parietes and Radii._--The state of the inner
surface of the parietes varies much; generally they are slightly
ribbed close to the basis, the ribs sometimes extending up to the
sheath; rarely the surface is quite smooth. The edges of the radii
are slightly crenated. The upper internal surfaces of the radii,
where overlying the alæ, are usually marked by feeble undulating
lines, nearly parallel to the basis. The alæ have very oblique
summits.
_Basis_: this is generally of a regular cup-like form, and about two
thirds as deep as the shell is high; sometimes it is pointed at the
bottom and distorted. The edge is feebly crenated, and rarely quite
smooth. It is often penetrated by small rounded irregular holes; and
I have seen specimens from the Cape of Good Hope with parts like a
sieve.
_Cirri_: in the first pair, the anterior ramus is nearly thrice as
long as the posterior ramus. The second cirrus is short, with one
ramus longer by three or four segments than the other ramus; the
terminal segments are truncated. The third cirrus is about one third
longer than the second cirrus. In the anterior ramus of the fourth
pair, the regular pairs of spines are rather crowded together in the
upper part of each segment, and the intermediate little spines and
dorsal tufts are rather long in comparison with those of the fifth
and sixth pairs of cirri: moreover, amongst the regular pairs, a few
very minute and thick spines, pointing upwards, could be perceived.
So that we here have the very unusual case of the fourth cirrus not
exactly resembling the fifth and sixth pairs; and we shall see, in
the following species, that this same anterior ramus of the fourth
cirrus presents in addition another very extraordinary character. In
the sixth cirrus there are on each segment four pairs of spines.
2. ACASTA SULCATA. Pl. 9, fig. 2 _a_-2 _d_.
ACASTA SULCATA. _Lamarck._ Animaux sans Vertèbres, 1818.
---- ---- _Deshayes_, in Guerin, Magasin de Zoologie, 1831, Tab. 24.
_Carino-lateral parietes about one sixth of width of lateral parietes:
inner surface of the parietes generally ribbed strongly: basis with the
edge strongly crenated: orifice of shell rather small: tergum with the
spur generally truncated and nearly half as wide as valve._
_Var._ (_a_) (fig. 2 _b_): _with the walls externally ribbed_.
_Var._ (_b_) (fig. 2 _a_): _with small membrane-covered clefts
between the edges of the walls, close above the basal cup_.
_Hab._--Sydney, Port Fairy, Moreton Bay in lat. 27° S., New South
Wales; Southern Australia; and, according to Lamarck, Western
Australia, in lat. 25° S. Mus. Brit., Cuming, Bowerbank, &c.
I am almost ashamed to admit this species, so small are its differences
compared with _A. spongites_; yet I think that it probably is a
distinct form.
In general appearance and character this species comes very near to
_A. spongites_. As in the latter, the parietes of the carino-lateral
compartments are narrow.[104] The orifice seems always to be smaller.
Internally, the parietes are generally much more strongly ribbed,
and the edges of the basal cup more plainly crenated. The sheath is
generally coloured of a brighter pink, sometimes tinged with orange.
The average largest specimens (from .3 to .5 of an inch in basal
diameter) are a little larger than the largest European specimens:
I have seen one specimen from Moreton Bay .4 in basal diameter, and
from the basal cup being very deep, actually .75 in height. The
_scutum_ has the articular ridge not so prominent and not so abruptly
cut off at the lower end, as in _A. spongites_: on the other hand,
the adductor ridge seems rather more prominent; but these differences
are trifling. In the _tergum_ the breadth of the spur (fig. 2 _c_, 2
_d_) varies in specimens taken out of the same branch of sponge; some
can hardly be distinguished from the same valves in _A. spongites_,
but generally the spur is broader and squarer.
[104] In Mr. Cuming's collection there is a specimen, from Sydney,
which I fully believe to be the present species, but cannot be
positively sure, as the opercular valves have been lost, which is
very remarkable from the walls of the carino-lateral compartments
being reduced to the thickness of a mere thread, not one twentieth
of the width of the lateral compartments; we here see the structure
of _Acasta sporillus_ prefigured.
This species presents some remarkable varieties: in one specimen,
from Sydney, the parietes were externally ribbed longitudinally
(fig. 2 _b_), the ribs being roughened with minute points, giving to
the shell an elegant appearance; and this without doubt is the _A.
sulcata_ of Lamarck, procured by Peron at Shark's Bay, lat. 25° S.,
on the opposite of the Australian continent: some specimens from Port
Lincoln, in South Australia, were not ribbed, only smoothly striated
in longitudinal lines; although both sets of specimens had almost
smooth scuta, and were thus different from common specimens, yet
there could be no doubt, from their similarity in all other points,
that they did not differ specifically from them, though the latter
had their scuta striated, but not their parietes. Hence we see that
there is no relation between the striæ on the parietes and on the
scuta. The Port Lincoln specimens, and some others, were remarkable
from the radii not extending down to the basal cup, a minute cleft,
covered only by membrane, being thus left along the sutures, low
down between the parietes (fig. 2 _a_); we shall see this singular
structure strongly developed in _A. perforata_. Owing apparently to
these clefts, the edge of the basal cup, exhibited traces of six
knob-like teeth, like those characteristic of _A. glans_.
_Cirri._--The cirri resemble those of _A. spongites_, with the
exception that the segments on the posterior pairs bear only
three main pairs of spines. With respect to the anterior ramus of
the fourth pair of cirri, the following very singular facts were
observed:--in a specimen from New South Wales (_var._ with the tergum
having a narrow spur), on two or three of the lower, but not on the
lowest, segments, the front margin was produced or developed into two
or three minute, thick teeth, slightly curved like hooks downwards:
in other specimens from New South Wales (_var._ with the tergum
having a broad spur, and inhabiting the same branch of sponge with
the last-mentioned variety), there was no trace of these teeth. But
again, in two other specimens with the tergum having a broad spur
(collected by different persons, near Sydney), and in another from
South Australia, this structure was carried to an extreme, for in
these (as represented, Pl. 29, fig. 2) there were beautifully formed
teeth on the fourteen lower segments (the twelve upper being without
them), and likewise on the upper segment of the pedicel. These teeth
are graduated in size on each segment; they are admirably adapted for
securing any prey; and, in fact, they convert each segment into a
mandible-like organ. On the segments, on which these teeth are well
developed, some of the regular pairs of spines are aborted.
_Diagnosis._--Finally, this species, if it be, as I believe,
distinct, differs from _A. spongites_ only in the internal surface of
the parietes being more strongly ribbed and brighter coloured,--in
the edge of the basal cup being more plainly crenated,--in the
articular ridge of the scutum being of a different shape,--in the
spur of the tergum being often broader,--in the segments of the
posterior cirri having only three pairs of main spines,--and, lastly,
in the occasional presence of the hook-like teeth on the anterior
ramus of the fourth cirrus.
3. ACASTA CYATHUS. Pl. 9, fig. 3 _a_-3 _c_.
_Carino-lateral parietes about one fourth of width of lateral parietes:
radii wider than the parietes: basis nearly flat, small: tergum with
the spur truncated, half as wide as valve._
_Hab._--Madeira, Mus. Lowe, and Bowerbank. West Indies, Mus.
Stutchbury. Hab. unknown, Mus. Brit.
I feel more confidence in this case, than in that of _A. sulcata_, that
we here have a distinct species, though at one time I treated it only
as a marked variety of _A. spongites_. I rely chiefly on the great
proportional width of the radii of all the compartments, and on the
width of the carino-lateral compartments, compared with the lateral
compartments, and on the general shape of the shell, which differs
considerably from that of the two previous forms: in consequence of
Acasta being attached to and imbedded in a yielding substance, such
as sponge, I believe external form to be of more value as a specific
character in this genus, than in most sessile cirripedes. I have
examined specimens taken out of the yielding _Spongia officinalis_ and
out of an unusually compact sponge, and they resembled each other in
every respect.
_General Appearance._--Colour pale pink, or that of flesh: basis
remarkably flat and rather small, with the walls above bulging out
a little. The radii are very wide, being wider than the parietes
to which they belong: the orifice is generally rather large. The
parietes of the carino-lateral compartments vary from one third to
one fourth of the width of the parietes of the lateral compartments.
Basal diameter of largest specimen .35 of an inch. Internally, the
parietes are generally more strongly ribbed than in _A. spongites_.
The _Opercular Valves_ are large, owing to the form of the shell. The
_Scuta_ present no particular character, and are not distinguishable
from those of _A. sulcata_; but the adductor ridge is perhaps rather
more developed. The _Terga_ (Pl. 9, fig. 3 _c_) are nearly as large
as the scuta, and this is an unusual circumstance; the spur is more
than half as wide as the valve; it is placed not quite close to the
basi-scutal angle; on the carinal side, the basal margin of the valve
slopes a little towards the spur. I may mention, that in several
specimens from Madeira, the scuta and terga, on one side, had grown
to a monstrous thickness.
_Cirri_: these resemble, in every respect, those of _A. spongites_,
with the remarkable exception that on the anterior ramus of the
fourth cirrus, several segments were furnished with the beautiful
downward curved, mandible-like teeth, as in _A. sulcata_; but
differently from in that species, there were none on the upper
segment of the pedicel. I should have thought this an excellent
specific character, had not these teeth been so extremely variable in
_A. sulcata_.
Finally, I think this species is more nearly related to _A. sulcata_
than to _A. spongites_.
4. ACASTA UNDULATA. Pl. 9, fig. 4.
_Shell, apparently, as in A. spongites, but larger: scutum marked by
longitudinal ridges, often in pairs, with the intermediate furrows
rather wide: spur of tergum nearly half as wide as valve._
_Fossil_ in Coralline Crag (Sutton), Mus. S. Wood.
I owe to Mr. Wood the inspection of a fine suite of separate valves.
Owing to the shell never having been found entire, its general shape is
not known, and, what is of more consequence, the relative proportional
width of the parietes of the carino-lateral compartment is unknown. I
have (but with doubt) given it a distinct specific name, owing to the
peculiar character of the furrows on the scuta, and to the large size
of the whole shell. In its other characters it comes nearest to _A.
spongites_, excepting in the spur of the tergum, which resembles that
of _A. sulcata_.
The compartments appear to have been rather smooth externally. The
radii are not wide, as in _A. cyathus_; and the basis is cup-formed.
Internally, the parietes are feebly ribbed, as in _A. spongites_.
Judging from the dimensions of the separated valves, this species
must have equalled and perhaps exceeded the size of the largest
living species, namely, _A. glans_, from Australia. Hence we may
infer, that the basal diameter probably exceeded .55 of an inch: I
may add, that the largest European specimens of _A. spongites_, from
Naples and Portugal, are only .3 of an inch in basal diameter.
_Scuta._--These seem to resemble the scuta of _A. spongites_ in all
respects, except in the longitudinal ridges standing much further
apart, and, consequently, in the furrows being much wider: each ridge
is generally double. Although there is a good deal of variability
in the character of these ridges in _A. undulata_, and likewise in
_A. spongites_, I have not seen any form intermediate between them.
It must, however, be confessed, that this is an extremely variable
character in many sessile cirripedes. In the _Terga_, the spur is
about half the width of the whole valve, and therefore rather wider
than in _A. spongites_.
5. ACASTA GLANS. Pl. 9, fig. 5 _a_-5 _c_.
ACASTA GLANS. _Lamarck._ Animaux sans Vertèbres, 1818.
_Parietes internally quite smooth, with the lateral margins of each
compartment inwardly prominent: basis with the edge rarely crenated,
but furnished with six inwardly prominent teeth: scutum strongly
striated longitudinally._
_Var._ (_a_) _with the edge of the basal cup finely crenated_.
_Hab._--New South Wales, Southern Australia; Mus. Brit., Stutchbury,
&c.
This fine species seems to be extremely common, imbedded in an open
porose sponge on the eastern and southern shores of Australia. It
is very distinct from the other species, with the exception of the
following _A. lævigata_, which, with some hesitation, I have allowed to
remain specifically separated.
_General Appearance._--Excepting in its larger size, this species
differs in external appearance but little from _A. spongites_; its
colour is pale dirty reddish. The surface is generally studded
with small calcareous points. The parietes of the carino-lateral
compartments are about one fourth of the width of the parietes of the
lateral compartments, and therefore proportionally of the same width
as in _A. cyathus_. The largest specimen which I have seen, was .55
of an inch in basal diameter.
_Scuta._--These are slightly narrower, thicker, and more convex than
in _A. spongites_: they are strongly striated in longitudinal lines.
The articular ridge is very feebly developed. _Terga_: in full-grown
specimens, the spur is half the width of the whole valve, and is
truncated; its basal edge being parallel to the basal margin of the
valve. The articular ridge and crests for the depressor muscles are
very feebly developed.
_Internal structure of the parietes._--The inner surface of the
parietes is quite smooth, without even a trace of ribs or teeth. But
the most important character is that the internal lateral margins on
both sides of each compartment, from the sheath to the basis, project
inwards and form a rim; so that when the shell is viewed from within
(Pl. 9, fig. 5 _b_, representing the lateral and carino-lateral
compartments, and part of the carina), the six sutures are seen to be
strengthened by six double columns.
_Basis._--This is moderately cup-formed. The edge, in order to
meet the basal points of the inwardly projecting lateral margins
of the six compartments, has six knob-like teeth. These are placed
at unequal distances, for two on each side stand near each other,
owing to the narrowness of the carino-lateral compartments. The
degree of their development varies extremely; when most developed,
as in the specimen figured (Pl. 9, fig. 5 _a_), each tooth is bifid
and a little hollowed out, so as to receive the points of the two
inwardly projecting margins which form each suture. Ridges, more or
less prominent, running from each of the six marginal teeth, extend
towards the centre of the cup. These six teeth cannot be seen from
the outside. The edge of the cup is rarely crenated; but I have seen
two instances in which this was the case.
_Cirri._--In the first pair, the rami are not quite so unequal as
in _A. spongites_; the longer ramus being about twice as long as
the shorter. In the third pair, there are some very minute, thick,
upwardly-pointing spines, which I did not notice in _A. spongites_.
In the fourth pair, the spines are a little more crowded, with longer
dorsal tufts, than in the sixth pair; and they are mingled with some
very minute, thick, upwardly pointing spines. In young individuals,
there are only three pairs of main spines on the segments of the
sixth pair, instead of four pairs.
6. ACASTA LÆVIGATA. Pl. 9 fig. 6 _a_, 6 _b_.
ACASTA LÆVIGATA. _J. E. Gray_ (!). Annals of Philosophy, (new
series), vol. 10, Aug. 1825.
_Parietes internally quite smooth, with the lateral margins of each
compartment inwardly prominent: basis with the edge strongly crenated,
and furnished with six inwardly prominent teeth: scutum feebly striated
longitudinally, or smooth._
_Var._ (_a_), _epidermis coloured dull orange_.--Red Sea.
_Hab._--Red Sea, Philippine Archipelago; Mus. Brit., Cuming, &c.
This species, of which I have examined many specimens from the above
two and other unknown localities, agrees in all essential points of
structure with _A. glans_, and consequently I for some time classed
them together; but the characters, though usually of small value, by
which this form differs from _A. glans_ being apparently constant, I
have with some doubt allowed it to remain specifically distinct. These
characters are, firstly, the much smaller size of the whole shell
in _A. lævigata_; secondly, the edge of its basal cup being always
crenated, which seems to be a rare accident in _A. glans_; thirdly,
though of secondary importance, the scutum being here less plainly
striated; and, lastly, the spur of the tergum being of less breadth,
and of a more rounded outline; on the other hand, it must be confessed,
that when small specimens of _A. glans_ are taken, there is hardly any
difference in the spurs of the terga.
_General Appearance and Structure of Shell._--The surface of the
shell is often very smooth, but is sometimes studded with some small
sharp calcareous points. The colour is white, or pale reddish-brown;
but in the specimens from the Red Sea, the tint is more orange,
with the upper part of the shell white. The orifice of the shell is
unusually small. The largest specimen which I have seen was only .25
of an inch in basal diameter, and therefore less than half the size
of _A. glans_. The internal surfaces of the _parietes_ are smooth,
with the two lateral margins inflected, as in _A. glans_. The edge of
the basal cup has six knob-like teeth, like those in _A. glans_, but
smaller; and, in addition, it is finely crenated.
_Scuta_: these differ only in being less plainly striated in
longitudinal lines; indeed, some specimens show hardly a trace of
this structure. _Terga_; these valves, in some varieties (Pl. 9, fig.
6 _b_) can hardly be distinguished from those of equal size from
young individuals of _A. glans_; other varieties have the spur (Pl.
9, fig. 6 _a_) not truncated, but broadly pointed, and therefore of
considerably different shape.
Neither in the mouth, nor cirri could I detect any difference with
_A. glans_.
7. ACASTA FENESTRATA. Pl. 9, fig. 7 _a_-7 _c_.
_Shell reddish, with six large, membrane-covered apertures between the
sutures, above the basal cup: carino-lateral parietes half as broad as
lateral parietes; internally, parietes and edge of basis smooth; tergum
with the articular ridge short and prominent; spur pointed._
_Hab._--Philippine Archipelago, Mus. Cuming.
_General Appearance._--Shell rather elongated or tubular; with the
upper part reddish, and the surface roughened with very minute
points. The basal cup is generally as deep as the shell is high,
ending downwards in a blunt point, often curved to one side. The
summits of the radii, as usual, are oblique. The parietes of the
carino-lateral compartments are about half as wide as the parietes of
the lateral compartments, and are therefore of greater proportional
width than in the foregoing or any other species of the genus.
The large membrane-covered openings, or, as they may be called,
windows, presently to be described, between the lower halves of the
compartments, is much the most remarkable character of this species.
The largest specimen which I have seen was only .23 in diameter, and
.6 of an inch in height, measured from the basal point of the cup to
the tips of the compartments.
_Scuta._--These barely exhibit a trace of longitudinal striæ. The
valve is rather thick and convex. The basi-tergal angle is much
rounded off. Internally, the articular ridge is thick and rather
prominent. _Terga_: the valve is furrowed in the line of the spur:
the spur is pointed and rather long; it is distinctly separated from
the basi-scutal angle of the valve, and the basal margin on the two
sides of the spur forms a straight line. The articular ridge is
prominent, and short.
_Structure of the Parietes, Radii, and Basis._--The parietes are
internally quite smooth down to its basis. The edges of the radii are
also smooth, as is the edge of the basal cup. The alæ project less
than usual. The radii are of moderate breadth, they extend downwards
only a little below the sheath, namely, about half way down the
shell, where they terminate, as usual, in a point. The increase in
width, during growth, of the radii, and their not extending down to
the basis, would necessarily cause a gap between the opposed edges
of the walls, in the portion beneath the radii; but besides this,
the edges of the walls themselves, beneath the radii, and on the
opposed side beneath the alæ, are hollowed out, but on the latter
side or beneath the alæ sometimes in a lesser degree. The result of
this is, that the compartments, in their lower halves, are separated
from each other by membrane-covered windows or apertures, arched at
their upper ends, and of considerable size, namely, about as wide as
the parietes of the carino-lateral compartments. I have only further
to remark, that during the downward growth of the parietes, the
apertures increase in size, but at the same time become closed up at
their upper ends; and the arched layers of shell added at these upper
ends, assume a very different aspect from the rest of the parietal
surface,--appearing like two wedges, with their points upwards, let
in, on one side of the suture, between the ordinary parietal surface
and the radius, and, on the other side of the suture, between the
ordinary parietal surface and the recipient furrow of the radius.
The _animal's body_ was in a bad state of preservation; but, as far
as I could make out, the cirri resembled those of _A. glans_.
_Affinities._--This species differs from _A. glans_ in not having the
internal margins of the compartments projecting inwards. It differs
from all the ordinary varieties of _A. spongites_, in the smoothness
of the basal edges of the parietes and of the edge of the cup; in
the greater width of the carino-lateral compartment, though this is
a variable point in _A. spongites_; slightly in the shape of the
scuta and terga; and, lastly, in the large, membrane-covered openings
between the compartments.
8. ACASTA PURPURATA. Pl. 9, fig. 8 _a_-8 _c_.
_Shell dull blueish-purple, with six small, membrane-covered apertures
between the sutures, close above the basis: tergum with the articular
ridge very short and prominent; spur very broad and rounded._
_Hab._--Sumatra; Philippine Archipelago; imbedded in the bark of an
Isis; Mus. Cuming, Stutchbury, Brit.
This species is perfectly distinct from the others, as shown by its
general appearance, its habits, and the structure of its opercular
valves: it is allied to _A. fenestrata_, in having membrane-covered
apertures between the compartments, and in some respects in its
opercular valves: it is also allied to _A. sulcata_ and _cyathus_ in
the parietes being often internally ribbed, in the basal cup having a
crenated edge, and in the anterior ramus of the fourth cirrus being
furnished with the minute hook-like spines.
_General Appearance._--Sub-globular, slightly compressed, with a
rather small orifice; smooth, but sometimes furnished with sharp
shelly points; dull purple, more or less dark, with the upper parts
of the walls often white. The radii are rather narrow, and generally
white, with their summits only slightly oblique, but variable in
this latter respect. The parietes of the carino-lateral compartments
are narrow, being only one sixth of the width of the parietes of the
lateral compartments. In some specimens there are membrane-covered
apertures of considerable size, in others mere narrow clefts, between
the basal halves of the compartments. The basal cup is moderately
deep. The largest specimen was only .16 of an inch in basal diameter.
_Scuta_, rather broad, externally convex, not longitudinally
striated: articular ridge prominent, short, not extending down above
one third of the length of the valve. Depression for the adductor
muscle deep. On the internal surface, near to the rostral angle, a
rather large purple spot of corium adhered to the valve. _Terga_,
broad, externally rather convex: scutal margin protuberant: carinal
margin slightly inflected, or furnished internally with a rim:
articular ridge prominent, very short, not extending down above one
fourth of the valve. Spur very broad, rounded, confluent with the
basi-scutal angle of the valve.
_Internal Structure of the Parietes, Radii, and Basis._--The
parietes, internally, are either quite smooth, or more commonly
ribbed, with the basal edge in consequence crenated; the ribs
are either placed at an unusual distance from each other, and
consequently are few in number, or are pretty close together. The
edge of the basal cup is either quite smooth, or closely crenated,
or distantly toothed, in conformity with the state of the internal
surface of the parietes. The radii have nearly smooth edges, with
their summits more or less oblique. They sometimes extend down only
three fourths, or only two thirds, of the length of the shell, and
the margins of the parietes under the radii being a little hollowed
out, the sutures are converted into clefts or apertures (of course
covered by membrane) like, but not so large as, those in _A.
fenestrata_. The margins of the parietes are hollowed out only on the
side of the radius, and not on both sides of the sutures, as is most
usual in _A. fenestrata_. In some specimens the radii extended down
close to the basal cup, and only very minute clefts were left between
the opposed edges of the parietes.
In the _animal's body_ the only noticeable character was, that on the
anterior ramus of the fourth pair of cirri, some of the segments were
furnished with very broad and thick, small, downwardly curved, teeth
or hooks, like those described in certain varieties of _A. sulcata_;
but they are here stronger and thicker. The segments in the three
posterior pairs of cirri are not so much elongated, as in the other
species.
9. ACASTA SPORILLUS. Pl. 9, fig. 9 _a_-9 _d_.
_Shell purplish-brown, with the parietes internally strongly ribbed
and reticulated: carino-lateral compartments extremely narrow, not
extending down to the basis._
_Hab._--Sooloo Islands, East Indian Archipelago; Mus. Dana.
I am indebted to Mr. Dana, the distinguished naturalist of the United
States Antarctic Expedition, for two specimens of this interesting
species, which, in the singular reticulated structure of the inner
surface of the walls, and in the almost rudimentary condition of the
carino-lateral compartments, not extending down to the basal cup, is
very distinct from the foregoing species. I have used Mr. Dana's very
appropriate MS. name of _sporillus_. The specimens were dredged up,
lying quite loose and unattached at the bottom of the Sooloo sea; the
one which I opened, must have long lain dead; but Mr. Dana assures me
that some were living, and he has sent me drawings of parts of the
mouth and cirri: I am much surprised at this circumstance; for analogy
would have made me believe that this species must have been imbedded
in some sponge-like body, such as the bark of a zoophyte, and that it
could not have lived unattached. I may add that a small fragment of a
brown leathery substance adhered to the upper end of one of the two
specimens, and this seems to indicate attachment.
_General Appearance._--Shell shaped like a pointed acorn; slightly
flattened; orifice extremely small; surface very finely punctured,
covered by a purplish-brown epidermis, with transverse stripes of
different shades, and with the apex dark; according to Mr. Dana, when
fresh, the colour was purplish-carmine. Radii narrow, white. The
carino-lateral compartments are extremely narrow; the wall-portion
(fig. 9 _b_) forming a mere linear rib, terminating downwards in
a sharp point, which does not reach the basal cup: hence this
compartment evidently tends to become rudimentary. The basal cup is
moderately deep and pointed. Basal diameter .16; height, from the
bottom of the cup to the top of the shell, .24 of an inch.
_Scuta_: narrow, with the upper part produced; not striated
longitudinally; coloured by a pale purple, longitudinal band.
Internally, there is scarcely a trace of an articular ridge, which,
in the other species, is always more or less developed. _Terga_ with
the spur bluntly pointed; nearly the whole basal margin, on the
carinal side, slopes towards the spur.
_Internal Structure of the Parietes, Radii, and Basal Cup._--The
parietes are strongly ribbed internally; and these ribs are connected
by very narrow, less prominent, transverse, slightly branched ridges,
giving a reticulated structure to the inner surface. Between several
of the main longitudinal ribs, in the lower part of the shell, new
ribs may be seen in process of formation, and these tend to convert
the reticulated structure into a double row of minute cells. I have
not met with an exactly similar structure in any other cirripede; but
I have no doubt that the little transverse ridges are homologous with
the transverse calcareous septa in the parietal pores of many Balani,
in the same manner as the internal longitudinal ribs, in this and
other species of Acasta, are homologous with the longitudinal septa
forming the above pores. The edge of the basal cup is pectinated with
teeth, which lock into the teeth formed by the ends of the internal
parietal ribs. The radii are narrow, and have smooth edges. The alæ
project beyond the parietes to a remarkably small extent. The sheath
is free, or hollow beneath. I have already described the almost
rudimentary condition of the carino-lateral compartments; this is
best exhibited in an internal view of the two compartments, as given
in Pl. 9, fig. 9 _b_.
_Animal's body_ unknown to me: from Mr. Dana's drawing the three
posterior pairs of cirri seem to have been much elongated: and the
rami of the first pair, as usual, unequal in length.
M. Deshayes has given an indifferent figure and imperfect description
of ACASTA TUBULOSA (Guerin, Magasin de Zoologie, 1831, Tab. 39; and
Guerin, Iconographie du Règne Animal, Mollusques, Tab. 38, fig. 4, but
here by a misprint called _A. spinulosa_); it is utterly impossible to
recognise the species of this genus from such materials.
3. _Genus_--TETRACLITA. Pl. 10, 11.
TETRACLITA. _Schumacher._ Essai d'un Nouveau Syst., &c., 1817.
CONIA. _Leach._ Journal de Physique, tom. 85, 1817.[105]
ASEMUS. _Ranzani._ Memoire di Storia Naturale, 1820.[106]
POLYTREMA. _De Ferussac._ Dict. Classique d'Histoire Naturelle, 1822.
LEPAS. _Gmelin._ Systema Naturæ, 1789.
BALANUS. _Bruguière._ Encyclop. Method., 1789.
---- _Lamarck._ Animaux sans Vertèbres, 1818.
[105] From a note by the Editor, it appears that Schumacher's essay
appeared before the number of the Journal containing Leach's paper,
so that Schumacher's name must be adopted.
[106] I have not seen a _complete_ copy of this work, and give the
title from a catalogue; the running heading of the part containing
the Cirripedia, is "Opuscoli Scientifici."
_Compartments four, sometimes externally calcified together: parietes
permeated by pores, generally forming several rows. Basis flat,
irregular, calcareous, or membranous._
_Hab._--Throughout the tropical and warmer temperate seas.
_General Appearance._--The shell is conical, more or less depressed,
and very rarely, even when growing in crowded groups, becomes
cylindrical or elongated. The orifice is seldom large, generally
diamond-shaped or oval. The colour is either nearly white or purple,
occasionally even inky black, or very dark green, and sometimes of a
pale pink peach-blossom. The surface is sometimes smooth, but more
commonly longitudinally ribbed; the outer lamina of shell is very often
wholly corroded away, excepting close to the basis, leaving the solidly
upfilled parietal tubes exposed: these give the shell a striated
appearance, or they appear like flattened tapering points adpressed to
its surface (Pl. 10, fig. 1 _b_): Lamarck attempted to express this
appearance, by using the specific name of _stalactiferus_. The colour
of the shell depends, to a considerable extent, on the colour of the
shelly matter in these exposed parietal tubes. We shall presently see
that the corrosion and disintegration, to which some of the species
are so liable, plays an important part during their growth. The radii
are either well developed, as in most of the species; or they are
entirely absent, as in the great majority of specimens of _T. porosa_
and _serrata_. In many individuals of _T. porosa_ and _purpurascens_
not only are the radii absent, but the four compartments are calcified
together without any trace, on the external surface, of the sutures.
The largest specimen which I have seen of _T. porosa_, which is the
largest species, was two inches in basal diameter, and nearly one inch
and a half in height.
_Scuta._--These valves are sub-triangular, and either longitudinally or
transversely elongated. Externally, the growth-ridges are moderately
prominent, and in _T. costata_ and _cœrulescens_ they are crossed by
longitudinal striæ. Along the occludent margin, the inflected extremity
of each alternate growth-ridge is generally much thickened,--a set of
teeth being thus formed, by which the two valves are locked together.
In _T. porosa_, this character is variable, for sometimes every
alternate ridge, and sometimes only two or three ridges, separated
from each other by several growth-ridges, are thus developed into
teeth. The _articular_ ridge is either moderately prominent, or is
extremely prominent, as in _T. cœrulescens_; but the lower edge in no
case depends as a free, hinge-like style, as sometimes in Balanus.
The _adductor_ ridge is generally well developed and distinct from
the articular ridge: in _T. purpurascens_ it is very blunt: in _T.
serrata_ it is united to the articular ridge half way up it, thus
forming a deep tubular cavity running up to the apex of the valve: in
_T. cœrulescens_, the adductor ridge is very short, and is united to,
or almost continuous with, the lower end of the articular ridge, a
small sub-cylindrical tubular cavity being thus formed. Small crests
exist for the attachment of the rostral and lateral depressor muscles,
in most of the species, excepting _T. purpurascens_ and _costata_, in
which, however, more especially in the former, there are, instead
of crests, minute pits for the attachment. These crests vary much in
prominence in the same species.
_Terga._--These valves present no essential differences from those of
Balanus; they are sometimes beaked, and the beak is hollow and occupied
by a thread of corium, as in that genus. The external surface of the
valve is often depressed in the line of the spur, but there is never a
longitudinal furrow with the edges folded in, as in Balanus. The spur
is very short in _T. purpurascens_. In _T. radiata_, the articular
ridge is remarkably prominent. The crests for the depressor muscles are
well developed in all the species. The shape of the terga is variable
in nearly all the species, and greatly so in _T. porosa_.
_Compartments._--Owing to there being only four compartments, the
lateral pair are large; the size of the carina relatively to the
rostrum varies, according as its alæ have been added to during
diametric growth. The walls are very thick, and are composed of
numerous tubes, in some species as many as fourteen or fifteen rows
being exposed on the basal margin (Pl. 10, fig. 1 _g_). The tubes
are generally angular, and slightly elongated in the ray of the
circular shell; sometimes they are nearly circular and small. New
tubes are formed only at the basal edge of the outer lamina, by the
bifurcation of the septa which form the tubes. In very young specimens
there is only a single row of tubes; and in _T. rosea_ this holds
good throughout life: in this species (fig. 3 _d_) the tubes, in the
single row, are large and quadrangular, and the outer lamina of shell
is strengthened by numerous, small, internal, longitudinal plates.
I believe the branching septa, which separate and form the parietal
tubes, correspond with the longitudinal septa in the more simple
walls of Balanus. The tubes become solidly filled up, in their upper
parts, with hard, and generally coloured shelly matter. The degree
to which they are filled up differs in the different species; the
external side of each tube is always first thus coated. The thin outer
lamina of shell, in several of the species, commonly disintegrates and
disappears; the upfilled parietal tubes being thus exposed. The inner
lamina of the walls is generally smooth, but in _T. radiata_ it is
longitudinally ribbed, as in most species of Balanus. The sheath is
generally dark-coloured; its lower edge does not project or overhang
the inner lamina, as is usual in Balanus, excepting in _T. serrata_,
and in some few varieties of _T. porosa_.
The _Radii_, when developed, are either narrow or broad, with their
summits either oblique or extending in a straight line from the top
of one compartment to that of another. In _T. serrata_, I have not
seen a single specimen with the radii developed; in _T. porosa_, they
are very seldom developed, and then, apparently, only in quite young
specimens, in which they are narrow; in _T. purpurascens_, they seem to
be about as often developed as not, and when present they are broad;
in _T. costata_, _cœrulescens_, and _radiata_, they are always largely
developed. In some specimens of the species, in which the radii are
not developed, even the sutures do not reach the external surface;
the outer lamina of the parietes being continuous all round, so that
the shell seems formed of a single piece. Even in such specimens, the
four compartments, viewed from within, can be seen to be distinct; and
the sutures can generally be traced across the whole thickness of the
parietal tubes; in this latter case, when the sutures are broken open,
the radii are seen to be represented (Pl. 10, 1 _h_) by a few small
sinuous ridges. Owing to the disintegration of the upper and outer part
of the shell, and the consequent exposure of parts of the sheath and
alæ, the radii sometimes appear as if developed, when such is not the
case. With respect to the internal structure of the radii, they are
formed, in _T. purpurascens_ and _costata_, of tubes, like those of the
parietes, and therefore according to the normal plan; whilst in the
other species they are formed by longitudinal sinuous ridges, sending
out on each side irregular denticuli; and the interspaces between
the ridges are filled up solidly during the growth of the radii, in
all the species, except in _T. radiata_, in which they are left to a
considerable extent open. These sinuous ridges, with their denticuli,
homologically represent the branching septa which form the parietal
tubes. The edges of the alæ are crenated in all the species, except in
_T. costata_.
_Diametric growth._--When first examining groups of _T. porosa_, in
none of which the radii had been developed, and in which, consequently,
the shell could not have grown in diameter, but only at its basal
margin, I was at first unable to comprehend, how the upper part of the
shell and the orifice could have acquired their proper proportional
width. The young shell, at its first formation, starts with an orifice
so small that a pin could hardly be inserted in it; and this in many
individuals is never increased in diameter by the diametric growth
of the shell; but in place of this, as the conical shell is added to
at its base, the whole upper part disintegrates and wears away, the
orifice becoming thus enlarged. We thus see that the corrosion and
wearing away of the upper part of the shell is a necessary element in
its growth. The development of the radii, which in some of the species,
as in _T. purpurascens_, at first seems to be quite capricious, really
depends upon the fact, whether the specimens have been exposed to
disintegration; for I have almost always found that when the outer
lamina of shell has been well preserved, the radii have been developed,
and the orifice has been enlarged by their growth, instead of by the
wearing down of the upper part of the conical shell.
_Basis._--This consists of a very thin, flat, though irregular,
translucent, calcareous plate, which towards the edges is sometimes
membranous. In _T. purpurascens_, the basis is entirely membranous.
When a portion of the calcareous base is dissolved in acid, a tissue
is left, composed of several laminæ, to which numerous bifurcating
cement-ducts are attached: even before dissolution, these delicate
bifurcating ducts can just be perceived by the aid of a simple lens.
_Mouth._--The several organs present no particular characters. There
are generally three teeth on each side of the notch in the labrum.
The palpi usually have parallel sides, but are club-shaped in _T.
purpurascens_ and _costata_. The mandibles have generally four teeth,
but there are five in _T. vitiata_, and only three in _T. costata_. The
maxillæ are notched. The outer maxillæ are bilobed in front.
_Cirri._--The segments of the three posterior pairs usually support
only three pairs of main spines, but there are four pairs in _T.
vitiata_ and _costata_: between each pair, there is either a tuft of
fine spines, or a single fine spine. The rami of the first cirrus
are unequal in length. In the third cirrus, the posterior ramus is
sometimes much elongated, but sometimes both rami are short and blunt.
Some of the segments in the third cirrus often support very coarsely
and doubly pectinated spines. Under the head of _T. porosa_, it will be
seen to what a remarkable degree the relative numbers of the segments
in the several cirri vary, even in specimens taken out of the same
cluster.
The _Branchiæ_ are well developed (at least in _T. porosa_ and
_costata_), as a large, plicated, tapering fold, with a small second
fold on the inner side at the base. In _T. porosa_ the _stomach_ is
destitute of cæca. The _vesiculæ seminales_ in this same species are
large, with their broad, blunt ends reflexed. The _ovarian_ tubes
surround the sack, and cover the basal plate.
_Distribution and Habitats._--This genus is confined to the tropics,
and to the warmer parts of the temperate seas: in the southern
hemisphere, it ranges south, to the Cape of Good Hope and to Van
Diemen's Land: in the northern hemisphere it does not appear to range
so far; I do not know of any authentic case of a species having been
found in the Mediterranean, or on the shores of the United States,
north of the West Indies. _Tetraclita porosa_ is found round the whole
world; _T. radiata_, also, has a very wide range, inhabiting the West
Indies, the East Indian Archipelago, and New South Wales. This latter
species, as well as _T. cœrulescens_, is often attached to the bottoms
of ships, adhering to _Balanus tintinnabulum_. The several species live
attached to tidal rocks, to littoral shells, or to massive corals. I
have met with two instances, in the West Indies and the Philippine
Archipelago, of _T. porosa_ adhering to wood. _Tetraclita porosa_
seems to feed chiefly on crustaceans: the number and the size of the
amphipods, isopods, and entomostracans, together with an annelid,
in the stomachs of some specimens from South America, was quite
surprising. As many as five species occur in the same region, in the
eastern half of the world; thus on the shores of New South Wales, we
have _T. porosa_, _vitiata_, _radiata_, _purpurascens_, and _rosea_; in
the Philippine Archipelago, we have _T. porosa_, _vitiata_, _costata_,
and _cœrulescens_.
I have not seen any species of this genus fossil.
_Variation._--The species vary in shape nearly, but not quite as much
as in Balanus, for we very rarely here see cylindrical varieties; but
in the same species, we have extremely depressed, steeply conical, and
convex forms; the orifice varies in relative size; the state of the
surface, whether ribbed or smooth, whether well preserved or corroded,
the upfilled parietal tubes being thus exposed, varies more than in
Balanus. The colour also varies; specimens of _T. porosa_ (Pl. 10, fig.
1 _a_ to 1 _f_) being dark purple, or even inky black, white, pale
pinkish-purple, and green: as far as I have seen, the pinkish varieties
of _T. porosa_ are confined to the eastern half of the world. Even the
ridges and crests on the under side of the scutum vary in some degree;
and the tergum in _T. porosa_ varies considerably, and in some of the
other species, slightly, in general shape: I believe that the tergum
becomes narrow and elongated, when the shell is steeply conical, with
the orifice of small diameter. After having spent several days in
disarticulating and closely examining the many specimens of _T. porosa_
in my possession, I concluded that its varieties formed at least four
species, these being in external appearance extremely distinct; but
ultimately the many intermediate forms compelled me to unite all into
a single species. Again, I may instance the conical, ribbed variety
of _T. purpurascens_ (Pl. 11, fig. 1 _b_), with the outer lamina of
the shell preserved and with the radii widely developed, as having not
the smallest resemblance to the other common depressed variety (fig. 1
_a_), having a granulated surface, produced by the exposed tips of the
upfilled parietal tubes, and without a trace of the radii or even of
the sutures; I may add that, according to the characters used by some
authors, these two varieties would be classed in two separate genera.
To distinguish the species of this genus, the chief reliance must be
placed (as in the case of most other sessile cirripedes) on the general
outline of the opercular valves, and on the ridges and crests on their
under sides: the internal structure, however, of the radii, and in
a lesser degree of the parietes, must not be overlooked. I have not
found the parts of the mouth, or the differences in the cirri, of much
service; and it will be shown under _T. porosa_, that the relative
numbers of the segments in the several cirri, and even their shape, is
extraordinarily variable.
_Affinities._--All the species are pretty closely allied, and there is
no ground for making any sectional division of the eight species, more
especially not on the ground whether or no the radii are developed.
_Tetraclita purpurascens_, taking into account all the characters,
including the animal's body, is perhaps the most distinct species in
the genus, but it is closely allied to _T. costata_. In the well-marked
character of the parietes being formed of a single row of large tubes,
_T. rosea_ differs from all the other species. The genus is closely
allied to Balanus; I can point out no difference in the animal's
body, nor any constant difference in the opercular valves. The ridges
and crests on the under sides of the scuta are more prominent in
Tetraclita; and all the species, excepting two, have crests (though
sometimes very slight) for the attachment of the rostral depressor
muscle, and these occur only in two species of Balanus: crests, also,
for the attachment of the lateral depressor muscle are common in
Tetraclita, but rare in Balanus. The main character, however, of the
genus, as compared with Balanus, is derived from the existence of
only four compartments, and in a lesser degree from the several rows
of parietal pores; but, as just stated, _T. rosea_ has only a single
row, and, on the other hand, in _Balanus crenatus_, there is a slight
tendency exhibited, in the dividing of the longitudinal septa near
the outer lamina, to form a second row of parietal pores; and in _B.
cariosus_, moreover, we actually have two or three rows of irregular
and very short pores. The thin, yet solid calcareous basis which
occurs in all the species of Tetraclita, excepting _T. purpurascens_,
resembles the basis in _Balanus flosculus_ and _imperator_, but I
suspect that the structure of the cement-ducts is different in the two
genera.
1. TETRACLITA POROSA. Pl. 10, fig. 1 _a_-1 _m_.
LEPAS POROSA. _Gmelin._ Syst. Naturæ, 1789.[107]
BALANUS SQUAMOSUS. _Bruguière._ Encyclop. Method., 1789, Pl. 165,
fig. 9, 10.
LEPAS FUNGITES. _Spengler._ Skrivter af Naturhist. Selskabet, 1 B.,
1790.
---- POROSA. _Wood's_ General Conchology, Pl. 9, fig. 4, 1815.
TETRACLITA SQUAMULOSA. _Schumacher._ Essai d'un Nouveau Syst., &c.,
1817.
BALANUS STALACTIFERUS. _Lamarck._ Animaux sans Vertèbres, 1818.
---- ---- _Chenu._ Illust. Conch., Pl. 4, fig. 6, 7.
ASEMUS POROSUS. _Ranzani._ Memoire di Storia Naturale, Tab. 3, fig.
32-35.
CONIA POROSA. _Sowerby._ Genera of Recent and Fossil Shells, Plate,
1823.
---- ---- _Leach._ (sine descript.) Encyclop. Brit. Supplement, vol.
3, 1824, Tab. 57.
[107] As Gmelin and Bruguière published the same year, I do not
know which comes first, but I have adopted the best known name.
Most authors give, amongst their references, Chemnitz, vol. 8,
Tab. 98, fig. 836, 837, but the accompanying description is more
applicable to _T. serrata_ of this work than to _T. porosa_;
without a figure or description, however, of the under side of the
scutum it is impossible to decide. Several authors, also, give
_Lepas cariosa_ of Pallas, as a synonym; this, however, is the
_Balanus cariosus_ of this work.
_Radii rarely present, when present narrow; even the sutures often
absent: shell steeply conical, with the surface generally corroded, and
having a stalactiferous appearance._
_Var._ (1) communis (Pl. 10, fig. 1 _a_): _outer lamina of shell
almost wholly removed; the portion preserved, as well as the exposed
parietal tubes, gray, or pale dirty brown, or dull purple_.
_Var._ (2) nigrescens (Pl. 10, fig. 1 _c_): _outer lamina of shell
almost wholly removed; the portion preserved, and the exposed
parietal tubes, very dark purple or inky black_.
_Var._ (3) viridis: _outer lamina of shell almost wholly removed;
the portion preserved, and the exposed parietal tubes, green; under
surface of opercular valves clouded green_.
_Var._ (4) rubescens (Pl. 10, fig. 1 _b_): _outer lamina of shell
almost wholly removed; the portion preserved, and the exposed
parietal tubes, pale reddish-purple, or peach-blossom purple; under
surfaces of opercular valves often reddish purple; terga often rather
narrow, with the spur somewhat pointed, and with the basal margin on
the carinal side sloping, or not making an angle with the spur_.
_Var._ (5) elegans (Pl. 10, fig. 1 _d_): _outer lamina preserved,
excepting sometimes near the summit of the shell; white, tinged
with yellowish brown from the epidermis; surface strongly ribbed
longitudinally; orifice rather small; sheath reddish-purple; terga
narrow, with the basal margin sloping as in var._ rubescens.
_Var._ (6) communis (young) (Pl. 10, fig. 1 _e_): _radii developed,
very narrow; outer lamina of shell preserved, gray or dull purple;
surface slightly ribbed longitudinally_.
_Var._ (7) patellaris (Pl. 10, fig. 1 _f_): _radii developed,
very narrow, white; outer lamina of shell preserved, generally
reddish-purple; steeply conical, with the orifice extremely small;
surface smooth, with longitudinal white ribs. Terga very narrow, with
the spur sharply pointed, and with the basal margin on the carinal
side sloping towards it, or not making an angle with it. Scuta, with
the adductor ridge very prominent. Attached to a ship's bottom_.
_Hab._--West Indies, Brazil, West Colombia, Panama, Galapagos
Archipelago, California, Philippine Archipelago, China, East coast of
Australia, Red Sea; generally attached to tidal rocks, sometimes to
shells, sometimes to wooden posts. Very common.
_General Appearance._--This, the widest-distributed and much
the commonest species of the genus, varies greatly in external
appearance. The usual shape is steeply conical, but some individuals
are much depressed. In the common varieties the outer lamina of
shell has been removed even close to the basal edge; the upfilled
parietal tubes being thus exposed (fig. 1 _b_), as flattened
adpressed points. These points are largest in large specimens, but
they vary somewhat in size in specimens of equal growth. When the
outer surface is preserved, it is generally ribbed longitudinally,
but is sometimes quite smooth. The most general colour is dirty gray
or dark purple; but many specimens are pale pinkish-purple, owing
to the exposure of the parietal tubes upfilled with shelly matter
of this tint: there are also, as given under the characters of the
_vars._, black, white and green varieties. The sheath is always
tinted by the prevailing colour. The radii are rarely developed,
but generally the four sutures are distinct; sometimes these are
externally quite obliterated, the shell, as seen from the outside,
consisting of a single piece like a patella or fissurella. When the
radii are developed they are very narrow, with their upper edges
oblique: their development seems always coincident with the more or
less perfect preservation of the shell, and their function is to
enlarge the orifice; the enlargement being usually effected by the
disintegration and removal of the whole upper part of the conical
shell. The size of the orifice varies considerably; in the seventh
variety it was extraordinarily small: in outline it varies from oval
to rounded trigonal or rhomboidal; in some specimens, with the radii
well developed, it was rounded pentagonal.
_Size._--_Tetraclita porosa_ is the largest species of the genus;
I have seen specimens attached to a large pebble of granite in the
British Museum, which measured two inches in basal diameter, and
nearly one inch and a half in height.
_Scuta_: these are sub-triangular and generally a little elongated,
but they vary slightly in relative breadth (fig. 1 _i_, 1 _l_), and
likewise in the degree to which the basi-tergal angle is rounded off.
The under surface is clouded with dull or pinkish-purple, or with
green, or is nearly white. The articular ridge is not prominent, and
the articular furrow is narrow. The adductor ridge is prominent, and
runs upwards for some distance close and parallel to the articular
ridge; and sometimes it extends nearly or quite up to the apex of
the valve; in one single specimen the adductor ridge had an abraded
appearance, and was very little developed. The crests for the rostral
and lateral depressores are sharp and distinct. Along the occludent
margin, the ends generally of the alternate lines of growth are
enlarged into knobs, serving to lock the two valves together; but in
many specimens only two or three knobs, at intervals of several lines
of growth (fig. 1 _b_), are developed.
_Terga_: when the upper end of the valve is not corroded, there is
a distinct beak, hollow within for a thread of corium. The scutal
margin is not much inflected, and the articular ridge not very
prominent. The spur is placed quite close to the basi-scutal angle
of the valve, so that there is no basal margin on that side of the
valve. The width of the valve and of the spur, and the acumination of
the extremity of the latter, varies in a remarkable manner. In the
broad and commonest variety (fig. 1 _k_), the basal margin of the
valve form an angle of about 130° with the carinal side of the spur,
and the basal end of the spur is broad and truncated. In the less
common and narrow variety (1 _m_), the basal margin in some extreme
cases forms very nearly a straight line with the carinal side of
the spur; and the spur itself is bluntly pointed: in _var._ 7 it is
sharply pointed.
_Structure of the parietes and radii._--In all cases the four sutures
are quite distinct, from top to bottom, on the internal lamina of
the shell, and generally they run through the whole thickness of
the walls, and are visible externally. Often they do not quite
reach the outer lamina, and then the shell externally consists of a
single piece, like a patella. Sometimes the sutures can be traced
running through the parietal tubes only for a short distance from
the internal surface; where they cease, the two walls of the suture
become fused together. When a perfect suture is split open, the
radius is represented (fig. 1 _h_) by a few narrow, sinuous ridges,
sending out on each side little branches or denticuli; these are
received into corresponding furrows in the opposed compartment. These
ridges run nearly parallel to each other, and somewhat obliquely,
from the outer lamina of the shell to the basis. When the radii are
developed, their edges are similarly formed, by sinuous denticulated
ridges, with the interspaces between them filled up solidly. The alæ
are but little prominent.
The _mouth_ does not deviate from the generic type. The _cirri_
are remarkable from the variability in the several pairs of the
relative numbers of their segments, as shown in the following table.
The segments do not correspond even on opposite sides of the same
individual, as may be seen in the two lower lines of the table. I
believe that variability to this degree is very uncommon in other
cirripedes, though, as stated in the Introduction, the number of the
segments always increases with the growth of the individual. The
terminal segments in the longer rami of both the first and third
cirrus are antenniformed,--being elongated, and of a different
shape, with fewer bristles, compared with the basal segments of
the same cirri. It is apparently these terminal segments which
are particularly liable to vary in number. In both rami of the
third cirrus, some of the segments, from the sixth to the eleventh
inclusive, (counting from the bottom), more especially the eighth,
ninth, and tenth, carry a few spines coarsely and doubly pectinated;
but as some of the adjoining segments carry spines which may be
called doubly serrated, it is not easy to draw an exact line of
demarcation. Sometimes, though rarely, a few of the nearly terminal
segments in the second cirrus are furnished with similar, doubly
pectinated spines.
_Numbers of the segments in the rami of the Cirri, in different
specimens._
+------------------+--------------+--------------+--------------+-------+
| |First cirrus. |Second cirrus.|Third cirrus. |Sixth |
| | | | |cirrus.|
| |--------------+--------------+--------------+-------|
| |Shorter|Longer|Shorter|Longer|Shorter|Longer|Either |
| |ramus. |ramus.|ramus. |ramus.|ramus. |ramus.|ramus. |
|------------------+-------+------+-------+------+-------+------+-------|
|Specimen from | | | | | | | |
|Pernambuco, Brazil| 9 | 19 | 10 | 11 | 15 | 27 | 26 |
|(_var._ 1) | | | | | | | |
|------------------+-------+------+-------+------+-------+------+-------|
|Another specimen | | | | | | | |
|from the same | 12 | 23 | 12 | 14 | 14 | 19 | 23 |
|cluster | | | | | | | |
|(_var._ 1) | | | | | | | |
|------------------+-------+------+-------+------+-------+------+-------|
|From the Galapagos| | | | | | | |
|Archipelago, | 10 | 16 | 10 | 11? | 11 | 16 | 25 |
|(_var._ 1) | | | | | | | |
|------------------+-------+------+-------+------+-------+------+-------|
|From the | | | | | | | |
|Philippine | 10 | ? | 10 | 10 | 11 | 19 | 18 |
|Archipelago, | | | | | | | |
|(_var._ 4) | | | | | | | |
|------------------+-------+------+-------+------+-------+------+-------|
|From California | 15 | 24 | - | - | - | - | 22 |
|(_var._ 5) | | | | | | | |
|------------------+-------+------+-------+------+-------+------+-------|
|From the bottom | | | | | | | |
|of a ship | - | - | 9 | 9 | 10 | 17 | 14 |
|(_var._ 7), | | | | | | | |
|young | | | | | | | |
|------------------+-------+------+-------+------+-------+------+-------|
|From South America| | | | | | | |
|(_var._ 1) | 11 | 18 | 11 | 12 | 13 | 21 | 22 |
|British Museum | | | | | | | |
|------------------+-------+------+-------+------+-------+------+-------|
|Another specimen | | | | | | | |
|from the | 16 | 25 | 13 | 18 | 15 | 23 | 27 |
|same cluster with | | | | | | | |
|the last specimen | | | | | | | |
|------------------+-------+------+-------+------+-------+------+-------|
|Same individual, | | | | | | | |
|but the cirri | 15 | 28 | 15 | 21 | 16 | 23 | - |
|from the opposite | | | | | | | |
|side of the body | | | | | | | |
+------------------+-------+------+-------+------+-------+------+-------+
_Varieties._--Under the generic description, I have stated that after
having spent some time in examining a very large suite of specimens
of _T. porosa_, I concluded that at least four of the varieties were
true species. It so happened that all the specimens which I first
examined of the _var._ (4) _rubescens_, had the narrow sloping terga,
and scuta with only two or three great teeth on their occludent
margins; but ultimately, in a group thus characterised, I found one
or two individuals with terga precisely of the shape of those in
_var._ (1) _communis_. Again, in a group of dull purple specimens of
_var. communis_, a few had the narrow sloping terga, and scuta with
teeth on their occludent margins, intermediate in size and number
between the varieties with only one or two great teeth, and those
with every alternate growth-ridge enlarged into a tooth. Hence _var.
rubescens_ completely broke down as a species.
With respect to _var._ (5) _elegans_ (a M.S. specific name of Leach)
I inferred at first, from external appearance alone, that it was
distinct; the outer lamina of the shell and even the epidermis is
preserved; the surface is strongly ribbed, and the whole shell,
excepting the sheath, is nearly white; the terga are narrow, with a
sloping basal margin as in _var._ (4) _rubescens_. Whole groups of
specimens are thus characterised. But as _var. communis_ is often
white, and as the surface, when the outer lamina is preserved, is
generally, as we shall presently see, ribbed longitudinally, the
differences in _var. elegans_ are quite unimportant.
The (6th) _var._ differs from _var. communis_ only in the narrow
radii having been developed, and consequently in the orifice being
pentagonal, and in the outer, longitudinally ribbed lamina of the
shell having been preserved. In the same group of specimens, I have
seen every intermediate stage between this and the common form. It
must not, however, be supposed that the young of _var. communis_
always pass through these stages, for such is not the case. In the
genus _Balanus_ it has been seen, how capricious in some species is
the development of the radii.
Of the other varieties, enumerated at the beginning, no further
mention is required, excepting with respect to _var._ (7), the most
singular of all. I have seen only three specimens, sent to me by Dr.
Aug. Gould, of Boston, United States, and these from the appearance
of their bases I have no doubt had been attached to a ship,--the
only instance which I have met with, in the present species. The
shell is steeply conical, with the orifice so small as to be reduced
to a mere pore; the radii are extremely narrow and white; the shell
is thin, with the surface smooth, but ribbed longitudinally and
regularly; the outer lamina of the shell and the epidermis are
perfectly preserved; the upper part of the shell is reddish purple,
which dies away towards the base: careful examination of the apex
shows that at the first growth the young shell was blueish-green.
The terga are narrow, with a sloping basal margin, as in _var.
rubescens_, but with the point of the spur sharper. I have formerly
remarked that the shape of the terga seems influenced by the size
of the orifice. The lower edge of the sheath depends freely: I have
seen no other instance of this latter structure, so common, but so
variable, in _Balanus_, in the present species, except to a partial
extent in one distorted specimen, in Mr. Stutchbury's collection,
adhering to _Balanus tintinnabulum_, and probably taken from a ship's
bottom. I may add that this distorted specimen was remarkable from
its radii being wider than in any other instance,--from its smooth
uncoloured surface without longitudinal ribs,--and from the perfect
preservation of the epidermis over its entire surface. Although Dr.
Gould's specimens, in external aspect, are absolutely and entirely
different from the common varieties of _T. porosa_, there are so many
intermediate forms, and the differences are so little important that
I feel no hesitation in attributing them to variation, consequent on
the individuals having been exposed to unusual conditions, attached
to the bottom of a ship.
2. TETRACLITA SERRATA. Pl. 10, fig. 2 _a_-2 _d_.
_Shell dark greenish-gray, with narrow, longitudinal, serrated ribs:
radii absent: scutum with the adductor and articular ridges forming a
cavity, which runs up to the apex of the valve._
_Hab._--Cape of Good Hope; Algoa Bay; attached to sandstone and to
Patellæ; Mus. Brit., Cuming, and Stutchbury.[108]
[108] I have seen three separate lots of this species all from the
Cape of Good Hope; one lot was collected by Dr. Krauss, at Algoa
Bay, and I strongly suspect is the species described by him in his
'Südafrikanischen Mollusken' as _Conia porosa_. If the species,
figured by Chemnitz, and mentioned in a note (p. 329), under _T.
porosa_, be the present species, the specimens probably did not
come from Tranquebar, on which point Chemnitz speaks only from
memory. I have seen one specimen ticketed New South Wales, it is
possible, considering the case of _T. rosea_, that this may be
correct, but I should like to have further confirmation before
giving it as a habitat.
_General Appearance._--Colour dark greenish gray; form steeply
conical; surface covered, especially in the lower half of the shell,
by numerous, narrow, sharp, longitudinal ridges, but slightly
prominent, and serrated or transversely divided into small teeth:
when the shell has been much disintegrated, the upper part of the
surface consists of the exposed, smooth, rather large, upfilled
parietal tubes. I have seen no instance of the development of
the radii; sometimes even the sutures are with great difficulty
distinguishable, though I believe they always reach the outer
surface; sometimes the sutures are wide from the disintegration of
the edges of the compartments. Orifice rounded or oval.
_Scuta._--The scuta and carinal half of the terga are blueish-green.
In the scuta neither the articular ridge or furrow are much
developed: the adductor ridge is prominent, and is united to the
articular ridge, about half way up the latter, thus forming a rather
large, triangular cavity, which runs up to the apex of the valve.
The _Terga_ are beaked. The spur, measured across the upper part,
is half as wide as the valve; it is bluntly pointed; it is placed
quite close to the basi-scutal angle of the valve, so that there
is no basal margin on that side; it curves towards the scutum, its
extremity extending beyond the basi-scutal angle.
_Structure of the Shell and Radii._--The parietal tubes are rather
large, especially those adjoining the inner lamina of the walls. The
shell is of singularly little specific gravity, which is due to the
parietal tubes not being filled up with shelly matter nearly to so
great an extent as in the other species; even in the uppermost part
the tubes are not solidly filled up, only their external sides are
thickly coated with greenish-black shell, which by corrosion becomes
grayish. The radii, as stated, are not developed: the shell breaks
with singular facility along the sutures, and the radii are then seen
to be most feebly represented by a few very small branching ridges.
The alæ have their edges plainly crenated. The sheath is dark green,
with the lower edge free.
The _Mouth_ presents no particular characters. With regard to the
_Cirri_, I am doubtful whether any confidence can be placed in the
numbers of the segments being constant; but I may state that the
second cirrus contained thirteen and sixteen segments in its two
rami; the third cirrus only fourteen in both rami; and the sixth
cirrus twenty-six segments in both rami. Whereas in every specimen
of _T. porosa_, the longer ramus of the third cirrus contained more
segments than either ramus of the second. About half the segments,
namely, those in the middle of both rami of the third cirrus, are
furnished with coarsely and doubly pectinated spines, like those in
_T. porosa_.
_Affinities._--Upon the whole, this species is more nearly allied
to _T. porosa_ than to any other. In the cavity formed by the
union of the adductor and articular ridges, it is allied to _T.
cœrulescens_. This species differs from all, in its little specific
gravity, consequent on the parietal tubes being only slightly filled
up, and in the peculiarly serrated, narrow, approximate ridges on
the external surface of the walls. The character derived from the
adductor ridge, just alluded to, is remarkable. In the shape of the
terga, in the absence of radii, and in the structure of the body,
this species approaches closely to _T. porosa_.
3. TETRACLITA ROSEA. Pl. 10, fig. 3 _a_-3 _d_.
CONIA ROSEA.[109] _Krauss_ (!). Die Südafrikanischen Mollusken, Tab.
6, fig. 28, 1848.
BALANUS CUMINGII. _Chenu._ Illust. Conch., Tab. 4, fig. 5.
[109] I am greatly indebted to Professor Krauss for having sent
me, for examination, the unique specimen collected by himself in
Algoa Bay. There can be no doubt of the identity of the African and
Australian specimens. It is a singular circumstance that the same
species should occur in these two distant places, and, as far as at
present known, not in the intermediate, more tropical coasts.
_Shell dirty white, tinged with pink; parietes formed by a single row
of large tubes: radii generally narrow: tergum with the spur rather
short and broad._
_Hab._--New South Wales, Moreton Bay in lat. 27°, Port Jackson, and
Twofold Bay; South Africa, Algoa Bay. Attached, in Australia, to
littoral rocks and shells; often associated with _T. purpurascens_,
_Chthamalus antennatus_, and _Catophragmus polymerus_; Mus. Brit.,
Cuming, Krauss, Darwin, Stutchbury.
_General Appearance._--Shell steeply conical, often rather convex;
dirty or brownish white, feebly tinted with pink; external surface
generally much disintegrated, and having in the upper part a pillared
appearance, owing to the exposure of the upfilled, large, square,
parietal tubes, and, in the lower part, a striated (and sometimes
serrated) appearance, from the exposure of the parallel, approximate
plates, with which the outer lamina of the shell is internally
strengthened. In only a few young specimens, the whole outer lamina
of the shell was well preserved; and in these the surface was very
smooth, and even glossy, giving to the specimens a quite different
aspect; even in partially corroded specimens, the lower part of the
shell sometimes is quite smooth. Generally, the radii are developed;
in most specimens they are narrow, but sometimes of moderate width;
their summits are oblique, and their edges often notched or toothed.
The recipient furrow, in each opposed compartment, is often almost
as wide as the radius itself, and is equally notched. In some much
corroded specimens there were no radii. Basal diameter of largest
specimen, 1.1 of an inch.
_Scuta_, generally thick, sometimes very thick, with the external
surface usually much corroded: articular furrow rather wide;
articular ridge not very prominent; adductor ridge prominent. The
rostral depressor muscle is attached in a small oblong pit, sometimes
including little crests; and the lateral depressor muscle is attached
to what may be described either as three or four parallel furrows or
crests.
_Terga_, with the spur placed close to the basi-scutal angle, so that
there is no basal margin on that side; spur short, with its lower end
truncated and rounded; broad, even exceeding, when measured across
the upper part, half the width of the valve. Articular furrow wide.
Apex not beaked.
_Structure of Walls and Radii._--This species differs from all the
others of the genus in having only a single row (fig. 3 _d_) of
parietal tubes; these are large, quadrangular, but elongated in the
ray of the circle. They are not filled up, even at the very top of
the shell, but they become thickly lined all round with compact
shelly matter. When the surface of the shell is disintegrated,
these upfilled tubes greatly affect, as already stated, the
external appearance. The outer lamina near the basis is internally
strengthened by longitudinal, sharp, approximate ridges or plates,
which, also, often affect, after corrosion, the external appearance.
The _radii_ have their sutural edges formed by a set of narrow,
branching ridges or septa; the ends of which, seen externally, often
give a notched outline to this edge; the recipient furrows in the
opposed compartments are deep, and their edges likewise are often
notched: the interspaces between the branching ridges are filled up
solidly. The _alæ_ have their edges coarsely crenated. The lower edge
of the sheath is not free.
The _mouth_ and _cirri_ present no particular characters: the third
cirrus has both its rami elongated, with the terminal segment
tapering. In the three posterior pairs of cirri, the tufts of little
spines between the main pairs are rather large.
_Affinities._--This species has no particular affinity with any
other. The circumstance of there being only a single row of parietal
tubes is not so important a difference as might at first be thought,
inasmuch as in the other species, during their quite early youth, the
walls are formed of only a single row of tubes or pores.
4. TETRACLITA PURPURASCENS. Pl. 11, fig. 1 _a_-1 _d_.
LEPAS PURPURASCENS.[110] _Wood's_ General Conchology, p. 55, Pl. 9,
fig. 42, 1815.
BALANUS PLICATUS. _Lamarck._ Animaux sans Vertèbres, 1818.
---- ---- ET PUNCTURATUS. _Chenu._ Illust. Conch., Tab. 4, fig. 3 et
12.
CONIA DEPRESSA (!). _J. E. Gray._ Appendix, Dieffenbach's Travels in
New Zealand, 1843 (sine descript. vel figurâ).
[110] The descriptions given by Wood and Lamarck are fuller and
more accurate than is usual in the case of Cirripedes, and I have
no doubt regarding these two names. The _Conia depressa_ of Dr.
J. E. Gray is, as I know from having seen the original specimens,
the young of this same species; the name is unaccompanied by any
description or figure.
_Shell depressed, pale purple or dirty white, with the surface
longitudinally ribbed, or corroded and granulated: radii or even
sutures none, or radii well developed and broad, with their summits
parallel to the basis: basis membranous: scutum transversely elongated:
tergum small, with the spur extremely short and rounded._
_Hab._--Sydney, New South Wales; Flinder's Lagoon, Sir C. Hardy's
Island, Barrier Reef; King George's Sound, Western Australia; Van
Diemen's Land; New Zealand, adhering to _Pollicipes spinosus_;
Mus. Brit., Cuming, Stutchbury, Darwin, &c. China (?) attached to
_Pollicipes mitella_, Mus. Brit. and Stutchbury. Generally attached
to tidal rocks, sometimes to shells. Very common.
_General Appearance._--Shell generally much depressed, in a few
cases rather steeply conical, in one single instance cylindrical,
but not much elongated. Colour, when alive, pale, but fine purple; I
presume, judging from some dried specimens, sometimes dirty white.
The state of the surface varies remarkably: about half the specimens
(fig. 1 _a_) which I have seen, had the outer lamina of shell quite
removed, and the surface granulated, owing to the projecting and
exposed tips of the upfilled parietal tubes; the radii are not
developed, and often even there is no trace of the four sutures;
the rather large orifice is somewhat rounded, and the two scuta,
with their surfaces disintegrated, have their middle parts deeply
indenting the terga. The shell, in the other and perhaps more common
condition (fig. 1 _b_), has the outer lamina preserved, and is
longitudinally ribbed with generally at least five or six ribs on
each compartment: the radii are here very wide, and extend from tip
to tip of the compartments, so that their summits are parallel to the
basis; they are generally covered by a brownish epidermis, thickly
clothed with little spines; the orifice is neatly diamond-shaped;
the apices of the opercular valves meet at a common point: these
specimens are almost always smaller and younger than the granulated
specimens. Altogether the specimens in the two opposite states have,
in their external appearance, nothing in common, and no one, without
careful examination, would ever suspect that they were specifically
identical; this, however, was proved by the intermediate forms,
and, in one instance, three of the compartments had their surfaces
granulated, and were entirely destitute of the radii, whilst the
fourth by some chance had been preserved from corrosion, was
longitudinally ribbed, and had its epidermis-covered radius fully
developed. The difference in the appearance of the opercular valves,
in the two states, is simply owing to the degradation of their upper
parts in the granulated specimens.
The basal diameter of the largest specimen was one inch, but the
height only .35 of an inch.
_Scuta_, transversely elongated, so that the basal margin is nearly
twice as long as the tergal margin: articular ridge very little
prominent; articular furrow wide but shallow; adductor ridge very
blunt, slightly prominent, sometimes almost absent, almost parallel
to the basal margin: there are no distinct crests for the rostral
or lateral depressor muscles, but some small irregular pits for the
latter. In one young specimen, the lines of growth were crenated,
showing a tendency in the valve to become longitudinally striated, as
in the allied _T. costata_. In some young and immature specimens, the
basal margin was deeply sinuous.
_Terga_, small in area, not above half that of the scuta: spur
extremely short, broad, placed close to the basi-scutal angle of the
valve, so that there is no basal margin on that side of the spur. The
lower end and sides of the spur form one uniform curve. Articular
ridge barely developed. Crests for the tergal depressores sharp and
prominent.
_Structure of the Shell and Radii._--The walls are very thick, and
the parietal tubes small and numerous; there are sometimes from
twelve to fifteen rows of tubes in the thickness of the wall. The
tubes in their whole upper part are filled up solidly; and, as
we have seen, are often exposed by disintegration. In very young
specimens, of that size in which in _T. porosa_ there would be only
a single row of parietal tubes, there were here two or three rows.
The development of the radii, as we have seen, is very capricious;
the sutures even sometimes being lost. The radii, when developed,
are broad, square on the summit, and covered by brownish hirsute
epidermis: internally they are formed of tubes like those forming the
parietes; in this respect differing from all the species except the
following, _T. costata_. The tubes in the radii run obliquely down
towards the basis; instead of in a transverse line, directly towards
the opposite compartment, as might have been expected from the
structure of the radii in Balanus. The alæ have their edges finely
crenated. The sheath in all the specimens which I have observed is
colourless; its lower edge is not free. The corium entering the
parietal tubes, and lining the opercular valves, the mouth, and the
anterior cirri, is generally of an extremely dark purple colour.
The _Basis_ is entirely membranous, in which respect this species
differs from all the others in the genus.
_Mouth_: all the trophi are unusually hairy or spinose. The labrum is
deeply notched and apparently destitute of teeth on the crest. The
palpi are club-shaped or enlarged at their extremities. The mandibles
have the fourth tooth rudimentary.
In the _Cirri_, the second and third pairs are remarkably short
and blunt. In one specimen the two rami of the first cirrus had
respectively six and sixteen segments; those of the second, six and
seven; those of the third, seven and seven; and those of the sixth
cirrus twenty rather elongated segments, with a small tuft of spines
between each main pair of spines.
_Affinities._--This species differs from all, in its membranous
basis, and in its transversely elongated scuta. It resembles _T.
costata_, and differs from all the other species, in its radii
(when developed) being square on the summit, and in being formed of
tubes,--in the smallness and number of the parietal tubes,--in the
absence of crests on the under side of the scuta for the rostral
and lateral depressor muscles,--in the shortness and rounded form
of the spur to the terga,--and, lastly, in the club-shaped palpi
and small size or absence of the fourth tooth in the mandibles. _T.
purpurascens_ differs from _T. costata_ in those points, namely, in
its membranous basis and transversely elongated scuta, in which it
differs from all the other species, and, moreover, in its scuta not
being plainly striated longitudinally, in having more ribs on the
external surface of the parietes of its shell, and in having only
three pairs of main spines on the three posterior cirri.
5. TETRACLITA COSTATA. Pl. 11, fig. 2 _a_-2 _c_.
_Shell depressed, whitish, generally with ten very prominent
longitudinal ribs: radii broad, with their summits parallel to the
basis: basis calcareous: scutum externally striated longitudinally:
tergum with the spur short and rounded._
_Hab._--Philippine Archipelago, Mus. Cuming. Attached to various
shells, within the tidal limit.
_General Appearance._--Shell whitish, probably tinged, when alive,
with reddish-purple; depressed; surface perfectly preserved, smooth,
but having longitudinal very prominent ribs, almost invariably ten
in number; namely, three on both the rostrum and carina, and two on
the two lateral compartments, with ten corresponding projections
round the basal margin. Orifice passing from rounded-trigonal to
diamond-shaped. The radii are very broad and square at the summit,
and extend from tip to tip of the compartments. Basal diameter of
largest specimen under half an inch, generally from .3 to .4 of an
inch.
_Scuta_, of the usual sub-triangular shape, and not transversely
elongated, as in _T. purpurascens_. External surface striated
longitudinally; in many specimens there is a medial depression, or
a row of very small pits, such as occur on the scuta of _Balanus
trigonus_ and _lævis_. The adductor ridge is moderately developed,
and runs nearly parallel to the occludent margin; there are no crests
for the rostral and lateral depressor muscles.
_Terga_: these in area equal two thirds of the scuta: the spur is
short and rounded, and placed as described under _T. purpurascens_;
but the articular ridge seems to be more prominent than in that
species.
_Structure of the Shell and Radii._--The parietal tubes are small,
and very numerous, as in _T. purpurascens_. The radii are wide,
square on the summits, but not so conspicuously covered by hirsute
epidermis as in that species. Internally, the tubes forming the radii
are smaller, and run more transversely than in _T. purpurascens_,
that is in the normal course, as in Balanus. The edges of the alæ are
nearly or quite smooth. The _Basis_ is as distinctly calcareous, as
in the other species of the genus.
_Mouth_: the trophi are not so hairy as in _T. purpurascens_; the
labrum seems destitute of teeth; the palpi are club-shaped at their
ends; the mandibles have only three teeth. The second and third
cirri are not so short and blunt relatively to the others as in _T.
purpurascens_. In the posterior cirri, the elongated segments carry
four main pairs of spines, between which there is no intermediate
tuft of fine spines.
The _Affinities_ of this species have been fully pointed out under
the last and closely related species. In external appearance, _T.
costata_ can at first hardly be distinguished from those young and
pale-coloured varieties of _T. purpurascens_, which have their
external surface not corroded, and their radii well developed.
6. TETRACLITA VITIATA. Pl. 11, fig. 3 _a_-3 _e_.
_Shell white, generally tinged in the upper part with pink; surface
irregular: parietal tubes very irregular: radii moderately wide, with
their summits oblique: alæ with very thick crenated sutural edges:
tergum with the spur not joined to the basi-scutal angle; spur with its
extremity equably rounded._
_Hab._--Philippine Archipelago; Barrier Reef, (Raine's Islet),
Australia; Mus. Cuming and Stutchbury. Attached to massive corals, to
coral-rock, to a Tridacna, and to _Tetraclita cœrulescens_.
_General Appearance._--Shell conical, moderately steep: white,
generally with a tinge of pinkish-purple in the upper part, owing to
the exposure of the tips of the upfilled parietal tubes. The lower
part of the surface is generally well preserved, and is formed by
very irregular, branching, longitudinal, slightly prominent, ridges.
Radii of moderate width, with their summits oblique. Orifice rather
large, rounded trigonal.
_Scuta_, rather narrow, with the upper part acuminated: the external
surface generally much disintegrated, and marked with some irregular
blotches of dark red. Articular ridge not prominent; articular furrow
rather deep: adductor ridge, distinct from the articular ridge,
pretty well developed, as are the crests for the rostral and lateral
depressor muscles. In young and well preserved scuta, there is an
external, medial, hyaline band, corresponding with the hollow under
the adductor ridge, and caused by the thinness of the valve along
this line.
_Terga_: externally, the carinal half is longitudinally and very
feebly striated. Internally, the articular furrow is very wide, but
shallow, and of unusual length, owing to the preservation of the
upper part of the valve; the articular ridge is not prominent. The
spur does not actually join, as in all the foregoing species, the
basi-scutal angle, but is separated from it by a short piece of basal
margin; its two sides are more nearly parallel than is usual, and
the end is regularly rounded. It is always rather narrow, though the
width varies considerably (fig. 3 _d_, 3 _e_). It extends in the same
straight line with the middle of the articular furrow. The terga,
though not possessing any striking characters, differ considerably in
appearance from those of the other species.
_Structure of the Shell and Radii._--The parietal tubes are
remarkable from their irregular shapes, and unequal sizes (fig. 3
_b_),--hardly two resembling each other. Sometimes a single elongated
tube will reach across the whole thickness of the walls. The septa
between the tubes are rather thick and rugged. The tubes are
generally darkly coloured from the adhering corium; they are solidly
upfilled, but only in the uppermost part, by dark chocolate-red
shelly matter. The radii are formed by irregularly branching ridges,
with the interspaces filled up solidly. The square edges of the alæ
are much thicker than in any other species, and are furnished with
transverse ridges, which are sometimes slightly branched. The inner
lamina of the walls near the basis, in most of the specimens, is
irregularly and longitudinally ribbed, the ribs being longitudinally
striated. The sheath and the upper part of the inner lamina of the
parietes are clouded with chocolate-red.
The animal's _Mouth_ and _Cirri_ were ill-preserved; but I was able
to make out that the labrum had some strong teeth, and that the
mandibles were furnished with five teeth, a greater number than in
any other species. The palpi had parallel sides as usual. In the
sixth cirrus, the segments had four pairs of main spines, instead of
the usual number of three.
_Affinities._--This species does not appear to be particularly
related to any other one: perhaps it is rather nearer to the two
following than to the foregoing species. The irregular parietal
tubes, thick-edged alæ, form of terga, five teeth to the mandibles,
and four pairs of spines to the segments of the posterior cirri, are
its chief characteristics.
7. TETRACLITA CŒRULESCENS. Pl. 11, fig. 4 _a_-4 _d_.
LEPAS CŒRULESCENS. _Spengler._ Skrivter af Selskabet, 1 Bind.,
1790.[111]
[111] The longitudinally folded walls, as described by Spengler,
the blue colour, the habitat, namely, associated with _B.
tintinnabulum_ from the East Indies, and more especially the
expression "Valvulæ operculi cardine dentato mobilis," apparently
referring to the highly prominent articular ridge of the scutum,
leave little doubt on my mind that I have rightly named the present
species.
_Shell with the upper part tinged greenish-blue, longitudinally ribbed:
radii moderately wide, with their summits oblique: scutum with a small
adductor and extremely prominent articular ridge, united together and
so forming a small sub-cylindrical cavity: tergum with the spur not
joined to the basi-scutal angle._
_Hab._--Philippine Archipelago, attached to _Balanus tintinnabulum_;
attached to a ship's bottom, and to _Balanus tintinnabulum_, both
from the Pacific Ocean; attached to a massive coral, and associated
with _T. vitiata_, and therefore from the tropical eastern seas; Mus.
Brit., Cuming, Stutchbury.
_General Appearance._--Shell conical, sometimes depressed; surface
with rather broad, smooth, longitudinal ridges; whitish, with the
upper part greenish-blue, sometimes very feebly tinted with pink;
radii white, or mottled with blueish-green, or with pink. When the
outer lamina of shell has been corroded, the upfilled parietal tubes,
of a dull blueish-gray colour, are exposed. The radii are moderately
wide, with their summits very oblique. In basal diameter, one
specimen was 1.8, and another 1.5 of an inch.
_Scuta_, externally furrowed very slightly in a longitudinal
direction, causing the lines of growth to become a little sinuous.
The valve is strong and thick; and the epidermis, when preserved,
is hirsute with spines. The articular ridge is extraordinarily
prominent; it projects, as measured from the external surface of
the valve, to an amount equalling half the width of the valve in
its widest part. The adductor ridge is very short, and is united
to the bottom of the articular ridge, thus forming a small, nearly
cylindrical tube, which runs up to near the apex of the valve. The
inflected occludent margin is broad and coarsely toothed. The crests
for both depressor muscles are not very prominent. When the scuta
and terga are articulated together, owing to the great projection of
the articular ridge of the scutum, its upper part is separated from
the tergum (fig. 4 _b_), by a remarkably wide and deep, fissure-like
hollow. The scutum, in some distorted specimens from a ship's bottom,
was narrow in proportion to its ordinary breadth.
_Terga._--These are large. Externally, there is a broad, longitudinal
depression, bounded on each side by a ridge. The carinal half is
feebly striated longitudinally. A large upper portion (fig. 4 _b_)
projects freely, and stands, when the two valves are articulated
together, above the apex of the scutum, but does not form a beak as
in _T. porosa_. Internally, the tergal margin is widely inflected;
the articular furrow is very deep, but the articular ridge is not
prominent. The spur is not very broad; it is separated from the
basi-scutal angle by a small space of basal margin, which forms a
straight line with the basal margin on the opposite side of the spur.
The end of the spur is truncated, and is parallel to the basal margin.
_Structure of the Shell and Radii._--The walls are thick: the
parietal tubes are rather large and regular; they become solidly
filled up in their upper parts. The sutural edges of the radii are
formed by unusually narrow, sinuous ridges, sending off delicate
denticuli on each side: the interspaces between these ridges are
solidly filled up. The crenated edges of the alæ are rather thick.
Neither the _mouth_ nor the _cirri_ offer any noticeable character;
but I may observe, that, in the mandibles, the third and fourth teeth
are close together; and that, in the three posterior cirri, the tufts
of small spines between the main pairs are small.
_Affinities._--This species is very distinct from the others, with
the exception of the following _T. radiata_, to which it is in
several respects allied. The under side of the scutum, with its great
articular ridge, and the cylindrical tube formed by the union of the
latter with the short adductor ridge, affords the most noticeable
character.
8. TETRACLITA RADIATA. Pl. 11, fig. 5 _a_-5 _d_.
CONIA RADIATA.[112] _De Blainville._ Dict. Sc. Nat., 1816-1830, Pl.
164, fig. 5, 5 _a_, (sine descript.)
---- LYONSII. _G. B. Sowerby._ Genera of Recent and Fossil Shells,
Plate, 1823, (sine descript.)
[112] The synonymy of this species is complicated. De Blainville
gives no description under the article Conia, published in 1818, or
in the vol. published in 1822; but I believe, from the figures of
the opercular valves, that I have correctly identified this species
with his _C. radiata_. Mr. Sowerby gives no description of _C.
Lyonsii_, or any figure of the opercular valves, but his drawing
of the shell is much better than de Blainville's, and I believe
it is the same species. Whether de Blainville's or Sowerby's
plate appeared first I cannot ascertain. In the second edition of
Lamarck, _Conia radiata_ of Blainville is given as a synonym to
_Balanus radiatus_ of that work; but this is quite erroneous. I
may add that if de Blainville's name does not apply to the present
species, it must to _T. cœrulescens_, and as the latter is the
older name it will be permanent. In this case, _T. radiata_ might
be allowed to stand as my own name, considering that Mr. Sowerby's
figure is imperfect and is not accompanied by any description.
At first I thought that the present species might be the _Lepas
mitra_ of Spengler, ('Skrivter af Naturhist. Selskabet,' 1790,
Tab. 6, fig. 5), but the parietes are not described as porose;
and the folds on the walls are too broad; on the other hand, his
description of the opercular valves makes me think this may be the
same species.
_Shell white, with numerous approximate longitudinal ribs: radii broad,
with their summits slightly oblique, internally porose: tergum with the
articular ridge extraordinarily prominent, with the spur not joined to
the basi-scutal angle._
_Hab._--West Indies, adhering to _Balanus eburneus_ and to _Lepas
anserifera_. New South Wales, adhering to _Tetraclita porosa_.
Attached to _Balanus tintinnabulum_, on a ship's bottom from Sumatra;
not rarely attached to _Balanus tintinnabulum_ on ships' bottoms;
Mus. Brit., Stutchbury, and Cuming.
_General Appearance._--Shell white, rather steeply conical, with
numerous, approximate, rather narrow, longitudinal, rounded ribs on
the walls: in a specimen half an inch in diameter, there were from
eight to twelve ribs on each compartment. The outer lamina of shell
seems always well preserved. Orifice rounded, trigonal. Radii white,
smooth, broad, with their summits only slightly oblique. I have
seen one specimen 1.2 in basal diameter, but quarter of an inch is
a common size, and very young specimens are unusually frequent in
collections.
_Scuta_ broad, externally not striated longitudinally. The articular
ridge is prominent, and the furrow deep, but not in so great a
degree as in _T. cœrulescens_. The adductor ridge is only slightly
prominent; it extends upwards only a little way above the lower end
of the articular ridge, and does not form with the latter a cavity.
There are no crests for the rostral depressor muscle, but there is a
little pit, formed by the folding over of the occludent margin.
_Terga._--These valves, when articulated with the scuta (fig. 5 _b_),
project above them to an extraordinary degree, and are separated
from them by a deep, fissure-like hollow, caused by the remarkable
prominence of the articular ridge of the terga. The upper part of
the tergum is not beaked, and does not project freely much above the
sack. The valve is large; externally there is a rounded longitudinal
furrow. The tergal margin is broadly inflected. The articular furrow
is deep, and the articular ridge far more prominent than in any other
sessile cirripede, for it projects, as measured from the outside
surface, more than half the width of the valve; and consequently
the valve, when viewed vertically from above, almost appears as if
formed by the union of three plates, viz., the articular ridge, and
the outside surface on each side of the spur. The spur is of moderate
width, with the corners rounded: it is placed near, but not close to
the basi-scutal angle, so that there is on this side a small portion
of basal margin, forming nearly a straight line with the margin on
the carinal side. In some young specimens, about the tenth of an
inch in diameter, from the West Indies and from New South Wales, the
spur (the position of which I found varied a little in some other
specimens) was placed nearly in the middle of the valve, and very
nearly at right angles to the basal margin; it is possible that these
may be a distinct species, but without larger specimens to judge
from, I think it more probable that this difference in the tergum is
due to variation and youth.
_Structure of Shell and Radii._--The parietal tubes are commonly
elongated in the ray of the circle: the septa are rather thick, and
strongly crenated at their basal edges. The inner lamina of the
walls is strongly ribbed longitudinally. The broad radii have their
sutural edges formed by ridges, with numerous and closely approximate
denticuli: the interspaces between the main ridges are not soon
filled up, and at the bottom, each interspace usually terminates in
a pore or tube; so that the radii are not solid, as in most of the
foregoing species, but porose. The alæ have their edges crenated.
_Basis_, calcareous, of unusual thickness; the inner, or upper
surface, is striated from the centre in rays, corresponding with the
ribs on the inner lamina of the walls. This striated or furrowed
structure in the basis, shows a tendency to its becoming tubular or
porose, as may be inferred from analogous cases in Balanus.
_Animal's body_ unknown.
_Affinities._--This species is rather more closely allied to the last
than to any other. There is a close analogy in the peculiar manner
in which the scuta and terga are articulated together in the two
species: in this species it is effected by the great development of
the articular ridge of the tergum, and in _T. cœrulescens_ by that
of the scutum. The internally striated calcareous basis, and the
internal tubular interspaces between the denticulated ridges of the
radii, are peculiar characters. The white colour, the narrowly and
closely ribbed parietes, and the broad radii, give this species an
aspect, by which it can be easily recognised.
4. _Genus_--ELMINIUS.
ELMINIUS. _Leach._ Zoological Journal, vol. 2, July, 1825.
_Compartments four: parietes not porose. Basis membranous._
_Distribution_, Southern temperate seas.
_General Appearance._--Shell conical, with a strong tendency in most
of the species to become cylindrical: orifice generally large. Walls
either thin and smooth, or thick and plicated longitudinally. Colours
various, pale purple, greenish, white, and, in _E. plicatus_, owing to
the exposure of an intermediate lamina of shell, bright orange-yellow.
Radii, either of considerable width, with their summits oblique and
rounded, as in the first two species of the genus, or very narrow, as
in the last two species. _Elminius plicatus_ is the largest species,
and is sometimes one inch in basal diameter. The outer surface of this
latter species is occasionally much corroded.
_Scuta_: these are of the usual shape; in _E. Kingii_ and _modestus_
there is no adductor ridge and no crests for the depressor muscles;
in _E. plicatus_ and _simplex_, on the other hand, there is a well
developed adductor ridge and crests for the lateral depressor muscles;
in some individuals, also, of _E. plicatus_ there are small crests for
the rostral depressores.
The _Terga_ are remarkable for their variability in all the species;
in many specimens of _E. Kingii_ and _modestus_ the basal margin on
the carinal side of the spur is deeply hollowed out. The width and
acumination of the spur varies in all the species. In _E. plicatus_ and
_simplex_ this valve is remarkably like that of _Tetraclita porosa_. In
some specimens of _E. Kingii_ the terga and scuta are firmly calcified
together.
_Structure of the Parietes and Radii._--As in Tetraclita, the two
lateral compartments are necessarily broad. The parietes are never
porose, but consist, in appearance, of a single layer of shell. In _E.
modestus_ the basal internal edges of the parietes are smooth, but in
the other species they are striated longitudinally with short ridges,
or sometimes with sub-cylindrical projections. In those specimens of
_E. plicatus_, which have externally suffered much corrosion, the walls
have been rendered extremely thick, by the inward production of these
ridges or plates; and in this case the ridges are not confined to the
basal edges, but extend upwards close to the sheath. The basal surfaces
of the walls in these latter specimens resemble those of Chelonobia,
but the walls in that genus have an internal lamina, which here is not
the case. The radii are wide in _E. Kingii_, and of moderate width in
_E. modestus_, with their summits oblique and smoothly rounded, and
their sutural edges not in the least crenated. In _E. simplex_ they
are extremely narrow, smooth-edged, and rounded: in _E. plicatus_ they
are narrow, and in this species alone the sutural edge is sinuous, and
sends inwards short ridges or teeth. The alæ, in all the species except
this last, are likewise smooth-edged. The lower edge of the sheath
depends, more or less freely, in all the species, except in _E. Kingii_.
_Basis_, membranous in all four species. In _E. modestus_, the true
basal membrane is extremely thin, and is divided into concentric slips:
on its inner surface there are attached numerous cement-ducts, varying
from 1/3000 to 1/2000 of an inch in diameter, repeatedly trifurcating,
rarely forming hexagonal or quadrangular loops, and with the branches
placed approximately parallel to each other. Beneath the true basal
membrane there is a complicated layer of cement, in the form of a
network, or of separate tubes, or in beads and patches. In _E. Kingii_,
the basal membrane presented a wonderfully complicated appearance, in
part due to the cement forming a mass of inosculating fibres; many of
these fibres seemed to end in circular discs of cement.
Neither the _Mouth_ or _Cirri_ offer any noticeable generic characters,
as distinct from Balanus and several other genera. The _Branchiæ_,
in _E. plicatus_, are well developed and moderately plicated. In _E.
modestus_ they are small, not plicated, but with a rounded sinuous
margin: in a specimen having a basal diameter of 25/100, the branchiæ
in total length were only 4/100 of an inch. At the bottom of the sack
I observed some inwardly pointed, tapering filaments, such as occur in
Balanus. In this same species I measured the ova, which were unusually
elongated, being 19/2000 in length; I may add, that the probosciformed
penis was actually thrice the length of the animal's body in some small
but mature specimens (with ova), having a shell with a basal diameter
of 16/100 of an inch.
_Distribution and Habitats._--This genus is remarkable, inasmuch as it
is not distributed over the whole globe: three of the species occur
very commonly on the shores of New South Wales, Van Diemen's Land,
and New Zealand; not extending, as far as I can judge, much north of
Sydney: the fourth species is confined to South America, ranging
from the Falkland Islands and Tierra del Fuego, as far north as
Chiloe. Elminius, therefore, appears to be strictly a southern genus.
_Elminius Kingii_ and _modestus_ represent each other on the American
and Australian continents; so I believe _E. plicatus_, in New Zealand,
represents _E. simplex_ in New South Wales and Van Diemen's Land.
The species are all attached to tidal rocks and shells. _E. Kingii_
is sometimes attached to floating wood. At the Falkland Islands, the
last-mentioned species adhered to some rocks, in a running brook of
fresh water, at most eighteen inches under high-water mark, so that for
the greater part of each tide it was exposed to absolutely fresh water.
At Sydney I found _E. modestus_ adhering to oysters in a muddy lagoon,
almost separated from the sea, and apparently very unfavorable for
cirripedes.
_Affinities._--This genus can be distinguished from Tetraclita only
by the four compartments not being porose, and by the basis being
always membranous; whereas, in _Tetraclita purpurascens_ alone it is
membranous. _Elminius Kingii_ and _modestus_, on the one hand, are
closely allied together, as are _E. simplex_ and _plicatus_ on the
other hand. The last-named species, in the characters of its opercular
valves and in its shell, comes nearest to Tetraclita. In _T. rosea_ we
have seen that there is only a single row of parietal tubes, and the
outer lamina of the shell is strengthened (as, indeed, it is in most of
the other species of the genus, and in Balanus) by small longitudinal
plates or ridges, which are similar and homologous to those on the
internal basal edges of the parietes in three of the species of
Elminius; so that the difference in the structure of the parietes, in
Tetraclita and Elminius, is small.
1. ELMINIUS KINGII. Pl. 11, fig. 6 _a_-6 _e_.
ELMINIUS KINGII. _J. E. Gray._ Zoological Miscellany, p. 13, 1831.
---- LEACHII. _King_ and _Broderip_. Zoological Journal, vol. 5,
1832-1834, p. 334, and appendix to King and Fitzroy's Voyages.
---- ---- _G. B. Sowerby._ Genera of Recent and Fossil Shells,
Plate.
_Shell smooth, gray or dirty white: radii broad, smooth-edged: scutum
without an adductor ridge; tergum with the spur distinct from the
basi-scutal angle: scutum and tergum sometimes calcified together._
_Hab._--Tierra del Fuego, Falkland Islands, Chiloe. Attached to tidal
rocks and sometimes to floating timber; Mus. Brit., Darwin, &c.
_General Appearance._--Shell fragile, either steeply conical with a
large orifice, or sub-cylindrical; surface smooth, grayish or white,
with large portions covered by pale brown epidermis. Radii broad with
their summits oblique, smooth, slightly arched, exhibiting a large
surface of the alæ. The alæ usually have their summits much less
oblique than those of the radii; the portion added during diametric
growth is of a dead white colour. The growth ridges on the scuta
are very little prominent, and are crossed by a very obscure band
of blueish-gray. The largest specimen which I have seen was .8 in
basal diameter, and the longest cylindrical variety .55 of an inch in
height.
The _Scuta_ are remarkable for not having any adductor ridge or
crests for the depressor muscles; the articular ridge is prominent,
but it is short, not extending down half the valve. I have mentioned
under the genus, that in many specimens at the Falkland Islands the
scuta and terga were calcified together.
The _Terga_ are rather small: the basal margin on the carinal side
of the spur is always hollowed out, but to a very variable degree,
as may be seen in the three figures (6 _c_-6 _e_); this margin is
generally dentated with one or two little points; and an inner lamina
of shell sometimes depends beneath the outer lamina, to which the
opercular membrane is attached, as may be seen in the figure (6 _d_)
of the external surface of the valve. The crests for the depressor
muscles are well developed. The tergal margin is broadly inflected,
and the articular ridge prominent, making the articular furrow deep.
The spur is rather narrow, and is either (6 _d_, 6 _e_) bluntly or
sharply pointed. The basal margin on the scutal side of the spur, is
hollowed out, but to a variable depth.
_Structure of the Parietes and Radii._--The parietes are thin; at
their internal basal edges they are finely striated in longitudinal
lines. The radii are solid, with quite smooth edges; they are
generally covered by the epidermis. The sutural edges of the alæ are
likewise smooth, these are added to largely during the diametric
growth; and their summits, as already stated, are much less oblique
than the summits of the radii. The internal surface of the shell
is smooth, and is tinted pale dull purple. The lower edge of the
sheath can hardly be said to be free. The carinal margins of the
compartments project a little inwards.
_Mouth_: the labrum is deeply notched, and supports five little teeth
on each side; the palpi are thickly clothed with spines on their
inner sides; the mandibles have five or only four teeth: the maxillæ
are notched, and the outer maxillæ bilobed.
_Cirri_: the first pair has one ramus nearly twice as long as the
other. The three posterior cirri are elongated, and each segment
supports five or six pairs of long spines, with a few minute
intermediate bristles.
2. ELMINIUS MODESTUS. Pl. 12, fig. 1 _a_-1 _e_.
_Shell folded longitudinally, greenish or white: radii of moderate
breadth, smooth edged: scutum without an adductor ridge: tergum narrow,
with the spur confluent with the basi-scutal angle._
_Hab._--New South Wales; Van Diemen's Land; New Zealand; very
commonly attached to littoral shells and rocks; associated with
_Balanus trigonus_ and _vestitus_; Mus. Brit., Cuming, Stutchbury,
Darwin.
_General Appearance._--Shell conical, generally rather steep,
occasionally depressed: walls longitudinally folded, sometimes very
deeply, sometimes only to a slight degree: colour dull greenish or
white. Radii of moderate width, with their summits very oblique,
smooth and slightly arched: alæ much exposed, with their summits
less oblique than those of the radii: the portion added to the alæ
during the diametric growth differs much in appearance from the other
portion. The scuta have the growth ridges but little prominent;
they are crossed by a faint longitudinal band of gray. The largest
specimen out of the many which I have seen, was under .4 of an inch
in basal diameter.
_Scuta_, destitute of an adductor ridge and of crests for the
depressor muscles: the articular ridge is moderate; but the
articular furrow is rather wide: the internal occludent margin is
much thickened. The _Terga_ are narrow and small; they are somewhat
variable in shape, caused by the degree to which the basal margin is
hollowed out (fig. 1 _c_-1 _e_), and likewise by the extent to which
the upper end of the valve has been worn away. No spur is apparent,
for it is confluent with the basi-scutal angle of the valve. The
articular ridge is very prominent, and runs down to the basi-scutal
angle; and as the valve in this part is extremely narrow, with the
spur not developed, it here assumes a channelled structure. The
basi-carinal corner of the valve is furnished with rather feeble
crests for the depressor muscles, and in those varieties in which the
basal margin is much hollowed out, this part is remarkably narrow.
_Structure of the Parietes and Radii._--The internal basal edges of
the parietes and the sutural edges of the radii and alæ, are all
smooth. The lower edge of the sheath depends freely. In the green
varieties the colour is most distinct on the internal surface of the
shell. The four compartments separate very easily when the shell has
been ill preserved in spirits, or after a very short immersion in
caustic potash.
_Mouth_, as in _E. Kingii_, excepting that there are only three teeth
on each side of the notch (which is deeper) on the labrum. The cirri
resemble those of _E. Kingii_; the segments in the sixth pair are
equally elongated, and bear five or six pairs of spines.
_Affinities._--This species is closely allied to its South American
representative _E. Kingii_; the differences consist in its smaller
size, often greenish colour, more folded walls, and narrower radii:
the internal basal edges, also, of the parietes are here smooth,
instead of being striated, as in _E. Kingii_. The terga present even
more obvious differences, in their narrowness, channelled under
surface, and in the absence of the spur, or more properly in its
confluence with the basi-scutal angle of the valve.
3. ELMINIUS PLICATUS. Pl. 12, fig. 2 _a_-2 _f_.
ELMINIUS PLICATUS. _J. E. Gray._ Appendix to Dieffenbach's Travels in
New Zealand, p. 269, 1843.
_Shell deeply folded longitudinally, corroded, coloured in parts
orange: radii very narrow, with their edges sinuous, and slightly
dentated: scutum having an adductor ridge._
_Hab._--New Zealand; New South Wales(?). Attached to rocks, often
coated by _Chamæsipho columna_; Mus. Brit. and Cuming.
_General Appearance._--Shell tubulo-conical, or conical, rarely
depressed; strong, rugged, coloured in parts bright orange; deeply
plicated longitudinally, but with the upper parts corroded and
smooth. Orifice large. The sutures are indistinct and almost
obliterated; the radii, when most developed, are narrow. Some
specimens have their whole surface deeply corroded; in which case
they are finely striated longitudinally, or pitted, and are of a
gray or brown colour. The largest specimens are one inch in basal
diameter, but one depressed specimen was 1.3 in diameter; another was
rather under one inch in diameter, and one inch in height.
_Scuta_; beginning with the common tubulo-conical and not much
corroded specimens, the valve (fig. 2 _c_) is moderately elongated,
but in a rather variable degree. A prominent adductor ridge runs,
from a little above a middle point of the basal margin, along
the slightly prominent articular ridge: the articular furrow
is moderately wide. There are distinct crests for the lateral
depressores. In the conical, corroded specimens, the scuta (fig. 2
_e_) are considerably broader, with the articular ridge much more
prominent, and the furrow wider: in one such specimen, there were
crests for the rostral depressor muscle.
The _Terga_, in the commoner variety, resemble those of _Tetraclita
porosa_; the spur adjoins the basi-scutal angle of the valve: the
articular ridge is moderately prominent, and the furrow moderately
deep. The valve is beaked, with an unusually large internal tube
for the thread of corium: the beak, however, is often worn away. In
the depressed much corroded specimens, the terga (fig. 2 _f_), like
the scuta, are broader and shorter than in the commoner variety;
and the spur more especially is broader. The scutal margin is much
more widely inflected, and the articular ridge much more prominent;
consequently the articular furrow is much deeper.
_Structure of the parietes and radii._--The orange or yolk-of-egg
colour, which is so conspicuous a character in the present species,
is due to a layer of shell between the inner and outer lamina, and
is exposed only by the corrosion of the latter. Hence the very base
of the shell is not of this colour; nor are the uppermost and still
more deeply corroded portions, for here the orange-coloured layer has
been removed. The sheath is orange-coloured, and the operculum, to a
certain extent, is similarly tinted. The epidermis on the parietes,
where preserved quite close to the basis, supports remarkably strong
spines, about 1/100th of an inch in length. The basal internal edges
of the walls are rather coarsely striated with irregular short ridges
and sub-cylindrical points; and the walls in most of the specimens
are regularly and deeply folded, which, with the little ridges, gives
the appearance represented in fig. 2 _b_, Pl. 12. I have stated,
under the Genus, that in the corroded and depressed specimens, the
walls are rendered extremely thick by the inward production and
upward extension of these same ridges and points; the under surface
of the shell acquiring almost the appearance of _Chelonobia caretta_.
The Radii are often not developed, even the sutures being obscure;
when most developed, they are narrow, with the outer lamina along the
growing edge sinuous, giving to the sutures a crenated appearance.
The sinuosities on the growing edge generally send inwards short
ridges or septa, like those on the sutural edges of the radii in most
Balanidæ, but of which there is no trace in the other species of
Elminius. In very minute, colourless specimens, about the 1/20th of
an inch in diameter, the radii are quite smooth-edged. The alæ have
their edges strongly crenated. The lower edge of the sheath depends
freely.
_Mouth_: the labrum shows some tendency to be bullate; the notch is
broad and shallow: the palpi have a thick brush of bristles on their
inner sides. The mandibles have four or five teeth. In the maxillæ,
the upper spines above the broad notch, are very strong. In the outer
maxillæ, the two lobes are widely separated.
_Cirri_: in the first pair, one ramus is about one fifth longer
than the other. In the third pair, the posterior ramus is one
fourth longer than the anterior ramus, and its terminal segments
are tapering, each having a single circle of bristles: the other
segments, and those of the shorter ramus, support many coarsely
pectinated spines. In the sixth cirrus, the segments are protuberant
in front, and carry four pairs of stout spines, with a tuft of fine
bristles between them.
_Affinities._--This species differs considerably from the first two
of the genus. In several characters it approaches nearer than the
other species to Tetraclita, especially to _T. porosa_;--namely,
in the scutum having an adductor ridge and crests for the lateral
depressores, in the whole form of the tergum, in the thick walls
liable to much corrosion, in the narrow radii, and in their edges,
as well as those of the alæ, being crenated; and, lastly, in the
character of the cirri, more especially of the third pair, with
its coarsely pectinated spines. It also approaches, in all its
characters, _Balanus imperator_ and _flosculus_.
4. ELMINIUS SIMPLEX. Pl. 12, fig. 3.
_Shell ribbed longitudinally, dirty white; radii extremely narrow,
smooth-edged; scutum having an adductor ridge._
_Hab._--New South Wales (Sydney and Twofold Bay); Van Diemen's Land;
tidal rocks, often attached to other Cirripedes, and associated
with _Balanus nigrescens_, _Tetraclita purpurascens_, _Catophragmus
polymerus_; Mus. Brit., Cuming, Stutchbury, and Darwin.
This species, of which I have seen specimens from the above three
localities, all exactly agreeing with each other, is perhaps the
Australian representative of _E. plicatus_, which seems to be confined
to New Zealand.[113] In all essential points it comes so near that
species, that I shall make the greater part of my description
comparative.
[113] I am bound to state that I have seen two specimens of _E.
plicatus_ marked Sydney, and one marked Moreton Bay, but in both
cases the collectors had visited New Zealand, so that a wrong
habitat by mistake might easily have got attached to the specimens
in question.
_General Appearance._--In external appearance there is considerable
difference from _E. plicatus_, for _E. simplex_ is generally of a
regular conical shape, of a dirty-white colour, with the surface well
preserved, having moderately wide, not very prominent longitudinal
ribs. The orifice is rather small and pentagonal. The radii are
extremely narrow or linear, with quite smooth edges; the sutures,
however, are always very distinct, and in the upper part, the alæ
are generally rather widely exposed, as viewed from the outside.
The largest specimen which I have seen was .7 of an inch in basal
diameter.
The _opercular valves_ are closely similar to those of _E. plicatus_,
but the _scutum_ is generally a little more elongated, and the
articular furrow not so deep: in accordance with this last fact,
the articular ridge in the _tergum_ is not so prominent as in _E.
plicatus_; but we have seen that these several characters are highly
variable in _E. plicatus_. The slope of the basal margin of the
tergum towards the spur varies in the present species, in a strictly
analogous manner, as it does in _Tetraclita porosa_.
_Structure of the Parietes and Radii._--The parietes are not so thick
as in _E. plicatus_; internally they are tinted pale purple; when
broken transversely, a row of microscopically minute orange-coloured
dots can generally be distinguished between the outer and inner
laminæ; and these evidently represent the orange-coloured layer in
_E. plicatus_. The sheath also exhibits a faint tinge of orange. The
radii are very narrow, and are quite smooth-edged, differently from
in _E. plicatus_. The edges of the alæ barely exhibit a trace of
being crenated.
In the body I could perceive no difference from _E. plicatus_,
excepting that in the third pair of cirri the two rami are like each
other, and do not support any coarsely pectinated, only serrated,
spines; but after what we have seen on the variability of these very
same characters in _Tetraclita porosa_, I dare not trust to them. The
three posterior pairs of cirri, also, seem here to be more elongated
in proportion to the others, than in _E. plicatus_.
_Affinities._--It is certain that this species is most closely allied
to _E. plicatus_; but as I have seen many specimens of the latter
brought by different persons from New Zealand, and as I have observed
in them no approach to the characters of _E. simplex_, which, in
specimens from three localities, also appear to be constant, I have
considered the two forms as specifically distinct. The present
species differs from _E. plicatus_, in its white, conical, moderately
ribbed, well preserved, smaller shell; and more especially in the
orange-coloured intermediate lamina of _E. plicatus_ being here
represented only by microscopically minute dots. But the radii being
smooth-edged, is the most important differential character, though in
_E. plicatus_, during its earliest growth, whilst still immature and
colourless, the radii are likewise smooth-edged.
5. _Genus_--PYRGOMA.
PYRGOMA. _Leach._ Journal de Physique, tom. 85, 1817.
BOSCIA. _Ferussac._ Dict. Classique d'Hist. Naturelle, 1822.
SAVIGNIUM. _Leach._ Zoological Journal, vol. 2, July, 1825.
MEGATREMA. _Ib._ Ib.
ADNA. _Ib._ Ib.
DARACIA. _J. E. Gray._ Annals of Phil. (new series), August, 1825.
CREUSIA. _De Blainville._ Dict. Sc. Nat., Pl. 116, 1816-30.
NOBIA. _G. B. Sowerby, junr._ Conchological Manual,[114] 1839.
[114] The name, Nobia, is given in this work on the authority of
Leach, but this must be a mistake, probably caused by some MS.
name, (that fertile source of error in nomenclature), having been
used.
_Shell formed of a single piece; basis cup-formed, or sub-cylindrical,
attached to corals._
_Distribution_, imbedded in corals, chiefly in the tropical seas,
round the world.
I feel no hesitation in including the above several genera in one
genus. In external appearance the _P. monticulariæ_ (Pl. 13, fig. 5
_a_), which forms the genus Daracia of Gray, is the most distinct, but
it is so intimately allied to the two, indeed to the three, foregoing
species, that it cannot be separated from them. Of the first five
species, _P. grande_, _conjugatum_, and _cancellatum_, form a graduated
series, but with the steps very distinct, the chief difference being
in the length of the spur of the tergum; for the fact of the scutum
and tergum being calcified together in _P. grande_ and _conjugatum_,
and distinct in _P. cancellatum_, is certainly unimportant, as may
be inferred from what we shall see in comparing together the last
four species of the genus, and from what we shall see in _Creusia
spinulosa_. The three above-mentioned graduated species are connected
with the last four species of the genus, by several points of
resemblance between _P. grande_ and _crenatum_. The first two species,
namely, _P. Anglicum_ and _Stokesii_, are the most closely related
together, and may indeed possibly be identical; these two, in all the
characters derived from the opercular valves, resemble Balanus and
other ordinary forms, and for this very reason they have some claims
to be generically separated from the other species of Pyrgoma; for in
these latter, the opercular valves seem to have broken loose from all
law, presenting a greater diversity in character than do all the other
species of Balaninæ and Chthamalinæ taken together.
_General Appearance._--The shell consists of a single piece, generally
without any suture, even on the internal surface; and this is the case,
at least in _P. Anglicum_, in extremely young colourless specimens:
nevertheless, in some specimens of this very species, and of _P.
conjugatum_, there were traces of two, but only two, sutures on the
sheath, one on each side towards its carinal end. The shell is much
depressed or actually flat; and I have seen specimens even slightly
concave; in _P. Anglicum_, however, the shell is steeply conical. The
outline is generally oval; but in _P. monticulariæ_ it is extremely
irregular. The surface is generally furnished with more or less
prominent ridges, radiating from the orifice, which is oval and small;
sometimes, as in _P. monticulariæ_, excessively small. The colour is
white, or pinkish-purple. Most of the species are small, but I have
seen specimens of _P. grande_ three quarters of an inch in diameter in
the longer axis, and, including the deep, almost tubular, basis, more
than three inches in length or depth.
_Opercular Valves._--In three species, viz., _P. conjugatum_, (Pl. 12,
fig. 7 _c_), _grande_, and _monticulariæ_, the scuta and terga, on
each side, are calcified together so perfectly, that there is no trace
of a suture or line of junction: in _P. milleporæ_, these valves are
generally slightly calcified together, but with the suture distinct.
The _Scuta_ differ so much in shape in the different species, that
little can be said of them in common: in _P. Anglicum_ and _Stokesii_
they resemble those of Balanus; but in the other species they are
much more elongated than is usual, and this is carried to an extreme
in the last four species; this elongation is due to a great increase
in breadth, as may be inferred from the position of the apex of the
valve, and from the direction of the lines of growth. But the two most
remarkable characters are, first,--the extraordinary development of
the adductor ridge, so that, in _P. conjugatum_, _cancellatum_ (Pl.
12, fig. 5 _c_), _grande_, and _crenatum_, it extends considerably
beneath the basal margin, being produced, in the first two species, at
the rostral angle, into a point; at the tergal end of the valve, the
adductor ridge, when thus much developed, blends into the articular
ridge. The second very remarkable character is the addition of a
special ledge along the occludent margin of the scutum, and along the
carinal margin of the tergum, which I will call the occludent ledges
(_limbus occludens_), and which serve to close the orifice leading
into the sack. The occludent ledge is small in _P. grande_, and is
clothed with thick yellow spines, giving it a brush-like appearance:
in _P. crenatum_ and _dentatum_ it is largely developed, the ledge on
the scutum being articulated with that on the tergum, as shown in Pl.
13, fig. 4 _a_, 4 _b_,--the ledges being here and elsewhere marked
by little bristly points. In _P. monticulariæ_, however, this ledge
arrives at its maximum development (Pl. 13, fig. 5 _f_), the rest of
the valve (the scuta and terga being here, as in _P. grande_, calcified
together) being reduced to a mere basal edge or border. Excepting for
the adductor muscle, the depressions or crests for the other muscles,
both on the scuta and terga, are hardly developed.
_Terga_: these, as in the case of the scuta, differ so much in shape
in the several species, that little can be said of them in common.
In _P. Anglicum_ and _Stokesii_, they are of the normal shape; in
_P. cancellatum_ (Pl. 12, fig. 5 _d_) this, to a certain extent, is
likewise the case, but the spur is produced to a quite extraordinary
length. In _P. grande_ (Pl. 13, fig. 1 _b_) there is no distinct spur,
and the whole valve is square. In _P. milleporæ_ (2 _f_) there is no
spur, and the valve is arched and triangular. In _P. crenatum_ the spur
is broad, rounded, and depressed (fig. 4 _b_), with the carino-basal
portion of the valve reduced to a mere border, barely distinguishable
from the great occludent ledge. In _P. monticulariæ_ there is no spur,
and the whole valve forms a mere border to the occludent ledge; and,
lastly, in _P. dentatum_, the valve is extremely variable in shape
(fig. 3 _c_, 3 _d_, 3 _f_), and on its internal surface (fig. 3 _g_)
there is an inwardly projecting, most singular and anomalous, tooth.
Hence we see how wonderfully variable the terga are in this genus.
_Structure of the Walls._--The shell consisting, as has been stated,
of a single piece, is generally thick. From the close alliance between
this genus and _Creusia_, it is probable that the shell, if examined
immediately after the metamorphosis, would be found to show traces
of being formed of four compartments. The walls are either solid
or porose; their basal margin is formed by strong crenated ridges,
answering to the longitudinal septa in Balanus; but these in _P.
monticulariæ_ are modified into a very irregular surface. The internal
surface of the shell is generally smooth, or only slightly ribbed. The
sheath has its lower edge free in _P. Anglicum_ and _Stokesii_, and
in a slight degree in _P. milleporæ_, but in the other species it is
closely attached to the walls. In _P. monticulariæ_ the sheath might
easily be overlooked. In _P. Anglicum_, _grande_, and _crenatum_, it
descends almost to the basal margin of the depressed shell, and as the
opercular valves and membrane are attached to the lower edge of the
sheath, the animal's body necessarily comes to be almost exclusively
lodged in the cup-formed basis. In _P. grande_ and _conjugatum_, the
lines of growth in the sheath are bent upwards on each side, at points
corresponding with the line of union between the scutum and tergum, in
a manner I have not seen in any other cirripede; and this sometimes
gives the appearance of two lateral sutures. I may here remark, that
the manner of growth in Pyrgoma is almost the converse of that in
Balanus, Tetraclita, and other allied genera; for in these latter, the
basis increases in diameter, and the shell chiefly in height; whilst in
Pyrgoma, the shell, from being so flat, increases almost exclusively in
diameter, whereas it is the basal cup which is added to in height or
depth.
_Basis._--This in all the species is more or less regularly cup-formed
or sub-cylindrical. In _P. grande_ it penetrates the coral to a
surprising depth; in _P. monticulariæ_ it is irregular in outline,
corresponding with the shell. The basis is generally almost wholly
imbedded in the coral; but this is not the case with _P. Anglicum_,
in which the basis is generally exserted, as it is in a slight degree
in _P. grande_. The shelly layer forming the basis, in most of the
species, is very thin, and is finely plicated owing to its edge
folding between the ridges or septa that form the basal edges of the
shell; this is very conspicuous in _P. cancellatum_. The basis is not
permeated by pores, except in _P. Anglicum_. In some sessile cirripedes
a cleft, covered only by membrane, may be observed all round between
the lower edge of the shell and the basis; a cleft of this nature
is rather conspicuous in _P. crenatum_, so that small portions of
the septa on the internal surface of the walls can be seen from the
outside. In _P. monticulariæ_, an analogous structure, developed to an
extreme degree, presents a very different and unique appearance; the
shell is nearly flat, and the smooth outer lamina does not nearly reach
to the circumference, a wide border being thus left exposed, which is
roughened (Pl. 13, fig. 5 _a_, 5 _c_, 5 _d_, 5 _e_) by the exposure of
the irregular septa. I have not seen a fresh specimen, but there cannot
be a doubt that this border is properly covered by membrane.
_Animal's Body._--From some cause, perhaps from the corals in which
the species of Pyrgoma are imbedded, long remaining damp, the internal
organs are generally badly preserved. I have received, in spirits of
wine, only _P. Anglicum_, but I have examined dry specimens, in a
tolerable condition, of _P. milleporæ_ and _crenatum_. As neither
the mouth nor cirri, in these three species, offer any noticeable
characters, distinct from those in Balanus or Acasta, my ignorance of
these organs in the other six species is not important. In the above
three species the labrum is deeply notched, with about three teeth on
each side of the notch, except in _P. milleporæ_, in which the number
is six. In all, the mandibles have five teeth, the two lower ones being
small: the maxillæ are not notched: the outer maxillæ are bilobed. In
the _Cirri_, the rami in the first pair are very unequal in length,
the segments being slightly protuberant in the shorter ramus. On the
segments of the posterior cirri there are four pairs of spines in _P.
Anglicum_, and three pairs (of which the second and third are short)
in _P. milleporæ_. At the dorsal basis of the penis there is a small
straight projecting point.
_Affinities._--The species (with the exception of the first two) are
much more distinct from each other, and more easily determined than
is usual with sessile cirripedes; it is, however, quite useless to
attempt naming the species without disarticulating and cleaning the
opercular valves. Although these valves differ so greatly in some of
the species from those of Balanus and the allied forms, the genus
itself, as a whole, does not differ much, except in the shell not being
divided into compartments, and in the basis being cup-formed and not
generally permeated by pores,--these latter characters being in common
with Acasta. With respect to the absence of separate compartments,
it should be remembered, that in the same species of Tetraclita we
have individuals with the four compartments distinct and furnished
with radii, and other individuals without any trace of a suture
externally,--the outer lamina of shell (though not the inner) having
become completely confluent all round. At the commencement of this
description, when giving my reasons for uniting the several proposed
genera into one genus, I gave a sketch of the affinities of the
species: I have only to add, that the following sub-genus Creusia is
closely, perhaps too closely, allied to Pyrgoma.
_Geographical and Geological Distribution._--Most of the species are
inhabitants of the hot coral-growing zones, in both the eastern and
western hemispheres, but more especially, as it would appear, in the
East Indian Archipelago. From the habits of the corals, most of the
species must be inhabitants of shallow water. _Pyrgoma Anglicum_,
however, is an inhabitant of deep water on the southern shores of
England, whereas at St. Jago, in the Cape de Verde Islands, I myself
collected it adhering to a Caryophyllia, within the tidal limits. This
same species existed on the shores of England during the Coralline Crag
period; and at this epoch it attained a larger size than at present.
Two species of the genus, according to Sismonda, are found in the
tertiary beds of Piedmont. Mr. Stutchbury, who is so well acquainted
with recent cirripedes, informs me that he has for many years examined
fossil Secondary corals, in the expectation of finding imbedded,
species of this genus or of some allied form, but without success. The
same species of Pyrgoma is by no means always confined to the same
coral: I have seen _P. crenatum_ on four or five different corals,
and _P. Anglicum_ on at least three kinds: on the other hand, I have
seen _P. milleporæ_ only on the _Millepora complanata_ (a member,
as I believe, of the vegetable kingdom), and _P. monticulariæ_ on a
_Monticularia_ from near Singapore.
1. PYRGOMA ANGLICUM. Pl. 12, fig. 4 _a_-4 _c_.
PYRGOMA ANGLICA. _G. B. Sowerby._ (sine descript.) Genera of Recent
and Fossil Shells, fig. 7, No. 18, Sept. 1823.
MEGATREMA (ADNA) ANGLICA. _J. E. Gray._ Annals of Philosoph. (new
series), vol. x, Aug. 1825.
PYRGOMA SULCATUM. _Philippi._ Enumeratio Molluscorum Siciliæ, Tab.
12, fig. 24, (1836).
---- ANGLICA. _Brown._ Illustrations of Conchology, (2d edit.,
1844), Tab. 53, fig. 27-29.
_Shell steeply conical, purplish red: orifice oval, narrow: basis
permeated by pores, generally exserted out of the coral: scutum and
tergum sub-triangular._
_Hab._--South coast of England and of Ireland, (12 to 45 fathoms,
Forbes and MacAndrew); Sicily; Madeira; St. Jago, Cape de Verde
Islands; generally attached to the edge of the cup of a Caryophyllia,
in deep water, but at St. Jago within the tidal limits; Mus. Brit.,
Cuming, Lowe, &c.
_Fossil_ in the Coralline Crag, Ramsholt; Mus. S. Wood.
_General Appearance._--Shell steeply conical, slightly compressed,
the lower part with rounded, approximate, radiating ribs: colour
dull purplish-red: orifice oval, small, and narrow. The basis is not
deeply conical, and occasionally is even flat. Generally it stands
exserted; but in the Coralline Crag specimens, it is almost wholly
imbedded. Externally it is furnished with ribs corresponding with
those on the shell. The largest recent specimen which I have seen,
from St. Jago, was .22 of an inch in basal diameter; but some few of
the British specimens are nearly as large, and one of the fossils
from the Coralline Crag a little larger.
The _Scuta_ and _Terga_ are of the ordinary shape of these valves in
Balanus and its allies. _Scuta_ triangular, with the basal margin a
little curved and protuberant: adductor and articular ridges distinct
from each other, moderately prominent: there is a small hollow for
the lateral depressor muscle. _Terga_ triangular, with the spur
rather narrow, moderately long, placed near, but not confluent with
the basi-scutal angle of the valve. The basal margin forms an angle
rather above a right angle with the spur. Internally, the articular
ridge and crests for the depressor muscles, feebly developed.
_Internal Structure of the Shell and Basis._--Internally, the shell
is ribbed more or less prominently. The lower edge of the sheath,
which is reddish, and extends far down the walls, seems always to
project freely. In several specimens there were on each side, at
the carinal end of the shell, a trace of a suture, which could
be perceived only on the sheath. The basis appears always to be
permeated by minute tubes or pores, though these are sometimes rather
difficult to be seen.
2. PYRGOMA STOKESII. Pl. 12, fig. 6.
MEGATREMA STOKESII. _J. E. Gray._ (sine descript. aut figurâ) Annals
of Philosophy, (new series), vol. 10, Aug. 1825.
_Shell moderately conical, pale-purplish red; orifice oval: basis not
permeated by pores, deeply imbedded in the coral: scutum and tergum
sub-triangular._
_Hab._--Imbedded in the Mycedia (Agaricia) agaricites; therefore from
the West Indies;[115] Brit. Mus. and Stutchbury.
[115] I am greatly indebted to Mr. Dana for having named for me the
coral in which this species was imbedded, and informing me that it
is a West Indian species.
This species comes so close to the last, that I am not sure that I have
acted rightly in retaining it, but I think that it is distinct; and in
this case, it is the representative, on the other side of the Atlantic,
of _P. Anglicum_ of our own side. It will be sufficient to point out
the few points of difference. The shell is much more depressed, with
the orifice oval, larger, and not so narrow. It is apparently of a
paler red, and the radiating ribs perhaps not so prominent. The basis
offers the most important difference, being deeply imbedded in the
coral; and there is not the least appearance of the thin shelly layer,
of which it is composed, being permeated by pores, as, we have seen,
is always the case with _P. Anglicum_. As in this latter species,
the sheath here depends freely. The opercular valves are closely
similar; but in the scutum, the adductor ridge occupies a rather more
central position; and in the tergum, the basal margin is more inclined
towards, or forms a greater angle with, the spur: these differences, by
themselves, I consider quite insufficient to characterise a species;
but conjoined with the flatter shell, the larger orifice, the more
deeply imbedded and non-porose basis, they may, I think, be admitted as
specific. In dimensions, this species seems to attain a slightly larger
size than _P. Anglicum_, for several specimens were .22 of an inch in
diameter.
3. PYRGOMA CANCELLATUM. Pl. 12, fig. 5 _a_-5 _f_.
PYRGOMA CANCELLATUM. _Leach_ (!). Encyclop. Brit., Supplement, vol.
3, Pl. 57, 1824.
---- LOBATA. _J. E. Gray_ (!). Annals of Philosophy, (new series),
vol. 10, 1825.
CREUSIA RAYONNANTE. _De Blainville._ Dict. Sc. Nat. (sine descript.),
Pl. 116, fig. 7, _a_, _b_.
_Shell with the circumference generally lobed: scutum elongated, with
the adductor ridge descending far below the basal margin, and produced
at the rostral end into a square point; tergum with the spur four times
as long as the upper part of the valve._
_Hab._--Imbedded in a Gemmipora, probably from the West Indies;[116]
Mus. Brit. and Cuming.
[116] In this case I am again indebted to Mr. Dana for naming the
coral; he informs me that the genus is found in the Pacific and
East Indies; the specimen sent he believes is the _G. cinerascens_.
_Appearance and Structure of Shell._--Shell nearly flat, sometimes
tinted dull dirty purple, with the surface marked by slight, broad,
approximate ridges, the ends of which form considerable projections,
giving to the shell, when not too much encrusted by coral, a lobed
border. In young specimens (.15 of an inch in diameter) some of
the points projected half as much as the semi-diameter of the
shell, giving it a radiating appearance. The orifice is oval and
rather small, but of variable size. The shell is thick, and, near
the outer lamina, is penetrated by pores: the internal surface is
smooth. The sheath (fig. 5 _b_) extends down but a short distance
from the orifice; it is closely attached to the walls: the lines of
growth, at a point on each side, bend a little downwards (instead of
upwards, as in the two following species), and hence the lower edge
of the sheath irregularly projects downwards on each side. The basal
cup, internally, is plicated, the hollows corresponding with the
projecting, longitudinal, parietal septa, which form the lobed border
of the shell. The largest specimen which I have seen, was rather
above .4 of an inch in diameter.
The _Scuta_ and _Terga_ are not calcified together: they are both
much elongated. _Scutum_ (fig. 5 _c_, 5 _e_).--For the first time in
the genus, or indeed in the family, this valve presents a remarkable
character in the adductor ridge being immensely developed, so as to
project far below the ordinary basal margin. At the rostral end, it
at first appears to project even more than it really does, for the
toothed occludent margin is in fact a prolongation of the true valve,
as distinct from the adductor plate. Excluding this very narrow,
prolonged, occludent margin, the adductor plate projects for a
length equalling the rest of the valve. Along the tergal margin, the
adductor plate is united to the articular ridge; and at the rostral
end, it is produced into a square tooth, whence a square-edged ridge
extends on the surface of the plate upwards to the ordinary basal
margin of the valve. The exact shape of this adductor plate varies
a little, as does the degree to which it is closely attached to the
ordinary basal edge of the valve. The valve, as distinct from the
adductor plate, is narrow, with the basal margin regularly curved.
The articular ridge is very prominent.
_Terga_ extremely narrow, linear, consisting in chief part of the
spur, which is fully four fifths of the entire length. Externally
(fig. 5 _d_) the valve is furrowed, with the edges more or less
folded in along the spur. The upper or ordinary part of the valve is
about one third wider than the spur. The basi-carinal angle is sharp,
owing to the basal margin being a little hollowed out. A special
plate of shell (fig. 5 _f_), hollow under its basal edge, runs from
the carinal margin to the articular ridge, which latter is situated
in the middle of the valve, and against which the articular ridge of
the scutum abuts. The spur is central; its end is bluntly pointed.
The total length of the tergum rather exceeds that of the scutum, the
produced adductor ridge being included in the latter.
_Affinities._--Under _P. grande_ I shall make a few remarks, showing
that in several characters _P. cancellatum_ and _grande_ are at
opposite ends of a short series, with _P. conjugatum_ intermediate
between them.
4. PYRGOMA CONJUGATUM.[117] Pl. 12, fig. 7 _a_-7 _c_.
[117] Dr. Gray thinks this is the _Pyrgoma stellata_, of Chenu,
('Illust. Conch.'); it may be so; but the figure given of the
shell will do equally well or rather better for the _Pyrgomum
dentatum_ of this work, and for some varieties of _P. crenatum_.
Without a careful description of the opercular valves, it is really
impossible to recognise, with any approach to certainty, the
species of this genus.
_Shell nearly flat with approximate radiating ridges: scutum and tergum
calcified together without any suture: scutum with the adductor ridge
descending below the basal margin, and produced at the rostral end
into a point: tergum with the spur about as large as the upper part of
valve._
_Hab._--Red Sea; Brit. Mus. and Cuming.
_Appearance and Structure of Shell._--Shell white, or with a tinge of
purple; nearly flat, with moderately prominent, narrow, approximate
ridges, radiating from the orifice, which is oval, rather narrow,
and not very small. In the largest specimens the ridges are less
prominent than in the figure given. The walls are thick, and not at
all porose: the sheath extends down almost to the base of the shell,
and its lower edge is closely attached to the walls: on each side,
towards the carina, there is a trace of a suture, and the lines of
growth on the sheath are here a little upturned. The basis is deeply
imbedded and internally furrowed; the calcareous layer forming it, is
thin. The largest specimen was .4 of an inch in the longer diameter.
The _Scuta_ and _Terga_ (fig. 7 _b_, 7 _c_), in all the specimens
which I have seen, are calcified together, with no trace of a
suture. There is, however, a slight furrow, which, I believe, marks
the normal line of separation between the two valves; and in the
following description this is assumed to be the case. The _Scutum_
has the adductor ridge greatly developed, so as to project below
the ordinary basal margin to a distance as great as the height of
the valve. At the rostral end, this adductor ridge or plate is
produced into a point; and at the tergal end, it is blended with the
articular ridge, and united to the inner face of the tergum. That
portion of the scutum which corresponds with the valve in ordinary
cases, and alone is externally visible as long as the operculum is
united by the opercular membrane to the sheath, is narrow, with the
basal margin considerably hollowed out: the occludent edge is formed
into thick teeth. The _Tergum_ is elongated, rather exceeding in
length the scutum, the latter being measured from the apex to the
rostral projection of the adductor plate. The surface of the valve is
depressed in the line of the spur, with the basal end of the latter
bluntly pointed. A very slight flexure (fig. 7 _c_) on the basal
margin indicates where we may believe the spur to commence, showing
that it rather exceeds in length the whole upper part of the valve.
The lines of growth obscurely indicate a tendency to the formation of
a slight "occludent ledge" along the carinal margin. Traces are just
visible of crests for the attachment of the tergal depressor muscles.
5. PYRGOMA GRANDE. Pl. 13, fig. 1 _a_-1 _d_.
NOBIA GRANDIS. _G. B. Sowerby, junr._ (sine descript.) Conchological
Manual, fig. 29, 1839.[118]
CREUSIA GRANDIS. _Chenu._ Illust. Conch., Tab. 1, fig. 2 _a_, sed
non, fig. 2.
[118] It is quite possible that this may be the _Balanus
duploconus_ of Lamarck, but with such a character as the following,
who can recognise a species? "_B. testæ parte supremâ univalvi,
indivisâ, convexâ: inferiore turbinatâ, non clausâ: aperturâ
ellipticâ._ L'exemplaire est sans opercule et incomplet."
_Shell moderately convex, nearly smooth: scutum and tergum calcified
together without any suture: scutum furnished with a small occludent
ledge, with the adductor ridge descending below the basal margin:
tergum square without a spur._
_Hab._--Singapore and East Indian Archipelago; Mus. Brit., Cuming,
Stutchbury; imbedded in two kinds of coral.
_Appearance and Structure of Shell and Basis._--The shell is conical,
though to a variable degree, and sometimes is much depressed. The
surface is smooth, with only traces of narrow approximate ridges.
The colour is white, often with a tinge of dark purple. The orifice
is oval, and moderately large. The shell and a small portion of
the basis usually stand exserted above the coral. The walls are
of variable thickness; when thick, the pores, by which they are
permeated, are but little apparent; sometimes there is more than a
single row of pores. The points of the septa on the basal edge of the
shell are small. The internal surface of the shell is smooth. The
sheath is closely attached to the walls, and descends nearly to the
basis; on each side its lines of growth are slightly upturned. The
basis is deeply cup-formed or cylindrical, and in section oval like
the shell; it penetrates the coral to a very remarkable depth,--in
one instance to three inches. The shelly layer forming it, is thin,
finely furrowed, and not permeated by pores. This is the largest
species in the genus; one specimen was three quarters of an inch in
its longer diameter, and above three inches in length.
The _Scuta_ and _Terga_ are calcified together, without any trace
of a suture; the line of junction can be inferred only from the
analogy of _P. conjugatum_, in which species the valves have a more
normal character, and are separated by a slight furrow. It may be
seen in the figure (Pl. 13, fig. 1 _d_) of the right and left hand
opercular valves, viewed from vertically above in their proper
relative positions, how abnormal their appearance is, which is partly
caused by the spinose occludent ledges, presently to be described,
but chiefly from the carinal margins of the two terga not being
straight and parallel, as in all other cirripedes, and therefore not
meeting each other, as is usual. In other genera, only the upper
part of the carinal margin of the two terga can be opened for the
exsertion of the cirri, the lower portion being united by membrane;
but here, I have little doubt, from the position of attachment of
the adductor muscle (fig. 1 _c_), so close to and almost on the
terga, that the whole length of the carinal margin of the two terga
is free or disunited for the protrusion of the cirri. This opening
between the two terga evidently cannot be closed, but is probably
filled up, and the animal thus protected, by the dorsal surfaces of
the curled-up cirri; such, I believe, being likewise the case with
some pedunculated cirripedes, as with Conchoderma. The _scutum_ has
a large adductor plate, which extends some little way (namely, about
one quarter of the height of the valve), below the ordinary basal
margin. This latter margin is slightly sinuous, and a little hollowed
out towards the tergal corner of the valve. I believe that the ridge,
which runs down to the basi-scutal corner of the tergum, though
appearing to be part of the scutum, really belongs to the tergum.
The adductor scutorum plate is not, as in the last two species,
produced into a point at the rostral angle; at the tergal end it
blends into the under surface of the tergum. The occludent margin
is coarsely toothed. Rather on the under side of this margin and in
the upper part, there is a narrow occludent ledge, which extends up
beyond the apex of the valve, and thence runs a little way along the
carinal margin of the tergum. This ledge is thickly clothed with
strong, yellowish-brown spines, and hence appears like a brush. It
is remarkable that the cavity for the adductor scutorum muscle is
situated almost on the tergum.
The _tergum_ is of large size, and nearly square; it is, in
appearance, separated from the scutum by a ridge running up to the
apex. The basal margin forms a right angle with the carinal margin,
along which latter margin the lines of growth are upturned, and blend
into the occludent ledge, which is common to the two valves. There
cannot be said to exist any spur, the whole basal margin being almost
straight; nevertheless, on close examination, the ridge which in
appearance separate the scutum and tergum, may, I think, be safely
considered as one side of the spur (which, it should be remembered,
has in all ordinary cases a longitudinal furrow or depression), and
the other side of the spur is, apparently, very feebly indicated by a
slight flexure in the middle of the basal margin. Hence, if the spur
had been developed, it would probably have been half as wide as the
valve. There are no crests for the tergal depressor muscles.
_Affinities._--The present species, with the last two, form an
interesting series. _Pyrgoma grande_ and _conjugatum_, however,
are more closely allied to each other than to _P. cancellatum_. In
the scutum, the whole valve is least elongated, with the adductor
plate least developed, in _P. grande_, and most elongated, with
the adductor plate most developed, in _P. cancellatum_. In the
outline of the tergum the range of shape is quite remarkable; in _P.
conjugatum_, which stands between the other two species, the spur is
rather long, whereas in _P. grande_ there is no spur at all--a very
unusual circumstance--and in _P. cancellatum_, at the other end of
this short series, the spur attains a length wholly unparalleled in
any other cirripede.
6. PYRGOMA MILLEPORÆ.[119] Pl. 13, fig. 2 _a_-2 _f_.
[119] From external aspect I suspect this species to be the
_Creusia madreporarum_, Leach (?), as given in Chenu, 'Illust.
Conch.,' Tab. 1, fig. 6. But I feel sure that Leach has nowhere
published this name; and it may be observed that Chenu gives it
with a mark of doubt. The shell in its imbedded state is only
figured; the opercular valves are not given; and no descriptive
details are added. Under these circumstances I have not adopted
this name; I have, perhaps, been in some degree influenced by the
fact that this species, judging from the many specimens examined by
me, is never imbedded in madrepores, but exclusively in millepores,
so that Chenu or Leach's specific name of _Madreporarum_ is
singularly inappropriate.
_Shell with the orifice narrowly ovate: sheath dark purple: Scutum much
elongated: tergum triangular, convex, without a spur._
_Hab._--Philippine Archipelago (Mindoro Island), Mus. Cuming. Mus.
Brit., Stutchbury, &c. Imbedded in _Millepora complanata_, sometimes
associated with _Balanus Ajax_.
_Appearance and Structure of Shell._--Shell oval, flat, coloured pale
dull purple, or white, with slight and narrow ridges, radiating, from
the orifice, which is not quite central, but is placed rather nearer
to the carinal than to the rostral end of the shell. The orifice is
small and narrow; the carinal end being rounded, and the rostral end
narrower and sharper,--this being the exact reverse of the usual
shape of the orifice in the Balanidæ. The walls are thick, and are
formed of large square tubes. The internal surface of the shell is
smooth. The sheath (fig. 2 _b_) is much more elongated than the
shell, for at the rostral end it extends to the basal margin, and at
the carinal end nearly to it, whereas on the two sides it is some way
distant from the base. The orifice is considerably out of the centre
of the sheath, being placed nearest to the carinal end. The sheath
has its lower edge slightly prominent or free; the lines of growth
are neither turned up nor down on the two sides, as in the last three
species. When nearly full grown the sheath is coloured dark purple,
but when young it is white, hence the upper part is white, surrounded
by an oblong purple ring, and this is surrounded by the white shell.
The basal cup is deep, and internally nearly smooth. The largest
specimens were .3 of an inch in their longer diameter. Great masses
of the Millepora are absolutely studded with this Pyrgoma, with
usually more specimens on one side of the plate or branch than on the
other. They stand in approximately parallel positions, the broad or
carinal ends of the orifices pointing upwards.
_Scuta._--The scuta and terga are closely united, and _are_ often
(perhaps always) in some slight, though variable degree, calcified
together; and hence they often break, rather than separate, at the
line of articulation. The external fissure or line of junction
between them (fig. 2 _c_) is oblique to the longer axis of the
scutum; in the uppermost part of the valve it is sometimes almost
obliterated. The two valves together are nearly as long as the
sheath, and consequently much longer than the orifice of the shell.
The scutum is much elongated, being fully four times as broad as
high. The valve narrows towards the rostral end, but in a variable
degree: the basal margin is hollowed out a little (but to a variable
amount), close to the basi-tergal corner. Along the occludent margin
a slip of the valve, widening downwards, is a little bent inwards,
and this inflected portion is separated from the rest of the surface
by a slightly angular ridge, running from the apex to near the
rostral angle. Internally, at the basi-tergal corner, a slight ridge,
parallel and close to the basal margin (and which can be seen only
when the basal edge of the valve is held upwards), represents the
adductor ridge, which we have seen so largely developed in the last
three species, and shall again see in _P. crenatum_. The articular
ridge (fig. 2 _d_) is extremely prominent, consisting of a more or
less rectangular shoulder.
_Terga_: these are rather small compared to the scuta: they are
triangular and much arched: there is no trace of a spur. Internally
(fig. 2 _f_), the articular ridge is central: there are some vestiges
of crests for the depressor muscle.
_Affinities._--This well-marked species, in the tendency of the
opercular valves to be soldered together, and in the remarkable
absence of a trace of a spur to the tergum, is allied to _P. grande_,
but it is more closely allied to the three following species.
7. PYRGOMA DENTATUM. Pl. 13, fig. 3 _a_-3 _g_.
_Scutum much elongated, with a tooth-like articular projection: tergum
convex, irregularly triangular, sometimes with an imperfect spur, and
on the internal surface with an inwardly projecting tooth; scutum and
tergum furnished with an occludent ledge._
_Var._ (1), 3 _c_, 3 _g_: _tergum, with a sharp internal tooth,
projecting rectangularly inwards_.
_Var._ (2), 3 _d_: _tergum, with a broad blunt internal tooth,
depending beneath the spur-like portion of the valve_.
_Var._ (3), 3 _f_: _tergum, with the basi-carinal end of the valve
truncated, with a small blunt internal tooth projecting rectangularly
inwards_.
_Hab._--Red Sea; Mus. Brit. and Cuming. Also associated with _Pyrgoma
crenatum_, and attached to _Meandrina spongiosa_.[120]
[120] Mr. Dana informs me that he believes that this coral comes
from the West Indies; though the specimens originally described
by him had no label. If this be so, both _Pyrgoma dentatum_ and
_crenatum_ have very wide ranges.
_Appearance and Structure of Shell._--Shell nearly flat, oval,
white or pink, with rather distant prominent ridges radiating from
the moderately large (for the genus) oval orifice. The ridges are
often obscured, and apparently sometimes almost obliterated by the
encrusting coral. Shell permeated near the outer lamina by short
imperfect pores: internal surface smooth: sheath inconspicuous,
descending rather more than half way down the walls; lower edge
closely attached to the walls. Basis deep. Diameter of largest
specimen .3 of an inch.
_Scuta_: these are elongated, but to a very variable degree, some
specimens being quite three times, and some barely twice as broad
as high. I observed this same variability in two sets of specimens,
differing, as we shall presently see, in the form of their terga:
it depends in part, but not wholly, on the varying width of the
occludent ledge, which is sometimes only a fourth, and sometimes
half as high as the rest of the valve. This is the first species
in the genus in which the occludent ledge--a structure peculiar
to the genus--has been amply developed. The basal margin of the
valve is slightly sinuous, and a very little hollowed out near the
basi-tergal corner; it is also very slightly reflexed, the reflexed
portion being separated from the upper part of the valve by a very
slight depression or even furrow. I notice this slightly reflexed
portion, simply as indicating a well marked feature in the basal
margin of the following closely allied species. Internally (3 _g_),
the adductor ridge is thick and slightly prominent, but does not
descend beneath the basal margin: it blends into the articular ridge,
which here projects in a remarkable manner and degree (3 _b_, 3 _e_,
3 _g_), like a rounded tooth. This tooth is in part a development of
the occludent ledge; it varies much in shape. The line of junction
between the scutum and tergum is nearly straight, and nearly at right
angles to their longer axes. In some specimens the scuta and terga
are partially calcified together.
_Terga_: in three sets of specimens the terga differed considerably,
but as in every other respect there was the closest resemblance,
I do not doubt that these are merely varieties. In all three, the
valve is rather small, irregularly sub-triangular in shape, and
externally somewhat convex; in all three, there is an occludent
ledge, of variable width as in the scuta; and in all three, there is
an internal tooth-like projection, of variable form, unlike anything
I have seen in any other cirripede. In the _first variety_ (Pl. 13,
fig. 3 _c_, 3 _g_), the basi-carinal corner of the valve is bluntly
pointed, and a slight flexure separates this portion of the valve
from the other and scutal half, which latter thus exhibits some
tendency to be converted into a spur: on the internal surface (3
_g_) of this spur-like portion of the valve, there is a rather long,
sharp tooth, which projects rectangularly inwards; it is flattened
in a plane at right angles to the longer axis of the scutum and
tergum together: it cannot be seen from the outside. In the _second
variety_, the shape of the valve is not very different (Pl. 13, fig.
3_d_), excepting that the flexure, separating the basi-carinal corner
of the valve from the spur, is deeper; but on the internal face of
the spur, the tooth is far broader than in the first variety, and
is flattened quite differently, viz., in a plane nearly parallel to
the surface of the valve, and instead of projecting rectangularly
inwards, it depends beneath the basal edge of the so-called spur, and
can be seen from the outside. In the _third variety_ (fig. 3 _f_),
the whole carinal end of the valve is cut off, and there can hardly
be said to be any trace of a spur, yet a slight furrow apparently
marks the line of separation between the basi-carinal portion of
the valve, here become very narrow, and the broad, irregular part,
which would have formed the spur had such been developed: on the
internal surface of the latter portion of the valve, there is a very
small, blunt, slightly flattened tooth, projecting inwards, and more
resembling that in the first than that in the second variety.
_Affinities_: under _P. crenatum_ I will point out the diagnosis and
relationship between this and that species.
8. PYRGOMA CRENATUM. Pl. 13, fig. 4 _a_, 4 _b_.
PYRGOMA CRENATUM. _G. B. Sowerby._ Genera of Recent and Fossil
Shells, (No. 218, Sept. 1823), fig. 1 to 6.
_Scutum much elongated, with the adductor ridge descending below the
reflexed basal margin: tergum with a broad depressed spur: scutum and
tergum furnished with a wide occludent ledge._
_Hab._--Philippine Archipelago; Singapore; Mus. Brit., Cuming,
Stutchbury; sometimes associated with _Creusia spinulosa_.
_Appearance and Structure of Shell._--Shell not distinguishable
from that of _P. dentatum_; nearly flat, oval, white, sometimes
pale pink, with rather distant prominent ridges radiating from the
moderately large oval orifice. Shell solid, or permeated near the
outer lamina by short imperfect pores: internal surface smooth. The
sheath descends nearly to the base of the walls; it is but little
conspicuous, and its lower edge is closely attached to the inner
surface of the shell. Basis deep. Diameter of largest specimen under
.3 of an inch.
_Scuta_: in this species the scutum is more abnormal than in any
other Cirripede in the whole family: this is owing both to the
adductor ridge descending far beneath the basal margin, and to the
great development of the occludent ledge; hence the middle and very
narrow portion of the valve alone answers to the scutum, as seen in
other genera. The whole valve, including the adductor ridge and the
occludent ledge, is narrow, being more than twice as broad as high;
but the proportional width varies, owing chiefly (as in the last
species) to the varying width of the occludent ledge. This ledge
commences a little way from the rostral point of the valve, and
gradually widening, extends to the apex, where it is either as high
or twice as high as the rest of the valve. It is articulated by a
convex surface, and by a hollow on its under side with the occludent
ledge of tergum. The scutum cannot be said to have any tergal margin;
without, indeed, the articular surface of the occludent ledge be
thus called. The basal margin is curved, and considerably reflexed,
of which peculiarity we have seen a vestige in the last species:
the reflexion is not well shown, owing to the foreshortening of the
reflexed edge, in fig. 4 _a_: this reflexed edge not being shown,
causes the lines of growth to appear as if they ran more transversely
to the longer axis of the valve, than they really do; for they run
nearly as in the scutum (fig. 3 _e_) of _P. dentatum_. The direction
of these lines of growth is of importance, for they show that the
elongation of the scuta is due to an inordinate increase in their
breadth, as compared to the same valves in ordinary species. The
adductor ridge, having a sinuous margin, runs from near the apex to
near the rostral angle: it descends below the basal margin about as
far as the height of the true valve, excluding the occludent ledge.
Of course this adductor ridge or plate lies beneath the membrane
connecting the opercular valves with the sheath, and is concealed by
it, as long as these valves remain within the shell. The edge of the
occludent ledge is straight, but the edge of the middle portion of
the valve, that is of the true valve, is much bowed.
_Terga_ (fig. 4 _b_): these are of so irregular a shape that they
can hardly be described; they may, however, be said to consist of
two portions joined together, of which the lower portion is a little
elongated transversely, of somewhat variable shape, with part of its
surface considerably depressed (compared with the rest of the valve),
sometimes being even concave: this concave portion apparently answers
to the spur in other cirripedes. At the scutal corner of the valve
there is a shoulder (perhaps answering to the inflected scutal margin
in an ordinary tergum), which locks into a hollow on the under side
of the occludent ledge of the scutum. The internal surface of the
so-called spur is rounded and convex. The upper part of the tergum
is in main part formed by a great occludent ledge; but this, on its
lower side, is bordered by a narrow irregular slip, which, as shown
by the lines of growth, represents the whole of the ordinary valve,
excepting, of course, the spur already described. The occludent
ledges of both valves support some fine spines.
_Affinities._--Observing how extraordinarily the terga varied in _P.
dentatum_, and that the shells were identical in that and the present
species, it occurred to me at first that they might, perhaps, be both
extreme varieties of one form: but in the scutum of _P. crenatum_,
the invariably great development of the adductor plate,--the marked
manner in which its basal margin is reflexed,--the absence of a
tooth-like articular projection,--and again in the tergum of _P.
crenatum_, the invariably large size of the concave spur, without
any internal tooth, altogether convince me that the two species must
be considered as distinct. This species is allied to _P. grande_, in
the scutum of that species having an occludent ledge, though small,
and a great adductor plate. I have only further to remark, that the
figure of the opercular valves, given in Sowerby's Genera of Recent
and Fossil Shells, is so good, that there can not be the least doubt
about the present identification.
9. PYRGOMA MONTICULARIÆ. Pl. 13, fig. 5 _a_-5 _f_.
PYRGOMA (DARACIA) MONTICULARIÆ. _J. E. Gray_ (!). Zoological
Miscellany, p. 6, 1831.
_Shell of an irregular shape, with a roughened exterior border: orifice
minute, circular: scutum and tergum both much elongated, calcified
together without any suture, both furnished with a broad occludent
ledge._
_Hab._--Singapore; Mus. Brit., Cuming, and Stutchbury. Sometimes
associated with _Creusia spinulosa_.
_Appearance and Structure of Shell._--Shell dull white, very
irregular in outline, sometimes rounded, more often unequally
elongated, and frequently star-shaped,--the projections being
quite irregular. Whole shell nearly flat, but with the central
part saddle-backed, or formed into a more or less prominent ridge,
extending in the line of the longitudinal axis of the animal's
body: the circumferential portions of the shell not unfrequently
are a little recurved upwards. Orifice extremely minute, circular.
The outer lamina of shell, which is smooth, does not extend to the
circumference, and consequently a rather broad, nearly equal border,
which is rough, surrounds the whole shell. I have no doubt that,
when the shell was alive, this border was covered by a membrane,
which, in drying, has curled up and been lost, in the same manner
as the strictly analogous but narrow open seam between the basal
edges of the shell and the basis in some cirripedes (as in the last
two species of _Pyrgoma_) is protected. The roughened border can
sometimes be plainly seen to be formed of normal (Pl. 13, fig. 5
_e_) longitudinal septa having crenated edges, with shorter septa
between the longer ones; but more often the septa are so irregular,
and so much branched (5 _d_), that the whole resembles a mass of
moss. Why the outer lamina of the shell in this one species does not
nearly reach the circumference of the walls, I cannot conjecture. The
extremely irregular, depressed shape of the shell, with the minute
circular orifice, and the singular rough circumferential and often
slightly reflexed border, together give to this species so peculiar
an aspect, that until close examination I did not believe that it
was a cirripede. The extreme irregularity of shape depends in great
part upon the irregular growth of the Monticularia, in which it is
imbedded.
Internally (fig. 5 _b_) the walls are smooth, but they are perforated
by many quite irregular, small orifices, which have the appearance
of having been formed by some boring animal, but really serve, as
I believe, to admit threads of corium into certain irregular pores
which penetrate the shell. The sheath descends but a very short
distance from the orifice: it is closely attached to the walls,
and might easily be overlooked. The basis is deep, of an irregular
outline, like that of the shell, and is formed by a very thin shelly
layer. The largest specimen which I have seen, measuring from the
extreme projecting points, was .4 of an inch in diameter.
The _Scuta_ and _Terga_ (fig. 5 _f_) are calcified together without
any trace of a suture; together they form a bow with the two ends
curled rather abruptly inwards; they are both extremely narrow, but
furnished with an occludent ledge, twice or thrice as high as the
proper valves themselves. This occludent ledge, which is finely
hirsute, begins at about one third of the length of the scutum
from the rostral angle, and runs to near the basi-carinal angle of
the tergum. The scutum itself is curved, with a slip, along the
true occludent margin (best seen at the rostral end), lying in a
different plane from the rest of the valve, much in the same way as
in the scutum of _P. milleporæ_. The adductor ridge descends a very
little below the basal margin of the valve, and extends for nearly
its entire length: this adductor ridge makes the proper valve even
narrower than it at first appears. The _Tergum_ is extremely narrow,
forming merely a border to the occludent ledge; but it is not short,
being about two thirds of the length of the scutum. There is no trace
of a spur; indeed, the valve is rather narrower where the spur should
have stood, than it is at the basi-carinal end. The scuta and terga
are calcified together by their apices.
_Affinities._--Although this species, as above stated, differs so
remarkably in external appearance from the other species of the
genus, and, indeed, of the whole family, yet the shell in no one
essential point of structure materially differs from its congeners;
and if we compare the opercular valves with those of the three
last species, we shall be struck with their close, yet graduated,
affinity: in _P. milleporæ_ the scuta and terga tend to become
calcified together, and the rostro-occludent end of the scutum is
bent into a different plane from the rest of the valve. In _P.
dentatum_ we have an occludent ledge exactly as in the present
species; but in that species the adductor plate is less developed
than in _P. monticulariæ_; on the other hand, in _P. crenatum_, the
adductor plate is more developed than in _P. monticulariæ_. If in
_P. crenatum_ we were to remove the spur from the tergum (and it is
much less developed in _P. dentatum_; and in _P. milleporæ_ it is
entirely absent) this valve would be almost identical with that of
_P. monticulariæ_. Under these circumstances I consider it impossible
to separate the present species as a distinct genus.
_Species Dubiæ._
The _Daracia Linnæi_ of J. E. Gray (Annals of Philosophy, new series,
vol. 10, 1825), was published without description or figure.
The _Megatrema semicostata_ of G. B. Sowerby, junr. (Conch. Manual,
fig. 33, 1839), is not described, and is very indifferently figured
without the opercular valves, and therefore can never be recognised.
There is an admirable figure of a Pyrgoma, without any specific name,
in the great 'Description d'Egypte,' but from the want of some details,
I cannot positively recognise the species; I am inclined to believe
that it is the _P. dentatum_ of this work.
M. Chenu, in his grand Illustrations Conchyliologiques, has given most
beautiful figures of several species of Pyrgoma, and of Creusia, but
unfortunately, from the opercular valves not having been figured, I
find it impossible to recognise them. The new species are _Pyrgoma
stellata_ (on which I have appended a note under _P. conjugatum_), and
_P. spongiarum_ and _P. corymbosa_ of Valenciennes.
6. _Sub-Genus_--CREUSIA. Pl. 13, 14.
CREUSIA. _Leach._ Journal de Physique, tom. 85, July, 1817.
_Compartments four, furnished with radii; basis cup-formed: attached to
corals._
_Distribution_, imbedded in corals throughout the tropical seas.
Creusia is closely allied to Pyrgoma; and had not this genus already
been adopted by several authors, I should not, I think, myself have
formed it; though, no doubt, it harmonises well with some of the other
genera of the family, which are perhaps all too intimately related.
Creusia differs from Pyrgoma only in the shell being separated by
sutures, into four compartments, with well developed radii: in other
respects, such as in general habit, in the cup-formed imbedded basis
(not permeated by pores), in the opercular valves, in the characters
derived from the mouth and cirri, there are no generic differences.
This affinity is more particularly evident when Creusia is compared
with the first few species of Pyrgoma: indeed, for a short time, I was
inclined to consider _var._ 10 of Creusia as identical with _Pyrgoma
conjugatum_. With respect to the species of Creusia, I confess I have
been much perplexed in determining whether there be only one, or half
a dozen. The latter conclusion would almost certainly be arrived at if
only a few specimens were examined; and it might naturally be thought
that some of the species were extremely well marked; the difficulty
of drawing any line between varieties and species, begins only when
some hundreds of specimens, from various parts of the world, are
disarticulated, cleaned, and carefully examined. Creusia, in this
respect, offers a striking contrast with Pyrgoma, in which nearly all
the species are strongly characterised. The shell differs very little
in the several varieties or species of Creusia; and the most marked
difference, namely, whether the walls are permeated by irregular
pores or not, seems certainly quite variable. It is in the opercular
valves, which in other genera offer by far the most reliable character,
that we encounter the chief cause of perplexity; for the characters
thus derived, though at first appearing very distinct, blend into
each other, and are not accompanied by any well marked differences
in the shell. Only a few of my specimens have any habitat; but the
geographical range, as far as it does go, throws no light on the
question which forms to regard as species and which as varieties. As is
generally the case with cirripedes, the variations are local, so that
the greater number of specimens imbedded in the same coral resemble
each other. Under these circumstances I have thought it best, after
repeated examinations of a very large suite of specimens, to describe
separately each variety, without attaching any name to it; but I will
first make a few general remarks on the structure of the shell. If
I do not thus throw much light on the subject, I shall at least not
burden it with error. I believe that the species will be definitely
made out only by persons resident in the coral-bearing zones. I have
given copious illustrations of the opercular valves; for, if my view be
correct, this genus offers a curious and striking case of variation;
if, on the other hand, I am wrong, the drawings, I hope, will aid
others in coming to a more correct conclusion.
1. CREUSIA SPINULOSA. Pl. 13, fig. 6 _a_-6 _h_: Pl. 14, 6 _i_-6 _u_, 6
U.
CREUSIA SPINULOSA. _Leach_ (!). Encyclop. Brit. Suppl., vol. 3, Pl.
57, 1824.
CREUSIA SPINULEUSE. _De Blainville._ Dict. Sc. Nat., Pl. 116, fig. 6.
CREUSIA GREGARIA. _G. B. Sowerby_ (!). Genera of Recent and Fossil
Shells, No. 18, Sept. 1823.
---- GRANDIS. _Chenu._ Illust. Conch. Tab. 1, fig. 2, sed non fig. 2
_a_ and _b_.
_Hab._--Philippine Archipelago, China, Singapore, Java, Red Sea, West
Indies; imbedded in various corals; Mus. Brit., Cuming, Stutchbury,
Dunker, &c.
_General Appearance._--The shell is oval, generally flat, sometimes
conical, with narrow and approximate ridges radiating from the
orifice (fig. 6 _a_). The ridges, however, are sometimes distant from
each other, and considerably prominent, projecting round the basal
border. The orifice is either neatly diamond-shaped or oval. The four
compartments are quite distinct; the radii are generally white, of
considerable width, and with their summits not oblique. The colour
is either white, or pale pinkish-purple; but in _var._ 11, bright
pink. Even in the white specimens, when well preserved, the sheath is
generally, but not always, either pale or dark purple. The largest
specimen which I have seen, from the West Indies, was above half an
inch in diameter; but from .3 to .4 of an inch is the more usual full
size. I believe that the size, as well as the great variability of
the present species, is partly determined by the rate of growth of
the various zoophytes in which the specimens are imbedded, for the
shell has to keep on nearly a level with the surface of the coral.
_Structure of Shell and Basis._--The walls are internally ribbed;
the ribs being usually prominent, sometimes to such a degree as
to deserve to be called plates. The outer lamina is of variable
thickness, and the prominence of the internal ribs appears in
considerable part to depend on the extent to which the outer lamina
has been thickened from within. In many specimens, instead of the
interspaces between these internal ribs or longitudinal septa
being solidly filled up, separate and successive laminæ have been
deposited, by which the shell is rendered cancellated or porose; but
the pores are very irregular; and sometimes they form two or three
irregular rows one behind the other: this structure seems eminently
variable. The edges of the radii are formed by crenated, and
occasionally branched, septa. That part of the alæ, which is added
during the diametric growth of the shell, is very thin. The lower
edge of the sheath seems always to be free. The shelly layer, forming
the basis, which is deeply cup-formed, is thin, more or less finely
furrowed in radiating lines, and not permeated by pores.
The opercular valves will be best described under the following
eleven varieties.
_Var._ (1), Pl. 13, fig. 6 _a_, 6 _b_, 6 _c_, 6 _d_.--_Hab._ Java,
and probably several other districts.--I will first describe a
typical sub-variety, by which I mean a sub-variety not presenting
any extreme character. The scutum is of a sub-triangular shape, with
the basi-tergal corner much rounded off (6 _d_), and generally but
not always hollowed out in a rather remarkable manner. The adductor
ridge is considerably prominent, and extends high up, parallel to
the articular ridge, which latter is rounded and prominent, but
to a variable degree. Near the rostral angle there is sometimes a
small tooth, or only a trace of one, depending beneath the basal
margin; this tooth we shall hereafter see much more developed. The
_Tergum_ is about two thirds of the width of the scutum. It is
often slightly beaked, but this is more conspicuous in some of the
following varieties. The spur is about half the width of the valve,
and its basal end is truncated, and nearly parallel to the basal
margin of the valve, but the truncated form passes insensibly into
a rounded outline. The shell in this variety is generally thick and
is not permeated by pores; the orifice is diamond-shaped. But in
another sub-variety the walls of the shell are always, or nearly
always, permeated by pores, and the tergum is very much narrower,
with the spur sharper, so that at first I concluded that these two
sub-varieties were specifically distinct: we shall, however, soon
see in _var._ 2 and in _var._ 4, that no confidence whatever can be
placed in the exact breadth of the tergum, or in the porosity of the
walls; hence I have been driven to consider the two varieties just
mentioned as merely sub-varieties.
_Var._ (2), fig. 6 _e_, 6 _f_, 6 _g_.--_Hab._ China; Red Sea.--The
shell is almost invariably permeated by pores, sometimes arranged
in two or three very irregular rows. In some specimens the scutum
exactly resembles that in _var._ 1, but with the tooth near the
rostral angle often rather larger: in other specimens the scutum is
much more elongated transversely (fig. 6 _e_), with the adductor
ridge more medial, and the basal margin not at all hollowed out
at the basi-tergal corner of the valve. The tergum, here, is the
remarkable feature, being sometimes excessively narrow, with a long
sharp spur, which often, but not always, terminates in a needle-like
point. In other specimens, from the same coral and certainly
belonging to this same variety, the valve is not so narrow (6 _g_),
and the spur not so pointed; consequently (as in several analogous
cases in other cirripedes) it is impossible to draw any line of
distinction between the specimens with the narrow and broad terga.
_Var._ (3), fig. 6 _h_ [_Creusia gregaria of G. B. Sowerby!_]--_Hab._
Unknown.--The scutum presents here exactly the same considerable
range of variation as in _var._ 2. The tergum is broad, as in _var._
1, but the spur is rounded, and from not being placed so immediately
close to the basi-scutal angle of the valve, gives to it a rather
different aspect. The breadth of the spur varies; an extreme variety
is given in fig. 6 _h_.
_Var._ (4), Pl. 14, fig. 6 _i_, 6 _k_, 6 _l_.--_Hab._ Philippine
Archipelago; West Indies.--The _scutum_ here presents the same
sub-varieties as heretofore, excepting that I have not seen any so
much elongated transversely. The shell is covered either with slight,
closely approximate ribs, as in the foregoing varieties, or with more
distant and more prominent ribs. In specimens taken out of the same
branch of coral the walls were either porose or solid. Sometimes the
sheath is bright purplish-pink. It is the _tergum_, again, which
presents a remarkable range of difference; for the longitudinal
depression or furrow which in the former varieties was quite open,
here has its edges more or less folded inwards, and is sometimes
quite closed. This same variation has been commonly observed in many
species of Balanus, in which it appears to be dependent on the age
of the individual; but this does not appear to be the case in the
present genus. As a consequence of the greater or less folding in of
the two sides of the furrow, the spur is rendered more or less narrow
and pointed, and thus becomes removed to a greater or less distance
from the basi-scutal angle of the valve. Further, as a consequence of
this folding in, the internal surface of the valve along the line of
the external furrow, is raised into a longitudinal ridge. The length
of the spur varies considerably. In some very young individuals, the
basal margin descends lower on the scutal than on the carinal side
of the spur. In one set of specimens (fig. 6 _l_), a plate extended
from the carinal margin to near the central longitudinal ridge
just mentioned: a similar structure was described under _Pyrgoma
cancellatum_.
_Var._ (5), Pl. 14, fig. 6 _m_.--_Hab._ Unknown.--We have seen in
_vars._ 2 and 3 that the scutum varies considerably in shape: here
it is unusually narrow, with the adductor ridge almost touching the
articular ridge. There is no little tooth near the rostral angle, and
the basi-scutal corner is not hollowed out. The tergum also varies;
in some individuals it is truncated and like that figured of _var._
1, but rather more rounded; in other specimens (from the same branch
of coral) the basal margin so blends into the spur that the latter
can hardly be discriminated (fig. 6 _m_); in other respects the
outline resembles pretty closely that of one of the _sub-vars._ (fig.
6 _f_) of _var._ 2. The shell is not porose; it is thick, with strong
internal ribs, and resembles that of _var._ 1; but it is of a pale
purplish colour.
_Var._ (6), _an. spec._? fig. 6 _n_-6 _q_.--_Hab._ Philippine
Archipelago.--This is a very remarkable variety; we have, imbedded
in the same coral, and with shells absolutely identical, specimens
with the scutum having three distinct but graduated forms. Firstly,
a scutum transversely elongated, in all external respects like some
of the varieties mentioned under _vars._ 2 and 3, with no rostral
tooth, and not hollowed out at the basi-tergal corner, but with the
adductor ridge more prominent. Secondly, a scutum of the same general
shape, but with the adductor ridge so much developed (fig. 6 _n_) as
to descend slightly beneath the basal margin, and to be seen when
the valve is viewed externally; there is a very slight tooth near
the rostral angle (as in some former sub-varieties), and which can
be here rather more clearly seen than hitherto, to be formed by the
adductor ridge (closely united to the external surface of the valve)
extending thus far, and being here produced a little downwards.
Thirdly (6 _p_), we have the adductor ridge immensely developed,
descending far below the basal margin of the ordinary valve; and
the basal margin at the basi-tergal corner is angularly and deeply
hollowed out. The appearance of the valve is widely different from
that in the first sub-variety, yet it is impossible to separate the
first and second sub-varieties, and it is almost equally certain that
the third sub-variety is only an exaggeration of the second. The
lower edge of the adductor ridge, in the third sub-variety, varies
a little in outline; it is deeply sinuous, and is produced at the
rostral angle into a point, of which we have heretofore seen only a
feeble representation. It would appear as if the great development of
the adductor plate had caused the exterior ordinary surface of the
valve to shrink or be less developed. There is a striking resemblance
in the structure here described with that in _Pyrgoma cancellatum_
and _conjugatum_. The terga belonging to the above scuta, also,
vary; the spur being sometimes square (6 _o_), and sometimes bluntly
pointed: when the spur runs in the same exact line (6 _q_) with the
scutal margin of the valve, a peculiar aspect is given to it, but
this is by no means always the case. Both opercular valves are often
partially coloured pinkish-purple. The shell is not porose; it is
thin, with remarkably prominent internal plates; it is apparently
always of small size, which I attribute to this variety inhabiting
a hard thin plate-like coral. The sheath is bright pinkish-purple,
of which we have had instances in some of the other varieties; and
the shell itself is sometimes pinkish. Taking the scutum of the
first sub-variety, together with the commonest accompanying variety
of tergum, I find it quite impossible to assign to it a specific
character; if, on the other hand, we consider the scutum of the
third sub-variety by itself, nothing can appear more distinct; but
I must repeat, there can be hardly a shadow of doubt that the three
sub-varieties of scutum here described, graduate into each other, and
are specifically identical.
_Var._ (7), fig. 6 _r_.--_Hab._ Probably Philippine Archipelago,
associated with _Balanus quadrivittatus_.--There can be hardly any
question of this being specifically identical with the last variety.
It inhabits a different coral. All the specimens were of small size.
The walls are not so thin, and the internal ribs not so prominent as
in _var._ 6. The sheath is either white or dull purple; I can, in
short, point out no difference in the shell from the typical _var._
1. The scutum is not so much elongated transversely as in _var._
6, and the basi-tergal corner is more cut off,--in which respect
it resembles the common varieties. The adductor ridge is largely
developed, so as to be just visible when the valve is viewed from the
outside, in a degree between the first and second sub-varieties of
_var._ 6: but the most singular character is the larger development
of the tooth near the rostral angle, and this was the case in the
same degree in all the specimens which I examined. The tergum
resembles that rather unusual sub-variety of _var._ 6 (fig. 6 _q_),
which has the scutal margin and the one side of the spur forming a
straight line. It appears to me that it would be absurd to consider
these slight differences, in parts unquestionably subject to much
variation, as specific, when we are almost forced to admit that the
much greater differences in the three sub-varieties of _var._ 6, are
not of specific value.
_Var._ (8).--_Hab._ Unknown, Mus. Cuming.--I have seen only a single
specimen of this, and refer to it on account of _var._ 11. The shell
is rather steeply conical, with distant and prominent ribs; the radii
are narrow; the walls are not permeated by pores; the colour is pale
purple. Altogether its external appearance is very different from
that of the foregoing varieties; but the scuta are identical with
those of _var._ 1, excepting that the rostral tooth is rather larger,
being nearly as large as in the last, _var._ 7. The tergum precisely
resembles that in some specimens of _var._ 2. Hence this variety
differs from the first two varieties only in the shade of colour, the
external shape, and the greater prominence of the external radiating
ribs of its shell. All these characters are variable in the several
foregoing varieties, and they have been found, as yet, insufficient
to discriminate species in any genus of sessile Cirripedes.
_Varieties With the Scuta and Terga Calcified Together._
_Var._ (9), Pl. 14, fig. 6 _s_ [_C. spinulosa_, of Leach (!)]--_Hab._
Unknown.--The shell is undistinguishable by a single character from
many specimens of the first, third, and fourth varieties; it is not
permeated by pores. The scutum and tergum, with the exception of the
one striking difference of their being calcified together without
any trace of a suture, are identical with those of _var._ 3, as may
be seen by comparing the figures 6 _h_ and 6 _s_. Hence to separate
this form specifically from _var._ 3, we should have to rely solely
on the calcification or union of the scuta and terga; but we have
seen this is a point which is variable in _Elminius Kingii_, _Pyrgoma
milleporæ_, and in some species of Balanus. The serial affinities,
moreover, in Pyrgoma, clearly show that this is a character of no
great importance. I must add that in several specimens of several of
the varieties, the scuta and terga were so closely joined, that until
careful examination, I was unable to detect the suture separating
them; such being the case it must be quite unimportant for any
functional purpose, whether or not the valves are calcified together.
I feel, consequently, hardly any doubt that I have acted right in
treating the present form as a mere variety.
_Var._ (10) _an. spec._? fig. 6 _t_.--_Hab._ Unknown.--This variety
bears nearly the same relation to _var._ 6, as the last variety did
to _var._ 3. The shell is rather stronger than in _var._ 6, with the
internal ribs not so prominent; and except in being tinted pale dull
purple, it differs in no respect from the shell of _var._ 1. If we
imagine the scutum and tergum in the third sub-variety of _var._ 6
(6 _p_, 6 _q_), in which the adductor ridge descends far beneath the
true basal margin of the valve, to be calcified together, without any
suture, we shall produce almost the identical valves of the present
variety. The scutum, however, here is not quite so much elongated
transversely, and the occludent margin is spinose and is furnished
with large teeth; these two characters give the valve a somewhat
different aspect, and hence I am more doubtful than in the foregoing
case, whether this form may not be specifically distinct. I must,
however, state that in _Tetraclita porosa_, I ascertained that the
teeth on the occludent margin of the scuta were even more variable
than here is supposed to be the case; and as for the shape of the
valve we have seen what wonderful variation there is in _var._ 6. The
tergum in this variety is about intermediate between the two common
forms, in the sub-varieties of _var._ 6. As for the calcification of
the two valves together, we have seen, under the last variety, how
little important a character it is.
_Var._ (11) fig. 6 U, 6 _u_ [_Creusia grandis_, of Chenu, Tab. 1,
fig. 2, but not fig. 2 _a_ and _b_].--_Hab._ Singapore, associated
with _Pyrgoma monticulariæ_.--This variety is very closely related
to the last. The shell, however, has a very peculiar aspect, which
made me for some time think it must be specifically distinct. It is
of a much brighter pink than in any of the foregoing varieties; the
surface is marked with very prominent, distant ribs, and the radii
are narrow, in which latter points, together with the tint (though
here brighter), this variety cannot be distinguished from _var._ 7.
The shell, however, is permeated by several rows of pores, in which
respect it resembles the shell in _var._ 2, and some specimens of
_vars._ 3 and 4. In the opercular valves there is a close general
resemblance with those of the last _var._ 10; the tooth, however,
near the rostral angle, is not so prominent; and in the tergum, the
spur is more truncated, shorter and broader than in _var._ 10, and
closely resembles that in _var._ 1 and 2. But I cannot consider any
of the points here specified of much weight.
The foregoing descriptions show how singularly the affinities of the
several varieties interlock in the most complicated manner. Hereafter
some one may, perhaps, succeed in grouping several of these forms as
species; but I am sure he ought not to attempt it without possessing
a very large suite of specimens, or without the great advantage of
comparing some two or three of the forms, fresh in their native site.
_Species dubiæ._--Under Pyrgoma, I have stated that though Chenu,
in his 'Illust. Conch.,' has given beautiful external figures of
the shells, imbedded in the coral, yet from the want of details on
the opercular valves and on the structure of the shell, I cannot
recognise his species. So it, likewise, is with Creusia. Chenu gives
the following new species; _C. radiata_, _multistriata_, _decorata_,
and _striata_. The _C. madreporarum_, I suspect to be the _Pyrgoma
milleporæ_ of this work, as there stated. The _C. grandis_ no doubt
is the _P. grande_ of this work, the _Nobia grandis_ of Sowerby. The
name _Creusia Childreni_ is given by Dr. Gray, without description or
figure, in the 'Annals of Philosophy,' vol. 10, new series, 1825.
7. _Genus_--CHELONOBIA. Pl. 14: Pl. 15, fig. 1.
CHELONOBIA. _Leach._ Journal de Physique, tom. 85, (1817).
CORONULA. _Lamarck._ Animaux sans Vertèbres, 1818.
---- _Ranzani._ Memoire di Storia Naturale, 1820.
---- _De Blainville._ Dict. des Sciences Naturelles.
ASTROLEPAS. _J. E. Gray._ (Klein) Annals of Philosophy, (new series),
vol. 10, (1825).
_Compartments extremely thick, six; but one of them, the rostrum,
internally is composed of three rudimentary compartments, united
together: basis membranous: scuta narrow, united to the terga by a
horny articular ridge._
_Distribution_, throughout the tropical, and warmer temperate seas of
the whole world; attached to turtles, crustacea, or smooth gasteropod
molluscs.
This is a distinct and well-defined genus. Several authors have
confounded it with Coronula, but this has been owing to an entire
misapprehension of the structure of the shell in the two genera. In
Coronula, the parietes are very thin, and are so deeply folded as to
appear like rays or septa connecting the outside of the shell (the
expanded ends of the folds) and the internal surface of the shell, but
the open spaces between the folds are occupied by the epidermis of the
Whale, and are external to the cirripede. In Chelonobia, the parietes
are remarkably thick; hence the plates or septa, connecting the outer
and inner lamina, are of unusual length; and the spaces between them,
though of course internal with regard to the cirripede and occupied by
the ovaria, have been compared to the spaces external to and between
the folded walls of Coronula. There is but little special affinity
between these genera; and I regret that they have come to be placed one
after the other in this work: but the elongated opercular valves,--the
thick and double opercular membrane,--the weak depressor muscles,--and
the peculiar manner in which the scutum is articulated by the aid of
a horny projection to the tergum, may indicate some real but slight
affinity to Coronula; the many points of difference, however, in the
structure of the shell and of the opercular valves, and especially in
the cementing apparatus of the basal membrane, and in the branchiæ,
all prove that the genera are very distinct. The singular structure of
the rostrum, which, in fact, consists of three compartments externally
blended together, and which three correspond in all essential respects
to the rostrum and two rostro-lateral compartments in the Chthamalinæ,
offers a very striking point of identity with that sub-family; but
neither in the mouth, cirri, or other part, can I detect any other
evidence of this relationship. Having so far discussed the affinities
of the genus, I may add, that the three species, though decidedly
distinct, are closely and nearly equally related to each other.
_General Appearance._--The shell is generally depressed, and broadly
oval or almost circular; in _C. testudinaria_ and _caretta_, it
has a massive appearance: the surface is generally smooth, or,
when disintegrated, finely striated: the colour is white. The six
compartments do not differ much in size: the rostrum is rather larger
than the carina, and the lateral compartments, than the carino-lateral
compartments. It is remarkable that in _C. caretta_ (Pl. 14, fig.
2), even in specimens which have not grown crowded together, the
compartments are almost invariably placed rather unsymmetrically,
the rostrum and carina not exactly facing each other.[121] The shell,
though so thick and massive, yields easily along the lines of suture.
The radii are moderately wide, or narrow, or not at all developed,
being represented by mere sutures: in this latter case, in _C.
caretta_, the orifice of the shell is enlarged, in the same manner as
we have seen in some species of Balanus and Tetraclita, by the gradual
wearing away of the upper part of the shell. In _C. testudinaria_,
the radii have a singular notched structure (fig. 1 _a_), and the
whole shell a star-like appearance. The orifice is not filled up by
the elongated opercular valves,--a considerable extent of opercular
membrane being visible on the two sides. The largest specimen which
I have seen, namely, of _C. testudinaria_, was nearly two and a half
inches in its longer diameter.
[121] In Mr. Stutchbury's collection there is a specimen of _C.
testudinaria_ in which there are only five compartments, one of the
lateral compartments having been aborted; of this I have seen no
other instance in any genus.
_Structure of the Parietes._--The parietes are of unequalled thickness,
especially in the first two species of the genus. From the outer lamina
(see Pl. 14, fig. 4, and the section in Pl. 15, fig. 1), numerous
vertical plates extend inwards, alternately to a less or greater
distance, some of them reaching to the inner lamina: these answer to
the longitudinal parietal septa in other genera, and the elongated
cavities between them (which occur in _C. testudinaria_ and _patula_)
answer to the parietal tubes or pores. The radiating plates or septa
have their sides finely channelled, and their basal edges generally
slightly sinuous and always finely toothed. The interspaces between
the plates in the uppermost part of the shell are filled up solidly,
and, in _C. caretta_, even down to near the basis: in this latter
species, the plates are irregular and much broken up, so as in parts
to consist of little, separate, flattened points. In _C. patula_, the
inner lamina of the parietes (_b_, in fig. 4. Pl. 14) can be best
made out to be distinct from the sheath (_e c e_ in fig. 4, and _c′_
in the section of _C. testudinaria_, fig. 1, Pl. 15). The sheath in
this genus descends in a very remarkable manner to the basal membrane,
and has its basal edge toothed like the basal edges of the radiating
septa. The inner lamina itself does not descend to the basal membrane.
In _C. testudinaria_, the inner lamina is of great thickness; but in
the section, (fig. 1) owing to its having been taken high up, the
inner lamina, (_b_), is not distinct from the shelly matter deposited
between the septa. In _C. caretta_, the line of separation between the
inner and outer laminæ can in no part be distinguished, owing to the
interspaces between the septa having been solidly filled up, close down
to the basis.
_Sheath._--The layer of shell surrounding the internal cavity (_e c
e_, fig. 4, Pl. 14), and extending down to the basal membrane, I have
called the sheath, owing to its being distinctly continuous with the
innermost layer in the upper part of the shell, to which the opercular
membrane is attached: this can be best seen by examining the alæ in
the separated compartments of _C. patula_. The sheath is not only
remarkable from thus descending to the basal membrane, but in _C.
testudinaria_ and _patula_ from its lower edge being perforated by
arched channels (under _c_, in fig. 4), allowing thick ribbons of
corium to reach the interspaces between the radiating septa. There
is one central arch or channel in the middle of the lower part of
the sheath of each compartment, and one on each line of suture, the
sheath being a little hollowed out on both sides of the sutures. As the
rostrum, as far as its internal structure is concerned, consists of
three compartments, we have altogether in the shell eight compartments
and eight sutures, and consequently altogether sixteen arches through
the lower part of the sheath, allowing sixteen thick ribbons of corium
to penetrate the parietes, and thus likewise reach the radii. There is,
however, sometimes a little variation in the number of these arched
channels. The upper part of the sheath is transversely marked by
zones of growth, to the lower one of which the opercular membrane is
attached. The line of attachment is not low down the sheath.
_Radii and Alæ._--The radii, when the compartments are disarticulated,
present a remarkable structure, from appearing to consist of a distinct
inner and outer portion. The radius normally consists of an inner and
outer lamina, united by septa parallel to the basis; but here the inner
portion is formed by a central ridge (_a a_, fig. 5, Pl. 14), sending
off on both sides little septa, often sub-branched; it is of nearly
uniform width; and there is no distinct inner lamina. The outer portion
(_b_), which often equals or exceeds in thickness the inner portion,
is, in fact, the normal outer lamina, developed to an unparalleled
degree. In most Cirripedes the edge of the radius is received in a
slight furrow in the opposed compartment, the lid of which furrow is
narrow, and matches the outer lamina of the radius; here the lid of the
recipient furrow is very broad, and resembles the outer lamina of the
radius in all its characters. In order to allow of growth between the
_thick_ opposed edges of the outer lamina of the radius and the lid of
the recipient furrow, the two surfaces are finely dentated (look in
fig. 5, under the pits, marked _b_), almost like the crown of a molar
tooth; thus allowing films of corium to enter. The structure here
described is common, in a greater or less degree, to all three species,
but is best seen in _C. testudinaria_. In this species, moreover, (fig.
5, _b_), the outer lamina, instead of being smooth and of either equal
or gradually increasing thickness from top to bottom, is generally, but
not always, (fig. 1 _a_), deeply pitted or notched in transverse lines,
the outer lamina being thus rendered alternately thicker and thinner,
and so formed into transverse ridges and valleys. Hence the lines of
suture become toothed, the points of the teeth facing each other, and
not interlocking. In the transverse section, fig. 1, Pl. 15, of the
same species, taken high up across the shell, (_f_) is the pitted outer
lamina, and (_e_) the inner portion of the radius. Although the radii
are thus specially added to in thickness, they are not so thick as the
very thick walls, and hence the lines of suture form furrows more or
less deep. The _alæ_ are of moderate thickness, and have their sutural
edges crenated by fine transverse septa.
_Rostrum._--I have already alluded to the peculiar compounded structure
of this compartment, unlike anything we have as yet seen.[122] The
thin outer lamina is quite continuous, and shows no trace of the
triple nature of the compartment; as may be seen by comparing the
drawing (Pl. 14, fig. 1 _a_) of the shell of _C. testudinaria_,
with the transverse section (Pl. 15, fig. 1) of the same species:
in this latter figure, _a_ is the outer lamina, and B A B the three
compartments of the rostrum. But when the outer lamina is worn away,
as is always the case with the upper part of the walls in _C. caretta_
(Pl. 14, fig. 2), the two fissures separating the three compartments
of the compounded rostrum, are plainly exhibited on the outside of
the upper part of the shell. On the internal surface, the sutures
separating the three compartments are always open, except in the upper
part of the sheath, above the attachment of the opercular membrane,
where they are partially obliterated by a thin continuous layer of
shell. That these three portions of the rostrum are in their essential
nature compartments, is well shown in _C. patula_ and _testudinaria_,
by the sheath or inner lamina having loopholes or channels (such as
before described) in the middle of each, and on each line of suture.
From the number of these channels, seven altogether, (the two between
the compound rostrum and lateral compartments being counted,) the
sheath of the compound rostrum is reduced to mere flattened pillars
between the several channels. By slight violence, the rostrum breaks
into the three portions; and the sutures between them are found to be
marked on both sides by sinuous, slight, calcareous ridges, those on
opposite sides locking into each; these represent the septa on the
edges of the radii and alæ and their recipient furrows, in the ordinary
compartments. The outline of the middle compartment of the three (best
seen in section Pl. 15, fig. 1, A), much resembles that of the carina;
in fact, if we suppose the growth of the two alæ of the carina to have
been arrested, no essential difference can be pointed out: in this
rudimentary compartment, therefore, we have a rostrum, characterised as
in the sub-family of the Chthamalinæ. In the two little rostro-lateral
compartments, moreover, (B B), a slight swelling on the side opposite
to the large existing radius, shows that if the development of these
compartments had not been prevented, each, probably, would have
had, exactly as in the Chthamalinæ, a radius on both sides. In the
introduction to the Balanidæ, I have argued, from the structure here
described and from some other facts, that in the Balaninæ the rostrum
is composed of the three anterior compartments, which we see in the
Chthamalinæ, indissolubly united together; hence in Chelonobia the
middle one of the three partially-blended compartments is properly the
rostrum, and those on the sides the rostro-lateral compartments.
[122] My attention was first called to this peculiar structure of
the rostrum by Mr. Stutchbury.
_Basis._--The basal membrane extends under the thick walls to the
outside. I was not able to make out the whole cementing apparatus.
The main trunk is remarkable from its small diameter, (_f f_ in fig.
2, Pl. 28), and from the distance at which the cement-glands stand
apart. Each gland gives rise to a pair of cement-ducts, which tend to
run in parallel groups; these ducts repeatedly bifurcate, occasionally
inosculate, and decrease in diameter; they debouch and allow the cement
to escape at numerous points, placed at irregular distances, on the
edges of each new slip of the basal membrane.
_Opercular Valves._--These are elongated; they do not fill the orifice
of the shell; they are attached by a strong opercular membrane a little
way down the sheath. The opercular membrane is generally double, for
the last-formed membrane is not immediately moulted as soon as a new
one is formed, as generally happens in Balanus. Externally, the valves
are marked by rather rugged, broad zones of growth. The _Scuta_ are
elongated in the line of the orifice of the sack; the occludent margin
(Pl. 14, fig. 1 _b_) is much inflected, and generally sinuous; along
this inflected portion a distinct square-edged ridge runs, which widens
from the apex downwards. The depression for the adductor muscle is
very deep: there are no pits or crests for the other muscles. When the
scutum is thoroughly cleaned and all the membrane removed by caustic
potash, the tergal margin is marked by a slight articular ridge and
furrow. This articular ridge is very remarkable from supporting a
prominent, flattened crest (fig. 1 _b_), composed of hard, yellow,
horny membrane, which overlaps the inner surface of the tergum,
and exactly corresponds, in shape and purpose, with the calcareous
articular ridge, when best developed in other genera. Beneath the basal
and generally slightly sinuous exterior margin of the valve, that is,
the margin to which the opercular membrane is attached, a slight
ledge depends (fig. 1 _d_), which narrows off towards the rostral end
of the valve. I should have thought that this had been a depending
adductor ridge, as in several species of Pyrgoma, had there not been
a nearly similar ledge beneath the middle part of the basal margin of
the tergum. The _Tergum_ is mitre-formed, with the summit more or less
truncated, and with the carinal margin generally more sloping than the
scutal margin: near the carinal margin there is a slight furrow (fig. 1
_d_), sometimes difficult to be distinguished, with the lines of growth
curving down to it on each side, and consequently with a very slight,
but variable, corresponding projection on the basal margin. This furrow
and slight projection, there can be hardly any doubt, represent the
spur, though here placed close to the carinal instead of to the scutal
margin, as in other genera. The tergum has a small articular ridge,
against which the overlapping horny articular ridge of the scutum
abuts. There are no crests for the tergal depressor muscles. Altogether
the scuta and terga are very peculiar. A portion dissolved in acid
exhibits no tubuli. On the opercular membrane there are no hairs. The
rostral depressor muscles of the scuta are singularly feeble, each
consisting of only one or two, or sometimes three or four, very thin
ribbon-like fasciæ; the lateral depressores of the scutum are twice as
strong as the rostral depressores; and the tergal depressores a little
stronger than the lateral depressores. All these muscles are plainly
marked with transverse striæ.
_Mouth._--The labrum is not in the least bullate, which character, as
well as some others that follow, I specify on account of the apparent
affinity of Chelonobia to the Chthamalinæ, as indicated by its rostrum.
The crest of the labrum is notched, and on each side of the notch there
is a long row of teeth, which, however, are sometimes worn away. The
mandibles have five main teeth; all excepting the first being laterally
double. The maxillæ are not notched. The outer maxillæ are bilobed on
their inner sides.
_Cirri._--The four posterior pairs, in proportion to the size of the
animal, are much elongated, and are remarkable from the number of their
short segments. The rami of the first cirrus are a little unequal in
length. The second cirrus is moderately short, with its segments
rather broad and protuberant, and thickly clothed with spines. The
third cirrus is of unusual length, being but little shorter than the
fourth pair; its segments, however, are broad, and are thickly clothed
with spines, as are the two segments of its pedicel: hence there is no
real approach to that important character of the Chthamalinæ, namely,
the similarity of the third with the three posterior pairs of cirri.
The numerous segments of the fourth, fifth, and sixth pairs of cirri
each support only two pairs of main spines; between each of these pairs
there is a little tuft of fine intermediate spines; the upper of the
two tufts on each segment is the longest. In a specimen of _C. patula_,
in which there were fifteen segments in one ramus of the second cirrus,
there were fifty segments in either ramus of the sixth cirrus. At the
exterior bases of the pedicels of some of the anterior cirri, there are
large tufts of finely plumose, delicate hairs.
_Branchiæ._--These are of large size: they consist of a single fold,
much plicated and sub-plicated.
_Ovaria._--The ovarian tubes run into the parietes, and fill up the
interspaces between the radiating septa.
_Range, &c._--The three species seem to range together, over the
tropical and warmer temperate seas of the whole world. _C. patula_
and _testudinaria_ are found in the Mediterranean, and the former at
Charleston, in the United States; I have not heard of specimens from
any point further north. _C. testudinaria_ and _caretta_ live attached
to turtles; whilst _C. patula_ always adheres to crustacea, to large
and smooth gasteropod mollusca, and, I believe, sometimes to ships'
bottoms. I have not heard of the discovery of any fossil species.
_Attachment._--_Chelonobia patula_ leaves no impression on the crabs
and shells to which it is attached. I have seen only a few specimens
of _C. testudinaria_ attached, and the carapaces of the turtle were
not at all, or scarcely at all, indented by them. The case is very
different with _C. caretta_, in which the shell, even of young
specimens, is always, as far as I have seen, imbedded to some depth,
and occasionally to a very great depth in the tortoise-shell. From
the extreme hardness of the tortoise-shell, when dry, the imbedment
appears more wonderful even than it really is. The younger shells have
the appearance of having grown from within the carapace, and then of
having burst through it, almost like little volcanos. I have seen
only one very young shell (1/10th of an inch in external diameter)
attached, and here there was nothing to countenance an idea which at
one time occurred to me, namely, that the larva perhaps fixed itself in
some little crack or cavity in the carapace, and there underwent its
metamorphosis. I believe that the imbedment is effected simply by the
sharp, growing, basal edges of the walls of the shell indenting the
surface, and finally rupturing the outer laminæ of the tortoise-shell,
through that same force by which the tender radicle of a plant
penetrates very hard ground. As soon as the surface was once ruptured,
the shell of the Chelonobia, growing outwards and downwards, would
easily, like a wedge, turn up the laminæ of the tortoise-shell; and
their ragged ends would surround the Chelonobia, as is seen actually to
be the case. In the genus Coronula and its allies, which are attached
to Cetaceans, we shall presently see, that the epidermis immediately
under the downward growing shell, and apparently in consequence of the
pressure thus exerted, is thinner than in the surrounding parts. In
two specimens of _Chelonobia caretta_, imbedded much more deeply than
usual--in one of which half the basal edge of the shell had fairly cut
through the carapace, and in the other was on the point of effecting
this--the tortoise-shell manifestly thinned out towards the line of
yielding; now this, I suppose, must be attributed either to absorption,
or to the living tortoise-shell being actually stretched till rendered
transparent and ready to burst or until bursted. On the latter view
of the tortoise-shell having been stretched, we must further suppose
that the pressure has prevented fresh layers of tortoise-shell being
deposited under the old and yielding laminæ. In one of the above two
specimens, the walls of the Chelonobia were deeply folded, nevertheless
the laminæ of the tortoise-shell followed every curvature, showing
that, though now so rigid, during the slow imbedment of the Cirripede
it must have been sufficiently pliant. A shell attached, as these two
specimens were, could never be removed, and, whether dead or alive,
would remain for ever imbedded in the tortoise-shell. Dr. R. Ball,
of Dublin, informs me, that he has seen specimens in which the shell
of the cirripede not only had penetrated the carapace, but likewise
the underlying bone, and had even entered some way into the body of
the turtle: it is well known that the tusk of a boar or the horn of a
ruminant, when curving in abnormally, will sometimes penetrate deeply
into the bones of the face or head; and this, I believe, is effected,
not by the fracture of the bone, but by the absorption of the point
pressed on: I conceive a similar process must have taken place in the
curious specimens examined by Dr. Ball.
1. CHELONOBIA TESTUDINARIA. Pl. 14, fig. 1 _a_-1 _d_, fig. 5; Pl. 15,
fig. 1.
LEPAS TESTUDINARIA. _Linn._ Syst. Naturæ, 1767.
VERRUCA TESTUDINARIA. _Ellis._ Phil. Transact., vol. 50, 1758, Pl.
34, fig. 12.
BALANUS POLYTHALAMIUS. _Bock._ Naturforscher, Stuch. 12, 1778, fig. 9.
LEPAS TESTUDINARIA. _Poli._ Testacea Utriusque Siciliæ (1795) Tab.
5, fig. 9-11.
CORONULA TESTUDINARIA. _Ranzani._ Memoire di Storia Naturale, Decade
1, (1820).
---- ---- _De Blainville._ Dict. des Sciences Nat., (1824), Tab.
117, fig. 2.
CHELONOBIA SAVIGNII (?) _Leach._ Encyclop. Brit. Suppl., vol. 3, 1824.
ASTROLEPAS ROTUNDARIUS. _J. E. Gray._ Annals of Philosoph. (new
series), vol. 10, 1825.
_Shell conical, depressed, heavy: radii rather narrow, depressed,
generally notched on both sides._
_Hab._--Mediterranean; west coast of Africa; north-east coast of
Australia; Low Archipelago, Pacific Ocean; west coast of Mexico, 23°
N. Attached to turtles.
It is impossible to feel sure which of the three species of the genus
Linnæus had in view, when describing his _Lepas testudinaria_; but
as Spengler has well discriminated the following species under the
specific name of _caretta_,[123] and Ranzani the third species under
_patula_, the present name may, without question, be retained for the
following species. In several respects this species is intermediate
between _C. caretta_ and _patula_, but it can most conveniently be
described first.
[123] I am indebted to Dr. J. E. Gray for having guided me to this
identification.
_General Appearance._--Shell strong, globulo-conical, depressed;
outline broadly oval; surface smooth, generally well preserved, but
when disintegrated, upper part finely striated; colour dead white;
orifice oval elongated, rather exceeding in length one third of the
longer basal diameter of the shell. The radii are rather narrow, and
deeply depressed; they have their summits square: their outer lamina,
as explained under the genus, on both sides of each suture is in most
specimens divided into teeth, the points of which face and touch
each other. These teeth or notches give quite a peculiar appearance
to the shell, and alone suffice to discriminate this species; they
are sometimes blunt and partially obliterated, but it is rare to
find a specimen in which some few teeth do not occur in some one of
the six radii. I have, however, seen two or three specimens with all
six radii perfectly smooth; in one of these the general shape of the
shell, without the aid of any internal characters, almost sufficed
to show that it belonged to the present species; but in another
specimen, which had unusually narrow radii, and the whole surface of
which had undergone considerable disintegration, and was consequently
striated, could only be distinguished from _C. caretta_ by internal
characters. I have seen several specimens having very irregularly
shaped compartments, but generally the appearance of the whole shell
is highly symmetrical, like a star; and the genus was appropriately
named by old Klein, _Astrolepas_. In some specimens, in Mr. Cuming's
collection, from the Low Archipelago, in the Pacific, taken off the
toe-nail of a turtle, the shape was almost cylindrical; the shell
almost resembling that of _Coronula diadema_. The largest specimen
which I have seen was 2.3 of an inch in basal diameter, but only .55
in height.
_Structure of Shell and Radii._--After the full generic description,
the only point to which I need allude is, that the radiating,
parietal septa, as well as the descending sheath, are much thicker
than in _C. patula_, and that their basal edges can be plainly seen
by the naked eye to be dentated with numerous points. The thickness
of these plates and of the sheath varies considerably. In _C.
caretta_, I may add, the plates are broken up into many separate
points, and in this species the descending sheath is not generally
perforated, excepting at the sutures, by loop-holes for the entrance
of ribbons of corium. The sheath and inner lamina of the parietes
in _C. testudinaria_, taken together, fully equal one third of the
entire thickness of the walls. Flattened cavities or tubes run up
between the parietal septa for about two thirds of the height of the
inside of the shell. With respect to the radii, I have only to add,
that the thickness of the inner portion relatively to the generally
notched outer lamina, varies considerably.
_Opercular Valves_ (fig. 1 _b_-1 _d_).--These closely resemble
each other in all three species, and have been almost sufficiently
described under the genus. The scutum has its occludent margin always
sinuous, but to a variable degree. In one specimen there was a deep
little pit at the rostral end for the rostral depressor muscle. The
terga vary considerably in shape, according as the basi-carinal
corner is more or less truncated. The external furrow, marking the
position of the rudimentary spur, varies much in distinctness,
chiefly depending upon its nearness to the carinal margin of the
valve.
_Mouth and Cirri._--The labrum sometimes has its teeth worn away.
The second pair of cirri, relatively to the others, is shorter and
thicker than in the two following species.
2. CHELONOBIA CARETTA. Pl. 14, fig. 2.
LEPAS CARETTA. _Spengler._ Skrifter, Naturhist. Selbskabet, Bd. 1,
1790, Tab. 6, fig. 4.
BALANUS CHELYTRYPETES. _Hincks_ (!) sine descript., Annals of Nat.
Hist., vol. 5, p. 333.
CORONULA SULCATA. _Chenu._ Illust. Conch., Tab. 1, fig. 1.
_Shell globulo-convex, extremely massive and heavy; upper part worn,
sub-striated: radii either not developed or very narrow: parietes
without cavities extending up between the interrupted, radiating septa._
_Hab._--West coast of Africa; northern Australia; common. Attached
to, and generally imbedded in, the carapaces of turtles.
_General Appearance._--Shell extremely strong, massive, heavy,
broadly oval, globulo-convex, though sometimes depressed; dirty
white: surface in the upper part always worn and disintegrated,
hence finely striated; in the lower part a little rugged, slightly
folded, and occasionally, in deeply imbedded specimens, much folded
or ribbed. Orifice oval, not at all angular, rather exceeding in
length one third of the basal diameter of the shell. Radii either not
at all developed, or very narrow; but even in the former case the six
lines of suture are plain; and in the latter case the radii lie at
some depth beneath the surface of the shell. It is remarkable that
the compartments are hardly ever arranged symmetrically, the rostrum
and carina not facing each other exactly; and this holds good in
specimens attached separately, without any apparent cause for this
want of symmetry. The largest specimen which I have seen was 2.1 in
basal diameter; and this, which was a steeply conical individual,
was 1.1 in height; and therefore nearly twice as high as an ordinary
large specimen of _C. testudinaria_.
_Structure of Shell and Radii._--The descending sheath and radiating
septa are of very variable thickness, and have their basal edges
finely dentated. The septa are not continuous, from the circumference
to the sheath, in unbroken plates, but are irregularly divided into
separate, often short portions, and even occasionally into mere
points. The sheath differs from that of the other two species in
having loopholes for the entrance of ribbons of corium only on the
eight lines of suture, and not (with rare exceptions) in the middle
of each compartment: this is evidently due to fewer filaments of
corium being here sufficient to supply the less deep interspaces
between the radiating septa; for in this species there are no
flattened cavities or tubes running far up the shell. The inner
lamina of the walls cannot be here distinguished, for a solid, flat,
calcareous surface extends from the circumference, between the
radiating septa, to the sheath. The sheath, had it not been from the
light thrown on this part by the other species, would have certainly
been mistaken for the inner lamina of the walls. The absence of the
flattened cavities or tubes extending up the parietes, seems to be
the least varying character; and serves to distinguish this species
from those worn and massive specimens of _C. testudinaria_, which
have narrow and not-notched radii.
In specimens in which the radii are not developed, no vestige of the
outer lamina can be detected, the lateral faces of the adjoining
compartments being closely pressed together; but in specimens in
which the radii have been developed and have grown, the outer lamina
of course is present, and is extremely thick, with the growing edge
having a branched and mammilated surface, as in _C. testudinaria_,
but with the external surface not pitted or notched as in that
species. The inner portion of the radius, whether or not developed,
has nearly the same appearance, consisting of rather thick transverse
septa, branching from a central ridge, which is sometimes obscure.
The _Alæ_ are remarkably little prominent, as least in those
specimens in which the radii are not developed, so that Spengler
seems to have thought that the structure of the shell was essentially
different from that of Balanus, which certainly is by no means the
case. The edges of the alæ are very thick, nearly as thick as the
inner portion of the radius.
The _Opercular Valves_ hardly present any essential difference,
compared with those of the other species; but the occludent margin
of the scutum is apt to be more sinuous, and its rostral end blunter
and squarer. The carinal end of the tergum is also squarer than in
any common variety of _C. testudinaria_; the external furrow or spur,
near the carinal margin, is very indistinct, and even sometimes is
quite absent.
Neither the mouth nor cirri present any deviations from the generic
character.
3. CHELONOBIA PATULA. Pl. 14, fig. 3 _a_, 3 _b_, 4.
CORONULA PATULA. _Ranzani._ Memoire di Storia Naturale (1820), Tab.
3, fig. 25-28.
ASTROLEPAS LÆVIS. _J. E. Gray_ (!). Annals of Philosophy (new
series), vol. 10, 1825.
VERRUCA CANCRI AMERICANI. _Ellis._ Phil. Trans., vol. 50, 1758, Pl.
34, fig. 13.
_Shell steeply conical, very smooth and light; orifice large, generally
exceeding half the basal diameter of the shell: radii broad, smooth,
only slightly depressed._
_Hab._--Mediterranean; Gambia, West Africa; Charlestown; Jamaica;
Honduras; Brazil; Australia. Attached to Crustacea, smooth univalve
shells, and apparently to ships' bottoms.
_General Appearance._--Shell white, very smooth, of little specific
gravity, steeply conical, but not high; orifice broadly oval,
polygonal, very large, namely, generally exceeding half the basal
diameter of the shell. The summits of the compartments are usually
perfectly preserved, pointed, and often a little recurved. The radii
are rather broad, very smooth, with their summits slightly oblique
and arched: they are seated only a little below the general level
of the parietes. I have seen one specimen rather more than one and
a half inch in basal diameter, but this is an unusual size; this
species not growing to so large a size as the two foregoing forms.
_Structure of Shell and Radii._--The walls are here thinner than in
the two foregoing species; and the basal surface of a compartment
rarely equals half the basal diameter of the internal cavity of the
shell, measured transversely to its longer axis. The radiating septa
are also much thinner, generally sinuous, and so finely dentated
along their basal edges, that the teeth can be clearly perceived
only by the aid of a lens. The interspaces between the septa run up
to nearly the summits of the compartments, and hence the lightness
of the whole shell. The inner lamina of the parietes is here not so
thick (fig. 4), and is more distinct from the descending sheath than
in the foregoing species. The sheath is thin, like the radiating
septa; the medial loophole in each compartment, for the entrance of
a filament of corium, is much wider than in _C. testudinaria_, for
it is generally as wide as the bordering plate on either hand; and
in not a few specimens, the medial loophole is so wide as hardly
any longer to deserve being so called, for the descending sheath
is reduced to mere flattened pillars or legs on the sides of the
sutures. Although the parietes are here not nearly so thick as in
the two foregoing species, yet as the radii stand but little beneath
the level of the parietes, the outer lamina of the radius has a
considerable thickness, and is sometimes separated from the inner
portion of the radius by an interval, in the same manner as in _C.
testudinaria_; and the growing edges of the outer lamina exhibit
traces of the same mamillated structure as figured in that species.
The _Opercular Valves_ are apt to be rather narrower and more
elongated than in the foregoing species, with the occludent margin
of the scutum generally, but not always, only slightly sinuous:
externally the scutum is sometimes feebly striated longitudinally. In
the tergum (Pl. 14, fig. 3 _b_) the longitudinal furrow is generally
plainer, from being more distant from the carinal margin, and the
rudimentary spur itself is more prominent.
The mandibles usually have five narrow teeth; but I have seen
one specimen having only four teeth, and with the inferior angle
truncated. The cirri present no particular character.
SECOND SECTION OF THE SUB-FAMILY OF BALANINÆ.
[_Scutum and tergum (when both are present) not overlapping each other,
or articulated together; basis membranous; parietes often deeply
folded, with the outer lamina, towards the basis, generally imperfect;
each branchia composed of two plicated folds; shell attached to living
vertebrata._]
8. _Genus_--CORONULA. Pl. 15, 16.
CORONULA. _Lamarck._ Annales du Museum, tom. 1 (1802).
DIADEMA. _Schumacher._ Essai d'un Nouveau Syst., &c., 1817.
CETOPIRUS (SED NON CORONULA). _Ranzani._ Memoire di Storia Naturale
(1820).
POLYLEPAS. _J. E. Gray, (Klein)._ Annals of Philosophy, (new series),
vol. 10, 1825.
_Compartments six, of equal sizes: walls thin, deeply folded, with
the folds forming cavities, open only on the under side of the shell:
opercular valves much smaller than the orifice of the shell._
_Distribution_, mundane, imbedded in Cetacea.
This genus and the three following, namely, Platylepas, Tubicinella,
and Xenobalanus, have very considerable claims to be separated as a
sub-family, as has been proposed by Drs. Leach and Gray. Although in
structure and habits they are certainly closely related together; yet
only few characters can be predicated of all four in common,--some
characters failing in one genus and some in another. All four, however,
differ from the foregoing genera in the opercular valves not being
articulated together, being simply united by tough horny membrane;
but Chelonobia makes some approach in this respect. All four have the
branchiæ composed of a double fold; but this can hardly be considered
a character of much weight, as may be inferred from the remarks made
on this subject at p. 153 of the Introduction, where the differences
of these four genera from the other genera of the family are discussed
at some length. Altogether I have been led to conclude, though with
much hesitation, that these genera had better not be separated as the
sub-family of Coronulinæ.
Our present genus, Coronula, is closely related to Platylepas, and
likewise to Xenobalanus, though this latter genus is so very different
in external aspect that it might easily be mistaken for a pedunculated
Cirripede. Lamarck and some other authors have placed the species of
Chelonobia under Coronula, but this has arisen, as explained under that
genus, from a misapprehension of their structure; the folded very thin
walls in Coronula having been compared with the radiating septa of
the very thick walls in Chelonobia. I may further add, that Coronula
has been divided into two genera by Ranzani, on palpably insufficient
grounds.
_General Appearance._--The three recent species of this genus have a
very handsome and striking appearance. The shell is highly symmetrical,
owing to the six compartments being of the same size and having
exactly the same outline. The general shape is either depressed or, as
expressed by the name, like a crown. The walls are longitudinally and
slightly ribbed, owing, as we shall presently see, to their wonderfully
folded structure; and the surface is marked by very fine longitudinal
striæ, crossed by finely beaded lines of growth. Hence the walls
offer a strong contrast in appearance with the six, smooth radii,
of equal breadth. The symmetrically hexagonal, or rounded-hexagonal
orifice of the shell is closed by a thick, nearly horizontal membrane,
supporting, towards the rostral end, the small opercular valves, with
a slit, having protuberant lips, in the middle, for the protrusion of
the cirri. The opercular membrane is attached all round, but a short
distance beneath the summit of the shell. In regard to size, I have
seen a specimen of _C. diadema_ two inches in height and two and a half
in diameter.
_Structure of the Shell._--The structure at first appears singularly
complicated, and quite unlike that of any other Cirripede; but the
whole results simply from the folding of the very thin walls, which
in all essential respects are constructed like those in Balanus.
In a young specimen of _C. balænaris_, having the orifice of the
shell 2/10ths of an inch in diameter, I found the upper part of each
compartment only slightly sinuous, not more so than is common in many
varieties of Balanus, but more symmetrical, for each compartment had
three slight furrows, making, for the whole shell, eighteen furrows.
The ensuing changes during the growth of the shell will be best
understood by looking at the diagrams _a_, _b_, _c_, in fig. 10, Pl.
15, which are supposed to represent the basal margins of the walls
of a single compartment: (_a_) shows the simply sinuous wall of the
young shell. As the shell grows, the furrows rapidly grow deeper and
deeper (_b_), and wider and wider; at the same time, the folds or
ridges between the furrows gradually become drawn out at their ends
into transverse loops (_c_), the extremities of which ultimately
become closely pressed together--the furrows being thus converted into
cavities, extending from the top to the base of the compartments. This
structure in the mature shell will be best understood by looking at the
transverse or horizontal section (Pl. 16, fig. 7) of the rostral end of
the shell of _C. diadema_, in which species the folding of the walls
is simpler than in _C. balænaris_ or _C. reginæ_: the walls (_e_) are
represented by a double line connected by little cross lines,--(_h_)
being one of the transverse loops at the outer end of one of the folds,
and (_f′_) one of the cavities between the folded walls, open at the
bottom of the shell, and occupied by the epidermis of the Whale. The
walls of the compartments, as here represented, (A being the rostrum,
C C the lateral compartments, D D the alæ of the carino-lateral
compartments,) are separated from each other by the broad radii; but
if the section had been taken low down near the basis, the end of the
folded wall of one compartment would have been separated from that
of the adjoining compartment only by the close suture: this will be
understood by a glance at the entire shell, given in fig. 3, Pl. 15.
The rostrum of the same species, viewed from the inside, is shown at
fig. 1, Pl. 16; here it may be observed, that the basal margin (_e e′
e′′_) of the folded wall is extremely oblique, the outer portion having
extended downwards much more than the inner portion: this obliquity
is more clearly shown in the lateral view (fig. 2) of a lateral
compartment, for this figure will equally well serve for a lateral view
of the rostrum (fig. 1), if the ala (_a′_) be supposed to be removed.
The section of the rostrum in fig. 7, (A), will now be intelligible in
relation to the view of the rostrum given in fig. 1, if it be borne in
mind that the section has been taken high up, near the letters _a_,
_b_, _c′′_ in fig. 1.
As above stated, the folds or ridges in the young shell become more
and more transversely drawn out at their ends into the transverse
loops, till the latter join and touch each other. In _C. balænaris_
the lines of junction are simple, though very close; in the other
species, the ends of the transverse loops, where touching, are finely
and elegantly toothed, and thus locked together. These teeth appear
single when the shell is viewed either externally or internally, but
when the walls along the lines of junction are forced apart, they are
seen really to consist of transverse rows of minute teeth. These teeth
are less distinct, forming only sinuous ridges in _C. barbara_ (fig.
6, Pl. 15); in all cases the teeth are formed by the modification
of the very minute beads, which ornament the lines of growth on the
external surface of the shell. In the rostrum of _C. diadema_, as seen
internally (fig. 1, Pl. 16), the serrated and closely-fitting lines of
junction (_f_) between the ends of the transverse loops of the folded
walls are plainly shown. It is the more or less rounded surfaces of the
transverse loops which give to the external surface of the parietes its
longitudinally ribbed structure: the ribs are plainest in _C. diadema_,
fig. 3, Pl. 15. The shell, in fact, as seen externally, consists of but
an extremely small portion of the external surface of the whole length
of wall, being exclusively formed of the transversely looped ends of
the radiating folds, together with the radii. Owing to the ends of the
transverse loops being so closely pressed together, the furrows are
practically converted (as already remarked) into cavities, open only on
the under side of the shell, and extending from the oblique bases of
the compartments up to their apices; and these are invariably filled
by the black epidermis of the Whale. Owing to this circumstance, the
skin of the Whale has been mistaken by some authors for parts of the
Cirripede! In _C. diadema_, in which the summit of the shell is often a
little disintegrated, the whale's skin is often there exposed, forming
three black spots at the top of each compartment. It should always be
remembered that these flattened and deep cavities are furrows, which
homologically ought to be open in longitudinal lines along the external
surface of the shell, from the top to the bottom.
As the shell increases in diameter, each of the original eighteen
transverse loops, forming the exterior surface of the shell, increases
in breadth; and they would have had to increase extremely, had not
some of the transverse loops become, during growth, divided into two
or three new transverse loops, in a manner strictly analogous with the
first formation of the eighteen folds in the young shell. In Pl. 15,
fig. 7, 8, 9, we see how one of the circumferential transverse loops,
by the formation of a medial furrow, or rather bay, becomes developed
into two transverse loops; and it is rather important to observe
that three new loops might equally well have been contemporaneously
formed. By the repeated formation of new circumferential loops and
the consequent formation of new folds, the wall of the shell, when
old, especially in _C. balænaris_, becomes folded in a wonderfully
complicated manner, as may be seen in Pl. 15, fig. 5, which is an
exact tracing of the extreme basal edge of the wall of a shell of
_C. balænaris_; to perceive the full amount of complication, it is
advisable to trace the wall of any one of the compartments, from one
suture (_s_) to another. In this figure the sutures are purposely
drawn a little open. It may be seen that the new transverse loops, and
consequently the new folds of the walls, have been, in this species,
mostly formed in symmetrical order, on both sides of the six sutures;
this results from the transverse loops on the sutures almost always
giving rise contemporaneously to three new transverse loops. In _C.
diadema_ the transverse loops on the sutures usually divide into only
two new loops, one on the rostral side and one still remaining at the
suture; hence the folding of the walls in this species is much less
symmetrical. The number, however, of the transverse loops and the exact
pattern of the folding is variable in all four species of the genus.
I may further add, to show the complication of the folds, that in a
shell of _C. balænaris_, having a basal diameter of two inches, and
which had the walls as little folded as ever they are, yet I found,
by careful measurement, that the entire basal edge of the wall, if
stretched straight, would have extended for a length of fifty-two
inches! Therefore, if the wall had not been folded, but had been simply
circular, as in ordinary cirripedes, the basal diameter of the specimen
would have been between sixteen and seventeen inches!
The central membranous basis is flat, but the bottom of the folded
walls of the shell is concave, which is caused by the outer ends of the
folded walls having grown at a greater rate than the inner ends. The
concavity is deep in _C. diadema_; in _C. balænaris_ it is much less
so, and here the inner hood-like ends of the folded walls are rather
abruptly, but in a variable degree, produced downwards, generally even
slightly beneath the level of the circumference of the shell; this
fact is of interest in relation to the peculiar, depending, spur-like
processes in the genus Platylepas. A lateral view of a compartment in
both these species, is given in Pl. 16, figs. 2 and 3; and by supposing
in each case a compartment to stand opposite, at a distance which may
be judged of from fig. 5, the vertical sectional outline of the whole
shell will be understood: in fig. 3 of _C. balænaris_, however, the
inner hood-like ends of the folded walls are not produced so much
downwards as is usual.
In the same manner, as the outside of the shell consists of the
transversely expanded ends of the folded walls, pressed closely
together, so the cavity in which the animal's body is lodged, is formed
by the inner and less closely joined ends of the folds, lined by the
thick sheath (_a_, fig. 1 and 7, Pl. 16), which latter extends down
very near to the basal membrane. The cavity for the body, is small
compared to the whole shell; in _C. diadema_ it is deeply cup-formed,
with a small, flat, membranous bottom or basis; in _C. balænaris_ it
is wider and shallower, with a broader bottom, and with the upper
edges of the walls more inflected. In both species, the thick membrane
connecting the opercular valves to the shell, is attached all round
near the summit of the sheath. The uppermost portion of the sheath is
not marked by concentric lines, as in most of the Balanidæ, owing to
the opercular membrane not being, as we shall presently see, regularly
moulted. A portion of a single wall, when closely examined, is found to
be formed of an outer and inner lamina, united by longitudinal septa,
and is thus permeated by minute, square, longitudinal pores,--exactly
as in the normal structure of Balanus. The walls are extremely thin,
and are striated longitudinally, owing to the slight projection, on
both the inner and outer surfaces, of the longitudinal septa; they are
thicker in the part forming the external transverse loops, being here,
in _C. balænaris_, as much as 15/1000ths of an inch in thickness; but
when forming one side of the spoke-like folds, the thickness is only
6/1000ths of an inch. The inner lamina is thicker, contrary to what is
usual, than the outer lamina; the sharp tips of the longitudinal septa
project a little beyond either lamina, giving to the basal edge of the
wall a serrated outline. It is singular that the thin outer lamina
is first formed as a rim or ledge on each side of the longitudinal
septa; these ledges being not closely united for some little space up
the wall, as is represented in the enlarged drawing of a bit of the
basal wall of _C. diadema_, Pl. 16, fig. 6. The open clefts thus left
are, of course, covered by the so-called epidermis, for otherwise
the included threads of corium would have been exposed. Each fresh
period of growth, in the case of _C. diadema_ (fig. 6) and _reginæ_,
and to a certain extent in _C. balænaris_, is marked by little knobs
on the longitudinal, slightly prominent, septa, and this prefigures
an analogous strongly marked structure in Tubicinella. A fine thread
of corium runs up each pore to the summit of the compartment; for
these pores are not, as in Balanus, cut off by transverse calcareous
septa, or have their upper ends solidly filled up with shelly matter.
As, however, the summit of the shell in Coronula is sometimes
disintegrated, the threads of corium within the pores would have
been exposed, had not each thread formed for itself, as I suppose, a
transverse membranous septum near the summit of the shell; at least
this is the case with the larger pores of the radii. The walls, where
closely pressed together in the spoke-like folds, are disunited at
the extreme base, but above this they are firmly calcified together.
A ribbon of corium runs along the basal edge of each spoke, and sends
threads of corium up the parietal pores on each side, and its upper
edge serves to deposit calcareous matter (homologous with the layers
of the sheath) and thus to unite the two walls firmly together. In
_C. diadema_, the walls of the terminal transverse loops are simply
calcified together like the spoke-like portions; but in _C. balænaris_
the opposite sides of the loops are united by septa (see the transverse
section in Pl. 15, fig. 2 _a_), making from five to eleven longitudinal
tubes within each transverse loop; these tubes being larger than the
parietal pores. When a piece of the shell is dissolved in acid, no
tubuli can be discovered, which may be accounted for by the thinness of
the walls; nor are there any spines on any of the external membranes.
The number of the pores, in the parietes of a moderately-sized specimen
of _C. balænaris_, I calculated was at 3400, each occupied by a thread
of corium springing from the eighteen branched ribbons, diverging from
the corium, surrounding the base of the sack. To this number must be
added between 300 and 400 larger threads of corium running up the
tubes in the transverse loops; and no less than about 2300 fringes and
threads occupying the pores in the six radii: thus we see that the
dermal system in Coronula is wonderfully complicated.
_Radii._--The radii are very wide in _C. diadema_. In all the species
their summits are square or parallel to the basis. Their internal
structure is remarkable: as the walls in this genus are extremely thin,
so are the _proper_ radii, for in fact they consist in this and all
cases, as we know, of one margin of the wall modified by its lateral
growth against the opposed compartment. But as the radii in Coronula
are not plicated, like the walls, the shell would have been excessively
weak along the six lines of suture, had not the radii been strengthened
by numerous sinuous plates, springing from the inner lamina of the
proper radius, and running downwards, attached to the folded wall
of the compartment to which the radius belongs, and with their free
edges pressed against the folded wall of the opposed compartment.
These plates give out short transverse denticuli, making altogether a
beautiful structure, as is best seen in Pl. 16, fig. 3, but also in
fig. 2 and 4, and _d_ in fig. 1. In the section, fig. 7, the proper
radius (_d_) is seen to be continuous with the wall (_e_), and to be
very thin, in fact forming but a small portion of the compound radius:
it is formed of an outer and inner approximate lamina, separated by
septa, which are nearly horizontal, and which consequently cannot be
shown in the transverse section fig. 7. The outer lamina of the radius
is imperfect, or does not reach quite to the suture, leaving the septa
a little exposed (imperfectly shown in Pl. 16, fig. 3), exactly as is
the case with the outer lamina of the parietes, at the basal margin
of the shell. In all common Balanidæ, a ribbon of corium runs up each
of the six sutures, and sends in fine threads between the septa of
the radii, but here a thread of corium runs up a minute, cylindrical
pore, situated on the line of junction between the radius and the
wall whence it arises; and from this longitudinal thread the finer
threads spring which pass between the horizontal septa of the proper
radius: this cylindrical pore is rather large in _C. balænaris_, but
excessively small in _C. diadema_ (see a black dot (_d′_) in section,
fig. 7), and is solidly filled up in the upper part of the shell. The
plates (fig. 3) which run down from the inner lamina of the proper
radius, and form the greater part of its thickness, are occupied by
fringes of corium, extending up from a ribbon of corium, running along
each suture, like those which run along the bases of the spoke-like
folded walls. This difference in the origin of the ribbons of corium,
occupying the interspaces between the plates and the pores in the
proper radii, shows the essential difference between the latter and the
thick inner portions of the compound radii. In _C. balænaris_ (fig. 3),
the compound radius extends from the outside of the shell to the sheath
and to near the basal edges of the folded walls: in _C. diadema_ (fig.
2), it does not extend so far inwards and downwards; and in _C. reginæ_
(fig. 4), even still less so. It ensues from this circumstance that
when, in these two latter species, a transverse section is made across
the middle of the shell, a large chamber (_v_, fig. 7), occupied by the
ovaria, is found on each line of suture (_s s_), separating the radii
(_d_) and alæ (_a_).
The _Alæ_ in _C. diadema_ (section, Pl. 16, fig. 7, _a′_) and _C.
reginæ_, are of remarkable thickness, nearly equal to that of the
radii, and in _C. balænaris_, of considerable thickness; this is
evidently to give strength to the shell, which is weakest along the
lines of suture, notwithstanding that the radii have been specially
thickened. The edge of each ala presents a miniature resemblance of
the edge of the radius, namely, a central ridge sending off on both
sides sinuous plates, themselves denticulated. In _C. balænaris_, the
ala rests almost entirely on the inner surface of the compound radius;
but in the other two species, in which, as already stated, the radii
and alæ are separated by chambers, the ala rests on a plate (_c′′_ in
fig. 1, 4, and 7), which extends from the top of the radius to the
bottom of the sheath, narrowing downwards (_c′′_, fig. 1), and is a
specially developed portion of the sheath for the radius to rest on.
In _C. diadema_ and _reginæ_, the sides of the folded walls, at the
ends of the chambers (_v_, in section, fig. 7), are strengthened by the
deposition of layers of shell in connexion with the sheath. The sheath
extends close down to the basal membrane in _C. balænaris_, and does
not project freely: in the other two species it depends freely, but
does not run quite so low down. I have only further to remark, that the
sutures, though very strongly united, are not calcified together; for
they easily separate after the action of caustic potash.
The _Basis_ (Pl. 28, fig. 1 _a_-1 _c_) is membranous and is formed of
concentric slips: each slip has eighteen angles, corresponding with
the open ends (see Pl. 16, fig. 5) of the eighteen folds of wall. A
ray of membrane runs under each of these folds, being prolonged from
the basal membrane; but these rays can hardly properly be called parts
of the basal membrane, for they split at each period of growth along
the middle, and the two halves are drawn from under the basal edges
of the walls, and thus come to invest their outer surfaces. The basal
membrane, and the whole cementing apparatus, which is much simpler than
in Balanus, has been fully described (p. 135) in the Introduction.
_Opercular Valves._--In the sessile cirripedes hitherto examined, the
four opercular valves are inclined towards each other, and nearly fill
up the orifice of the shell, being united to the walls by a more or
less narrow circular border of membrane; this membrane being attached
rather low down to the sheath. In Coronula the opercular membrane is
stretched like the skin of a drum, almost horizontally across the top
of the shell; it is, however, generally attached to the sheath rather
lower down at the carinal than at the rostral end: hence the animal's
body, as remarked by Burmeister, is attached almost horizontally;
but this we shall see, under Xenobalanus and Tubicinella, cannot be
considered a character of much importance. The valves are quite small
compared with the opercular membrane, and certainly are of little
functional importance. The scuta in _C. balænaris_ and _diadema_ (Pl.
15, fig. 3 _b_) are sub-triangular; but the under or growing surface
is elongated and arched. These valves stand almost at the rostral
end of the orifice, instead of on each side of it. In _C. balænaris_
the terga are small, with the under surface oval: in _C. diadema_
they are either quite aborted, or are represented by a barely visible
plate of shell, parallel and close to the tergal margin of the scutum.
The aperture leading into the sack is formed by a nearly medial slit
of considerable length, furnished with irregularly protuberant,
inwardly inclined lips. These lips are formed by the development of
an inner fold or crest of membrane, which can be just detected in
most Cirripedes: the lips include a double fold of corium, and are
covered by a delicate tunic, continuous with that lining the sack, and
homologically continuous with the opercular membrane. The opercular
membrane is very thick, tough, and yellowish; it is, in parts, finely
plicated in lines radiating from the apices of the valves: these plicæ
consist of membrane in an altered condition, being harder, more horny,
and of a browner yellow; the plicæ are large at the rostral end of
the scuta, and projecting beyond these valves, they afford attachment
to the rostral depressor muscles. Rims of similarly modified membrane
(Pl. 15, fig. 2 _b_) connect the scuta and terga together. The rims and
plicæ are occasionally moulted together with the opercular membrane.
In almost all hitherto described Balanidæ, a new opercular membrane is
formed at each period of exuviation, and as soon as formed, the old one
is generally moulted, together with the other membranes of the body:
the case is very different in Coronula, in which it is evident, from
the lines of growth on the valves and sheath, that a new membrane is
formed only at rather long intervals, and that it is formed in some
degree extensible, so as to allow of some growth in the shell. Two or
even three of these membranes are retained at the same time, one over
the other; and thus, by their joint thickness, they afford protection
to the included animal's body, and compensate for the smallness of the
opercular valves. In a large specimen of _C. balænaris_, two inches in
diameter, there had been formed, since its existence as a very young
shell, not more than eight opercular membranes, whereas the other
membranes must have been moulted within this same period at least
thirty times. In a young specimen of this species, having the orifice
of the shell only two tenths of an inch in diameter, I found the
opercular membrane, as usual, double. This membrane is not furnished
with spines, nor is it penetrated by tubuli as in most other genera.
The tissue left after the opercular valves have been dissolved in acid,
presents no tubuli, or any other recognisable character, and is not
divided (as is usual) into layers.
_Muscles of the Sack._--These muscles differ considerably from
those of other sessile cirripedes hitherto described. There are the
usual five (or properly six, the two tergal muscles being here, as
elsewhere, confluent) bundles of fasciæ; but they hardly can be called
bundles, they are so much spread out and thin. In _C. balænaris_,
each rostral muscle consists, in different specimens, of from three to
five principal fasciæ; these, at their upper ends, are attached to the
cartilaginous plicæ at the rostral extremities of the scuta; at their
lower ends, they do not reach (as in all the previous genera) to the
basal membrane, but after converging, they diverge again into a little
fan of fibres, which are firmly attached to the corium low down on the
sides of the sack. The lateral depressores of the scuta consist each
of about three fasciæ, and they terminate downwards like the rostral
muscles. The tergal depressores are spread out into a thin sheet;
upwards they reach to the basal edges of the lips of the sack-aperture,
and downwards they curve a little towards the rostral and opposite end
of the shell, and extend nearly to the basal membrane. These tergal
muscles include two fasciæ, larger than the others, which extend rather
further, both upwards and downwards, than the other fasciæ. But the
most novel character in these several muscles is that in their lower
portions they do not exhibit transverse striæ, thus showing a tendency
to become involuntary as in pedunculated cirripedes. This circumstance,
and their feebleness, is easily accounted for by the thick unyielding
nature of the opercular membrane, and the feebly developed character
of the opercular valves. In _C. diadema_, the tergal muscles are much
spread out, having, as in _C. balænaris_, a larger fascia on each side;
but the lateral depressores of the scuta form a well defined nearly
cylindrical bundle; the rostral pair are extremely weak and spread out:
I could perceive only feeble transverse striæ on some of these muscles,
and on others there was not the least trace of striæ. I may add that
the adductor scutorum muscle is well developed, as are the eight pairs
of muscles which unite the animal's body to the under surfaces of
the scuta. The action of the adductor scutorum serves to close the
sack-aperture, but towards the carinal end necessarily with very small
force; protection in this part can only be afforded by the protuberant,
valvular lips, and by the dorsal surfaces of the inwardly curled cirri,
with their tufts of bristles.
_Mouth._--The mouth (Pl. 26, figs. 3, 4) is much elongated
transversely, and does not differ essentially from that of most other
Balanidæ. The labrum is notched and not in the least bullate, though
in _C. diadema_ there is a slight prominence outside close beneath
the notch. The palpi are of large size, with their bristly apices
touching each other. The mandibles are very strong, with from four to
five main teeth, which are remarkable by presenting only rudiments
of being laterally double; but between the second and third, and
between the third and fourth main teeth, there is a small intermediate
tooth,--these I have not met with in any sessile cirripede hitherto
described. The maxillæ are small, with the two upper spines remarkably
strong. The outer maxillæ are on their inner faces bilobed. Between
these organs there is a minute projection, or mentum, flattened in the
longitudinal axis of the body; I have not noticed this in any previous
Cirripede.
The _Cirri_ are short and extremely much flattened: the three anterior
pairs have their rami unequal in length by two or three segments; the
posterior edges of their pedicels are fringed by tufts of extremely
fine hairs. The pedicel of the first cirrus is very broad; its rami are
short, with the segments very broad. The rami of the second and third
cirri are short, with the segments protuberant in front and thickly
clothed with spines; the terminal segments have some short, thick,
claw-like spines. The three posterior pairs have protuberant segments,
each supporting three or four pairs of short, strong, main spines, with
a small intermediate tuft: the dorsal tuft is small.
The prosoma is of large size. The _stomach_ is large, without cæca, but
with some internal longitudinal plaits (in _C. balænaris_ at least),
showing a tendency to the formation of cæca. In the stomach of _C.
balænaris_ I found a considerable quantity of a conferva, too much, I
think, to have got in accidentally.
_Generative system._--The vesiculæ seminales are large; and at their
broad blunt ends, in _C. balænaris_, four separate vasa deferentia
enter, of which fact I have seen no other instance. The ovarian tubes
do not extend up the sides of the sack, but lie at the bottom, over
the basal membrane; in _C. balænaris_ they send six short ray-like
prolongations into the six sutures; in _C. diadema_ they send similar
prolongations into the sutures, and fill up, as I believe, the six
chambers (Pl. 16, fig. 7, _v_) lying between the radii and alæ: I have
examined only one specimen of _C. diadema_ in spirits, and this had
the ovarian tubes in an early state of development, when they can with
difficulty be distinguished from the pulpy corium; the orange-coloured
masses, however, which filled the six cavities, resembled the layer
which certainly consisted of undeveloped ovarian tubes and cæca, lying
over the basal membrane. The ova are wonderfully numerous; their length
is 3/400ths of an inch. The larvæ have been noticed in the anatomical
introduction.
_Branchiæ._--These are immensely developed, covering almost four fifths
of the area of each side of the sack. Each consists of two nearly
equal folds, attached vertically to the carinal end of the sack, and
transversely across the upper end extending to the animal's body. The
outline of the free part is rounded. Both folds of both branchiæ are
deeply plicated on both sides; hence the superficies of the whole is
very great. We shall find that this structure is common to the three
following allied genera, but with these exceptions, I have observed
double branchiæ only in one species of one other genus, namely, in
_Chthamalus dentatus_.
_Attachment._--The shells adhere with remarkable strength to the
whale's skin. Having, until recently, examined only separated
specimens, and observing portions of the whale's skin adhering to
the outside, and solidly filling up the cavities on the under side,
I did not doubt that the shell had the power of forming, by its own
action, a deep cavity in the skin of the whale. Inspection of the basal
outline of the walls of the shell (Pl. 16, fig. 5) of _C. balænaris_,
will show how singularly unfitted its structure is for any burrowing
process; and I was led to speculate on the possibility of the pupa
being able to bury itself deeply in the skin, but rejected this
view as opposed to what is known of the habits and structure of the
pupæ of other Cirripedes. Having now examined several specimens of
_C. diadema_ adhering, in a group, to a large piece of skin, in Mr.
Stutchbury's collection, it has become evident that the attachment
is as much owing to the upward growth of the whale's skin, as to the
downward growth of the Coronula. In Pl. 15, fig. 4, a vertical section
is given of the whale's skin, taken through the place whence a shell
of _C. diadema_ has been removed; consequently we here see nothing
but the whale's skin: the upper black layer is the dark horn-coloured
epidermis, forming the general surface of the whale's body, and
resting on an orange-coloured fibrous layer, which is lightly shaded
in the drawing. The two horns in the section are two of the eighteen
projections, formed entirely of the dark epidermic layer, which fill up
the eighteen flattened cavities produced by the folding of the walls.
Outside the horns we see the section of a circular furrow, in which the
circumferential margin of the shell was lodged; and between the horns,
there is the central hollow, within which, when lined with shell, the
cirripede's body was included. The circular furrow is formed in main
part by the epidermic layer being thinner there than on either side,
and partly by the orange-coloured, underlying fibrous layer curving a
little downwards, from having apparently yielded to the pressure of the
circumferential margin of the shell. With respect to the cause of the
thinness of the epidermis under the circular furrow, I do not know how
much to attribute to mere mechanical compression or stretching, and how
much to the pressure of the shell, having checked[124] its formation.
In the case of very young and small shells, it is hardly possible that
their pressure can have in any way influenced the formation of new
epidermic layers under the thick old layers; and we must believe, at
least in these cases, that the whole effect is mechanical, the sharp
basal edges of the shell having indented the epidermis; but this is
not more surprising than that the radicle of a plant should penetrate
hard ground. Whether the indented epidermis in the circular furrow
becomes ruptured, I am not sure; ragged layers may commonly be observed
outside the shell, but it is very possible that these may be the ends
of layers of epidermis which have been preserved by the covering of
the shell, whereas, on the surrounding parts of the whale's body,
these same layers have been removed by disintegration. To return to
the section,--the outline of the boss of orange-coloured fibrous
tissue, under the central hollow, clearly shows that it must have
been formed by its own upward growth, for it stands above the general
surrounding level of the corresponding layer. This same conclusion is
still more obvious with respect to the eighteen flattened prominent
horns, formed of the dark epidermis; the manner in which the epidermis
has been forced, moulded, and packed into the eighteen flattened and
curved cavities of the shell, so as to adhere to them with considerable
tenacity, is extremely curious. The prominence of these horns is so
great that it appears to me quite impossible to account for them,
excepting by a special formation of epidermis beneath each cavity.
The basal membrane of the Coronula, which lies at the bottom of the
central hollow, adheres by its own cementing apparatus; and when the
larva first attaches itself, this adhesion must be very important, as
it allows the basal edges of the shell, during their slow downward
growth, to press firmly on the whale's skin, and so slowly indent it
with the circular furrow. The final cause, probably, of the cavities
on the under side of the shell in this genus, formed by the singularly
convoluted parietes, is to allow of the upward growth into them of the
epidermis of the whale, thus securing a firm attachment and allowing
the shell to exert a strong downward pressure, and thus effect its
partial imbedment, and protection from the enormous force of the waves
to which it must be exposed.
[124] Formerly I was inclined to believe, as stated in my former
volume on the Lepadidæ, that the cement injured the true skin of
the supporting animal, but this, at least in such cases as the
present, I do not now at all believe.
With respect to _C. balænaris_, I have seen only specimens, preserved
on shrunk and twisted whale's skin, with the underlying fibrous
layer not preserved; but the cavities in the shell were filled by
horns of epidermis, exactly as in _C. diadema_. There is, however,
this difference in the attachment of the two species, that in _C.
balænaris_, owing to its depressed form, the circumference of the shell
indents the whale's skin, not vertically downwards, as in _C. diadema_,
but very obliquely outwards; and, consequently, buries itself much more
completely, but less deeply, under a folded and apparently ruptured
flap of the epidermis. In young specimens, of the size of a shilling,
the entire shell, with the exception of the operculum, is thus covered
up and protected whilst young and tender.
_Geographical Distribution._--The genus is found wherever whales
occur, that is, from the Arctic to the Equatorial regions, in both
hemispheres. It is asserted that sometimes as many as a couple of
hundred specimens will adhere to a single whale. _Coronula barbara_,
a form closely allied to _C. diadema_, existed during the Red Crag
period; and Bronn has described some fossil specimens from Italy.
_Affinities._--In the wonderfully convoluted shell,--in the parietal
tubes not being either filled up by calcareous layers or being crossed
by calcareous septa,--in the outer lamina of the shell between the
longitudinal septa near the basis being imperfect, Coronula differs
from all the foregoing genera; in the two latter respects it agrees
with the three following allied genera, viz., Platylepas, Tubicinella,
and Xenobalanus. The equal size of all six compartments of the shell,
has been observed in very few genera besides Coronula. In a new
opercular membrane not being formed at each exuviation, and in two or
three of these membranes being persistent, and in their being attached
high up the sheath, this genus agrees with the three following genera
alone. In the valves tending to be rudimentary, and in the protuberant
lips of the sack-aperture, we have a close alliance with Xenobalanus.
The muscles of the sack being spread out, and tending to lose their
transversely striated character, are great peculiarities in Coronula,
Tubicinella, and Xenobalanus. The simplicity of the cement-ducts is
a remarkable character, observed in two of the following genera, but
not in Tubicinella. The double branchiæ is a peculiarity common to all
four genera. Neither the mouth nor cirri offer any new characters of
much importance in Coronula or in the three allied genera: the lower
teeth of the mandibles not being laterally double, but having small
intermediate teeth, is the newest feature in the mouth.
1. CORONULA BALÆNARIS. Pl. 15, fig. 2, 2 _b_: Pl. 16, fig. 3, 5.
LEPAS BALÆNARIS. _Gmelin._ Systema Naturæ (1789).
---- ---- _Chemnitz._ Conch., vol. 8, Tab. 99, fig. 845, 846 (1785).
BALANUS ---- _Bruguière._ Encyclop. Method., Pl. 164, fig. 13-18
(1789).
CORONULA ---- _Lamarck._ Annales du Museum, vol. 1, Tab. 30, fig. 2-4
(1802).
---- ---- _Chenu._ Illust. Conch., Plate, fig. 1 and 4.
---- ---- _De Blainville._ Dict. des Scien. Nat., 1818 and 1824,
Tab. 117, fig. 3, 3 _a_.
_Shell much depressed, with longitudinal flattened ribs, having simple
edges; orifice rounded-hexagonal: radii very thick, almost equalling
the shell in thickness: opercular valves four._
_Hab._--Attached to whales in the Southern Ocean.
Having described, under the genus, in so much detail the structure of
the whole shell, it will be sufficient here to point out the characters
by which this species differs from the others.
_General Appearance._--The shell is generally much depressed, though
sometimes, in large specimens, from the turning in of the basal
edges, a considerable degree of convexity is acquired. The radii
are moderately wide, and give a star-like appearance to the shell.
The surface of the whole upper part of the shell is smooth; the
broad and much flattened ribs (_i. e._ the terminal, transverse
loops of the folded walls) of which each compartment is formed,
generally divide at a short distance from the apex. The close sutures
separating these ribs are straight, and not finely serrated, by which
character alone this species can at once be discriminated from the
others. The surface, when closely examined, is found to be finely
striated longitudinally, and is transversely crossed, chiefly in
the lower part, by minutely beaded growth-ridges. The orifice is
rounded-hexagonal, and is small compared with the whole shell; it is
also smaller than the basal edge of the internal cavity of the shell;
consequently the whole basal edge cannot be seen through the orifice
from one point of view,--the operculum of course having been removed.
An unusually large specimen was 2.75 of an inch in diameter; this
individual was one of the more convex varieties, and yet its entire
thickness, from top to base, was only .9 of an inch.
_Operculum._--Having described, under the genus, the general
structure of the opercular valve, I will here only add a few details
on their shape. They are small compared with the whole extent of
the opercular membrane. The scuta (Pl. 15, fig. 2 _b_, scutum to
the left) stand near each other at the rostral end of the aperture
leading into the sack, with their two rostral ends united by the
yellowish-brown, longitudinally plicated, horny membrane, described
under the genus. The terga (fig. 2 _b_) stand a little apart from
the scuta, on the sides of nearly the middle of the sack-aperture.
The lips are protuberant and moderately developed all round. The
scuta are elongated, and a little curved; including the upper
imbedded portion, they are almost sub-triangular: but the under
growing surface is much elongated, nearly flat, with the two ends
of nearly the same width, truncated and rounded. The _Tergum_, in
rather small specimens is, if the upper imbedded portion be included,
sub-triangular, with the growing surface oval, and between one third
and one half of the length of the scutum; but in large specimens,
the tergum becomes style-formed, lying parallel to the tergal margin
of the scutum, with the growing surface proportionally much smaller,
and not above one fifth of the length of the growing surface of
the scutum. Hence we see some tendency in the tergum to become
rudimentary, as it is in the next species. The brown, horny, plicated
substance in which the terga are imbedded, extends considerably
beyond the valves themselves.
_Structure of the Shell and Radii._--I have already so fully
discussed this subject, that I will here only enumerate the points in
which this species differs from _C. diadema_ and _reginæ_:--Firstly,
the more symmetrically folded walls (Pl. 16, fig. 5), new folds
arising on both sides of all six sutures. Secondly, the inner ends of
the folded walls, which surround the internal cavity, being almost
square, but with their angles rounded; their inner ends descend some
little way beneath the basal edge of the sheath, as low, or lower,
than the circumference of the shell. Thirdly, the external lines of
junction between the transverse terminal loops being smooth or not
serrated. Fourthly, these loops being flatter towards the outside,
and being internally filled by septa (Pl. 15, fig. 2 _a_); tubes
being thus formed larger than the proper parietal pores. Sixthly, the
sutural edges of the compound radii (Pl. 16, fig. 3) being very much
broader in the inner and upper part of each compartment than in the
outer and lower part; for in the inner and upper part they stretch
from the outside of the shell to the sheath, so that the alæ rest on
them, and hence no large open cavity is left between the radii and
alæ; in the inner and upper part, also, the radii extend down nearly
to the basal edges of the folded walls. The septa, of which the radii
are formed, stand further apart than in _C. diadema_ and _reginæ_.
Seventhly, the alæ are only 1/5th or 1/6th of the thickness of the
radii, whereas in the two other species they are very much thicker,
being nearly as thick as the radii: the alæ are also here (fig. 3,
_a′_) squarer than in those two species, that is, their basal margins
are not so short compared with their upper margins; their edges
present, also, a slightly different structure. Eighthly, the basal
edge of the sheath does not here project freely, and does not descend
quite so close to the basal membrane.
_Mouth._--The palpi are furnished with a row, extending along the
whole basal exterior margin, of very long upwardly pointing spines;
on their upper edges there is a brush of small spines. The edge of
the labrum is clothed with extremely fine spines, and is furnished
with a few minute teeth. The mandibles have four teeth, and the lower
end is broadly rounded and coarsely spinose: the second and third
teeth are double at their points, and between the second and third,
and again between the third and fourth, there is a small intermediate
tooth.
Regarding the _Cirri_, I have nothing especial to remark.
_Geographical Distribution._--I have examined nine sets of specimens,
having localities attached to them: three from off New South Wales;
two off the Cape of Good Hope; two from the west coast of South
America; and two marked only "South Sea." Hence I am led to conclude
that this species is confined to the southern hemisphere, or if it
extends into the northern hemisphere, it is probably only in the
Pacific Ocean. I do not believe (though so stated in some works) that
this species occurs on the shores of Europe. It seems often to be
associated with Tubicinella. Some specimens thus associated, sent by
Mr. Bennett to Professor Owen, were said to have been attached to the
_Balæna australis_.
2. CORONULA DIADEMA. Pl. 15, fig. 3, 3 _a_, 3 _b_; Pl. 16, fig. 1, 2, 7.
LEPAS DIADEMA. _Linn._ Systema Naturæ, 1767.
---- ---- _Chemnitz._ Conch., vol. 8, Tab. 99, figs. 843, 844.
BALANUS DIADEMA. _Bruguière._ Encyclop. Method., n. 164, fig. 13, 14
(1789).
CORONULA ---- _De Blainville._ Dict. des Sc. Nat. (1824), Tab. 117,
fig. 4.
---- ---- _Leach._ Encyclop. Brit. Suppl., vol. iii, 1824.
---- ---- _Chenu._ Illust. Conch., Plate, fig. 3.
---- ---- _Burmeister._ Beiträge zur Naturgeschichte der
Rankenfüsser, 1834, Tab. 2, fig. 1-14, 18.
_Shell crown-shaped, with longitudinal convex ribs, having their edges
crenated; orifice hexagonal: radii moderately thick, very broad: terga
absent or rudimentary._
_Hab._--Attached to whales, in the Arctic Seas; United States and
Great Britain; Gulf-Stream, Atlantic Ocean; New Zealand(?).
_General Appearance._--As previously remarked, owing to the fulness
of the generic description, minute details on structure, excepting
those characteristic of the present species, need not here be given.
The crown-like shape of the shell is well expressed by its name of
_Diadema_, but the crown tends to pass into a cylinder. The radii are
extremely broad. The orifice is large, and neatly hexagonal: when
the operculum is removed the whole inside of the cup-formed shell
can be seen at once, for the flat membranous basis is much smaller
than the orifice. The under side of the shell is deeply concave.
The outside of each compartment is formed by broad, rounded, and
somewhat prominent, rarely divided, ribs (_i. e._ the transverse
ends of the folded walls); these ribs are closely united together
by finely serrated lines of junction (Pl. 16, fig. 1, _f_). Their
surfaces outside are delicately striated longitudinally, and plainly
crossed (more plainly than in the foregoing and the next species) by
irregular, transverse ridges, especially in the lower part of the
shell. The largest specimen which I have seen was two and a half
inches in diameter and two in height.
_Scuta._--These are placed close together at the rostral end of the
orifice, and are imbedded in the brownish, tough, longitudinally
plicated, horny substance, which extends far beyond both ends of the
valves. In outline (fig. 3 _b_) they are mitre-shaped, or rounded
and sub-triangular, a little curved, and more or less elongated,
being most so in young specimens; they are, however, less elongated
and rather more massive than in _C. balænaris_. _Terga_,--these seem
entirely absent in most specimens; but in one (fig. 3 _a_) I found
a rudiment, namely, a short thin plate of shell, barely visible to
the naked eye, extending parallel and near to the tergal margin of
the scutum. The lips of the aperture of the sack are prominent, and
highly so towards the carinal end.
_Structure of Shell and Radii._--Owing to the shell not spreading
much at the base, new folds in the walls are much seldomer formed,
and therefore the external longitudinal ribs (_i. e._ the terminal
transversely elongated loops), are much seldomer divided, than in _C.
balænaris_ or _reginæ_; even rather large specimens sometimes having
only the original eighteen folds.[125] When new folds are formed,
only one is formed on one, viz., the rostral, side of each suture,
instead of on both sides, as in _C. balænaris_. The inner ends of the
folded walls, surrounding the basal membrane, are narrow, instead
of being almost square, as in _C. balænaris_. The lower edge of the
sheath, which projects freely, descends almost to the level of the
basal edges of the walls. The outer ends of the folded walls, forming
the transverse loops, are internally filled up solidly by calcareous
matter, instead of by septa forming tubes, as in _C. balænaris_.
The radii (Pl. 16, fig. 2) are a little thicker in the lower and
outer than in the upper and inner part of each compartment; in the
middle, they do not reach the sheath by about half the thickness of
the compartment, and consequently they are separated from the plates
(_c′′_ in fig. 1 and 7) on which the alæ rest, by large chambers (_v_
in fig. 7), which extend up to nearly the apices of the compartments:
the extent, however, to which the upper ends of these chambers have
been solidly filled up, varies a little. The sinuous plates forming
the main portion of the compound radii are rather thinner and closer
together than in _C. balænaris_. The alæ are thick, being thickest in
the middle part, and there equal the radii in thickness; their lower
margins are very short compared with their upper margins, hence they
are almost wedge-formed.
[125] These should be counted on the under side of the shell,
for if counted from the outside, the number would be thought to
be twenty-four, as on the side of each compartment bordering the
radius, a half-fold has the appearance of being a whole fold, so
that in appearance six folds are added to the real number of the
folds. This caution is necessary whatever the number of real folds
may be, that is, whether or not the original eighteen folds have
been increased by subsequent plications. These remarks, also, are
applicable to the other species.
_Mouth._--The teeth and fine hairs on the labrum are sometimes
obscure, and sometimes plain: close outside the bottom of the medial
notch, there is a small hard prominence. The palpi are broad; on
their basal exterior margins there is a short row of spines, which
do not equal in length the width of the palpi, and therefore are not
so long as in _C. balænaris_. The mandibles have five main teeth, of
which the second and third show only an obscure rudiment of being
double; between these two teeth, and between the third and fourth
tooth, there is a small intermediate tooth: the inferior angle is
narrow, rounded, and spinose. The extremity of the apodeme of the
maxillæ is expanded.
_Cirri._--I have only to remark, that the pedicel of the first cirrus
is extremely broad, and that the rami are set on in an unusually
crooked manner: the basal segment of the shorter and broader ramus of
this cirrus has its dorsal surface produced into a plate fringed with
very fine hairs.
_Geographical Distribution._--I have received only four specimens
with certain localities attached to them: namely, the Arctic Seas of
Scandinavia; the coast of the United States, and of Britain; and the
Gulf-Stream. There is also a specimen in the British Museum, sent by
Mr. Stephenson, mingled with shells of Mollusca from New Zealand;
but a Coronula, procured from a floating whale in the early part
of the outward voyage, might so easily be sent home with specimens
subsequently collected in another county, that I do not as yet fully
admit that this species is an inhabitant of the Southern Pacific
Ocean: I am less willing to admit this, from suspecting that _C.
reginæ_ in the Pacific, replaces the _C. diadema_ of our Northern
Seas.
3. CORONULA REGINÆ. Pl. 15, fig. 5; Pl. 16, fig. 4.
_Shell globulo-conical or depressed, with longitudinal, much flattened
ribs, having their edges crenated, and their surfaces striated and
granulated; orifice hexagonal: radii thin, not exceeding one fifth of
the thickness of a compartment: terga absent._
_Hab._--Attached to whales, Pacific Ocean; Mus. Cuming, Stutchbury,
and Darwin.
I have seen eight specimens of this form--two in Mr. Cuming's
collection, one of which was procured at Iquique in Peru, the other
from an unknown locality; one was given me in Chile, and was stated
to have come from the Pacific; two others are in Mr. Stutchbury's
collection, from unknown but distinct localities, and three in a
group in the British Museum. These eight specimens, of which six were
collected separately, all strikingly agree in general aspect and in
details of structure, so that I can easily recognise the shell, and can
at once pick out a single compartment, when mingled with those of the
two foregoing species; nevertheless, the differences are so small, that
I at first hesitated whether to name the species; but, upon reflection,
I am convinced that it is distinct. _Coronula reginæ_ is much more
closely related to _C. diadema_ than to _C. balænaris_, and I suspect
that it replaces in the Pacific the former of these two species.
_General Appearance_; shell conical, straight-sided; some specimens
being nearly as much depressed as _C. balænaris_, and considerably
more depressed than any variety of _C. diadema_; other specimens
being globulo-conical, but rather less globular than the ordinary
form of _C. diadema_. I may here remark, that shape is of more value
in this genus, in which the shell is attached to the yielding skin
of whales, than in those genera in which it adheres to rocks. The
orifice is neatly hexagonal, and the whole internal cavity, when the
opercular membrane is removed, can be seen from one point of view,
owing to the contraction of the lower part of the cavity and small
size of the basal membrane. The surface of the shell is smooth, but
has, even up to the summit, a peculiar frosted appearance, different
from that of the foregoing species, caused by the longitudinal
striæ being rather more distinct, and being crossed by beaded, very
delicate transverse lines of growth. The ends of the transverse
loops, forming the exterior surfaces of the compartments, are much
flattened, even more so than in _C. balænaris_. The lines of junction
between the loops, are finely serrated, as in _C. diadema_; and
internally they are solidly filled up, instead of being formed into a
set of tubes by longitudinal septa, as in _C. balænaris_. The under
side of the shell, with its folded walls, presents an appearance
intermediate between the variable appearance of this part in the
above two species.
The sutural edges of the radii (Pl. 16, fig. 4) offer by far the
most remarkable character, in their thinness from top to bottom; for
they hardly exceed one fifth of the thickness of the compartment,
measured from the external surface to the base of the sheath; hence
a very large cavity is left between the radii and alæ: in the
thickness of the radii the three species already described form a
series, _C. reginæ_ at one end and _C. balænaris_ at the other. The
sinuous plates, forming the lower part of the radius, are coarser and
stand rather further apart than in _C. diadema_. The alæ are thick,
and have the same outline, being narrow at their basal margins and
broad at top, as in _C. diadema_, with their sutural edges similarly
constructed: the basal edge of the sheath likewise projects freely.
_Operculum._--This resembles most closely that of _C. diadema_. There
are no rudiments of terga. The scuta cannot be distinguished from
those of _C. diadema_.
_Mouth._--The labrum has a row of inwardly curved little teeth along
the whole crest, and these I did not notice in _C. diadema_: there
is only a trace of the prominence on the outside at the bottom of
the central notch. The hairs on the basal exterior margin of the
palpi are moderately long. The mandibles have five teeth. In the
_Cirri_, the first pair resembles in its peculiar structure those of
_C. diadema_. In the sixth pair, however, the segments support only
three pairs of main spines: but the specimen was not very large, and
probably in old specimens there would have been four pairs.
_Summary._--The present species differs from both the foregoing
only in its rather more conical and straight-sided outline, smooth,
frosted surface, and in the narrowness of the sutural edges of the
radii, and consequent large size of the chambers between the radii
and alæ. It resembles _C. diadema_, as far as the shell is concerned,
in the external ribs or transverse loops having their lines of
junction serrated and in being solidly filled up--in the shape of
the orifice and of the internal cavity of the shell--in the shape
and structure of the alæ--and in the basal edge of the sheath being
free. It comes nearer to _C. diadema_ than to _C. balænaris_ in the
structure of the radii. It differs from _C. diadema_, and comes
nearer to _C. balænaris_, in the external ribs being flattened,
instead of being convex, and in the lines of growth being very
delicate. But as it resembles _C. diadema_ in the several foregoing
characters of its shell, in the opercular valves, in all parts of the
mouth (excepting the labrum), and in the cirri, it is very much more
nearly related to that species than to _C. balænaris_.
4. CORONULA BARBARA. Pl. 15, fig. 6.
CORONULITES DIADEMA (?) _Parkinson_, Organic Remains (1811), vol. 3,
p. 240, Pl. 16, fig. 19.
_Shell (probably) crown-shaped, with longitudinal convex ribs, having
their edges crenated, and their surfaces rugged, both externally and
internally, with transverse ridges: radii moderately thick; the spaces
between the radii and the alæ solidly filled up._
_Fossil_ in Red Crag, (Bawdsey and Sutton); Mus. S. Wood and
Geological Society.
The species here described, though near to _C. diadema_ and easily
confounded with it, I have no doubt is distinct. I owe to the kindness
of the Rev. Mr. Image an examination of the original specimen figured
by Parkinson; and in Mr. Stutchbury's collection there is a similar and
more perfect specimen; both of these resemble _C. diadema_ in general
form, but have been too much worn to be positively identified. The
following description is drawn up from some compartments collected by
Mr. Searles Wood, belonging certainly to three, and probably to four
individuals, one of which was young; as these specimens agree in all
essential respects, I feel pretty confident that the characters, by
which the present species differs from _C. diadema_, are of specific
value.
_Structure of Shell._--The longitudinal ribs on each compartment
(_i. e._ the terminal transverse loops), are convex and prominent,
as in _C. diadema_, but they are crossed by more prominent ridges
of growth than even in the roughest varieties of that species, so
that the surface of the shell is more rugged. In the three previous
species, the surface of the wall entirely round the cavities occupied
by the whale's skin, is striated only by longitudinal very fine
lines; but here, the outer portion, or that (fig. 6) formed by each
transverse loop, is crossed by transverse ridges of growth, like,
but less prominent than, those on the external surface of the shell.
The minute teeth, along the lines of junction between the transverse
loops, are here less regular, and can hardly be said to exist; for
the two edges are locked together by what may be more strictly
called minute zig-zag ridges than teeth. The exact number of the
circumferential plications in the walls is variable in the same
manner as in the three foregoing species. The sutural edges of the
radii are about as thick as, or rather thicker than, in _C. diadema_.
As in this latter species, and in _C. reginæ_, each ala rests, not
on the internal surface (as in _C. balænaris_, and in all other
Balanidæ) of the radius, but on a special plate; but in _C. barbara_,
instead of a deep chamber, running up to the apex of the compartment,
being left between the radius and ala, this part is filled up almost
entirely by solid shell. Although the extent to which this chamber is
filled up varies a little, and although its depth varies a little in
_C. diadema_, yet there is a marked difference between the specimens
of this latter species, in which the chamber is most filled up, and
those of _C. barbara_, in which it is least filled up. The alæ are
thick, as in _C. diadema_, and their sutural edges have a central
ridge, sending off on both sides sinuous ridges. The basal margins
of the alæ are not short compared with their upper margins, and
therefore the whole ala is not wedge-formed; and in this rather
important respect _C. barbara_ resembles _C. balænaris_ (Pl. 16,
fig. 3), and differs from _C. diadema_ (fig. 2). The lower edge of
the sheath does not seem to have projected freely,--in this respect,
also, resembling _C. balænaris_. From the basal margin of the alæ not
being narrow, and from the inner ends of the folded walls being, as
it would appear, also broad, I have little doubt that the cavity in
which the animal's body was lodged, resembled in shape that in _C.
balænaris_, the membranous basis being larger than the orifice of the
shell.
Opercular valves unknown.
_Summary._--This species is more nearly related to _C. diadema_ than
to the others; but in some points, just specified, it resembles _C.
balænaris_. The characters by which it differs from all the species
are, firstly, the more prominent transverse ridges on the external
surface of the shell, and more especially on the surfaces bounding
the outer sides of the cavities occupied by the epidermis of the
whale. Secondly, the character of the teeth, or rather ridges, along
the lines of junction between the transverse loops. And, thirdly, the
spaces between the radii and the plates on which the alæ rest, being
solidly filled up.
_Species Dubiæ._
The _Coronula bifida_ is an Italian tertiary species, so named by
Bronn, in his "Italiens Tertiär-Gebilde" (1831), p. 126. It is very
possible that this may be identical with _C. barbara_, but Bronn does
not seem to have been aware of the absolute necessity of giving minute
details in his descriptions of fossil cirripedes. The chief character
of _C. bifida_ is thus given:--"Eine tiefe Furche oder Spalte theilt
die Längenrippe von oben herab bis zur Hälfte, welche bei der sonst
ähnlichen _C. diadema_ entweder ganz fehlt, oder nur zuweilen kurz
angedeutet ist." Had it been stated that the longitudinal ribs were
divided from the middle down to the base, instead of from the top to
the middle, the description would have been intelligible to me, though
the character thus afforded would not have been of specific value, as
this dividing of the ribs occasionally occurs in all four species, and
is produced by the formation of new folds to the walls.
9. _Genus_--PLATYLEPAS. Pl. 17, fig. 1 _a_-2 _b_.
PLATYLEPAS. _J. E. Gray._ Annals of Philosophy, (new series), vol. x
(1825).
CORONULA. _De Blainville._ Dict. des Sciences Nat. (1824).
COLUMELLINA. _Bivona_ (fide _Philippi_). Nuovi generi, &c. di
Molluschi, Palermo (1832).
_Compartments six, each bilobed and inwardly produced, so as to form
six midribs, which support the outwardly convex, membranous basis._
_Distribution_, throughout the tropical and warmer temperate seas.
Imbedded in turtles, sea-snakes and manatee.
This small genus, consisting of three species (though I have named
only two), is a very natural one, yet closely allied to Coronula. Most
authors have united these genera, but in doing so they destroy two very
natural little groups. Platylepas, moreover, when all its characters
are considered, has as good a claim to be generically separated from
Coronula, as has Tubicinella,--a genus universally admitted. I shall
presently recur to this subject.
_General Appearance._--The most remarkable character, and which
gives a peculiar aspect to the whole shell, is that each of the six
compartments has a medial fissure or fold; the shell thus consisting
of twelve lobes. Each fold extends from the summit to the base of
the shell, and is produced inwards in the shape of a midrib (Pl. 17,
fig. 1 _a_). The membranous basis is supported by the basal edges of
the six midribs, and is thus rendered convex. As the midrib on the
rostrum would have interfered, if fully developed, with that part of
the animal's thorax which I have called the prosoma, it is of less size
than the other midribs; and from this same cause the whole rostrum is
pushed a little on one side, generally to the left, so that the shell
is not perfectly symmetrical. The midrib of the carina is less than
the lateral midribs. The orifice is oval, and is more or less plainly
indented by six angular points,--the worn-down summits of the six
midribs. The narrow elongated opercular valves stretch from one end of
the orifice to the other. The rostrum is much broader than the carina:
the lateral compartments are but little broader than the carino-lateral
compartments. The radii are narrow, though variable in width. The
general shape of the shell varies much in the same species, being
either much depressed, or rather steeply conical; either circular, or
more commonly oval. The (so called) epidermis is usually persistent
in the lower part of the shell. The shell itself is white. The basal
diameter of the largest specimen of _P. bissexlobata_ was nearly
three-quarters of an inch.
_Structure of the Parietes, Radii, &c._--The walls of the shell in _P.
bissexlobata_ are permeated by minute pores, and have exactly the same
structure as in Coronula; the pores being completed by the union of
ledges on the outer sides of the longitudinal septa, and the latter
have little knobs formed on them at each period of growth. In _P.
decorata_ the walls are solid. Each compartment, as already stated, has
a deep nearly medial furrow, the sides of which are closely pressed
together, thus forming the midribs, and thus differing from the furrows
in Coronula, which not being pressed together, form cavities on the
under side of the shell. The inner longitudinal margin of each midrib
is thickly coated by a layer of shell, and is thus rendered much more
prominent than it would have been if formed exclusively by the inward
folding of the wall. The upper part of each midrib forms a slight,
longitudinal ridge (fig. 1 _d_) on the sheath, having become encased
by the sheath during its downward growth. Owing to the small size of
the midrib of the rostrum, there is scarcely any ridge on the sheath
of this compartment. The sheath does not descend half-way down the
walls in _P. bissexlobata_, but further down in _P. decorata_; but
perhaps the thickened inner margins of the midribs may be considered as
the downward prolongations of the sheath, and on this view the sheath
descends actually to the basal membrane. The radii are narrow: their
edges have simple septa, with the interspaces filled up solidly. The
alæ have their sutural edges smooth.
The _Basis_ is membranous, and more or less convex. This convexity
is rendered possible by the support afforded by the basal edges of
the six midribs. The so-called epidermis or membrane which covers the
shell, is prolonged in six flaps under the midribs; and hence the
proper basal membrane has six deep indentations, receiving the above
flaps. The cementing apparatus is constructed on the same simple type
as in Coronula, as has been described in the Introduction.
_Scuta and Terga._--These are narrow and long, stretching the whole
length of the orifice of the shell. The two valves touch each other,
but are not properly articulated together: their upper surfaces
scale off in layers: they resemble each other very closely in shape
(fig. 1 _c_), being simply oblong. The membrane covering the valves
supports some very minute spines, 2/1000th of an inch in length. When
a portion of valve is dissolved in acid it is seen to be penetrated
by very numerous tubuli. The opercular membrane is rather thick: in
_P. decorata_ I found it occasionally double; in this species it
supports some very minute spines; it is attached about one third down
the sheath, of which the upper part is not marked (as in Balanus) by
regular concentric rings. The aperture between the valves, leading
into the sack, has protuberant lips, as in Coronula and Tubicinella.
The normal five bundles of muscles surrounding the sack, differently
from in the two genera just mentioned, do not thin out and spread
out downwards, but retaining their full thickness, reach the basal
membrane, to which they are attached.
_Mouth_: the parts present no difference from those in Coronula;
between the outer maxillæ we have the same small flattened mentum-like
projection; and in the mandibles, the same small intermediate teeth
between the second and third, and between the third and fourth main
teeth, as in that genus. The main teeth here are more plainly double
laterally. There are teeth on each side of the central notch of the
labrum. The cirri differ little from those of Coronula, but the
segments are not nearly so much flattened and broad, and the second and
third pairs are not so short. The segments of the sixth pair support
four pairs of main spines, with a few minute intermediate spines, but
with no intermediate tuft.
_Branchiæ_: these consist on each side (at least in _P. decorata_) of a
double fold, much less plicated than in Coronula; the fold nearest to
the animal's body is rounder, and not quite so large as the outer fold.
_Range and Habits._--The _P. bissexlobata_ ranges from the
Mediterranean and west coast of Africa to the West Indies, to the
north-east coast of Australia, and, judging from the name given by
Chenu, to California: the second species, _P. decorata_, inhabits the
Pacific Ocean; and the third imperfectly known species, was deeply
buried in the skin of a sea-snake, off Borneo. The _P. bissexlobata_
is attached both to turtles and to the skin of the manatee: some
specimens in the British Museum, attached to the latter, from the west
coast of Africa, were entirely buried in the skin, with the exception
of the operculum. The specimens of _P. decorata_, which I obtained at
the Galapagos Archipelago, were buried about half their depth in the
softer parts of the skin of the green turtle. I do not doubt that the
imbedment is effected in the same manner as in the case of Chelonobia.
_Affinities._--This genus, in general structure and habits, approaches
closely to Coronula; but it differs from all the species of that genus
in the following respects, which appears to me fully to justify its
generic separation:--The outline is often oval, instead of circular;
the compartments are not of equal sizes and shapes; each compartment
has here only one inward furrow or fold, and this has its sides pressed
closely together, and is encased, but not obliterated, by the sheath;
the rostrum is not quite medial; and the radii are narrow and nearly
solid. The opercular valves, again, are here more developed, and
their microscopical structure is different. The opercular membrane is
less constantly double; and, what is much more important, the muscles
of the sack are not here spread out, and show no tendency to become
rudimentary and lose their transverse striæ. The cirri differ only in
not being so broad and flattened. The branchiæ are not quite so much
developed. Finally, I may add, that in several of the above respects in
which Platylepas differs from Coronula, it approaches ordinary sessile
cirripedes.
1. PLATYLEPAS BISSEXLOBATA. Pl. 17, fig. 1 _a_-1 _d_.
CORONULA BISSEXLOBATA. _De Blainville._ Dict. des Sciences
Naturelles, tom. 32, (1824), Tab. 117, fig. 1.
PLATYLEPAS PULCHRA. _J. E. Gray_ (!). Annals of Philosophy, (new
series), vol. 10, (1825).
COLUMELLINA BISSEXLOBATA. _Bivona_ (fide _Philippi_). Nuovi generi di
Mollusch. (1832), Tab. 3, fig. 1.[126]
CORONULA CALIFORNIENSIS. _Chenu_ (!). Illust. Conch., Tab. 1, fig. 4.
[126] This memoir was published in the 'Effemeridi Scientifiche e
Litt. per la Sicilia,' according to the 'Bibliographia Zoologiæ et
Geologiæ,' by Agassiz and Strickland.
_Shell with the transverse lines of growth conspicuous: parietes
permeated by pores; sheath descending barely half-way down the
parietes._
_Hab._--Mediterranean, attached to turtles. River Gambia, attached
to manatee. Honduras, attached to manatee. Moreton Bay, lat. 27°
S., Australia, apparently attached to the dugong of that coast.
California(?) Mus. Brit., Stutchbury, and Cuming.
_General Appearance._--Shell generally much depressed, and broadly
oval or circular; sometimes steeply conical. Orifice oval, generally
not large. Surface rather plainly marked by closely approximate
lines of growth, which with the lobed outline gives to the whole an
elegantly sculptured appearance: occasionally the longitudinal ridges
formed by the parietal septa are distinct. Basal diameter of largest
specimen three quarters of an inch.
_Structure of Shell._--I have nothing material to add to the generic
description. The midribs are not so prominent as in the following
species, and hence the basal membrane is less convex. The origin
of the midrib as a fold is very plain. The sheath descends barely
half-way down the walls, and is a little hollow on its under margin,
on each side of the midrib. Beneath the sheath the parietes are
finely ribbed (fig. 1 _d_), but to a variable degree. I may here
remark, that in the specimens taken from manatee, on the coast of
Africa and at Honduras, the internal ribs extended further up and
were plainer, and the opercular valves seemed to be a little narrower
than in the other specimens, so that I at first suspected that they
were specifically distinct, but I could make out no other than these
small and variable points of difference.
_Scuta_: oblong (fig. 1 _c_), about twice as long as broad, with the
rostral end rounded, rather narrower than the other end, and curled a
little inwards. _Terga_, of nearly the same shape and nearly as long
as the scuta: the carinal end is rather more pointed than the scutal
end of the valve, and when viewed internally, the growing surface
of this end is seen to be bluntly pointed. In both valves the upper
layers of shell usually scale off.
_Mouth and Cirri._ The only differences, compared with the following
species, that I perceived, were that the hairs at the end of the
palpi were here rather longer, and the fourth tooth in the mandibles
rather less distinct. The rami of the first cirrus are very unequal
in length.
2. PLATYLEPAS DECORATA. Pl. 17, fig. 2 _a_-2 _b_.
_Shell with fine longitudinal ridges, ornamented in the lower part
by minute beads; parietes not porose: basal membrane equalling in
convexity the shell._
_Hab._--Pacific Ocean; Galapagos Archipelago; Lord Hood's Island, Low
Archipelago; Mus. Cuming, Darwin.
_General Appearance and Structure._--Shell oval, with the orifice
large. The walls are thick, and of less height from top to base than
in the last species. The surface (and this is the chief external
character) is marked by fine longitudinal ridges, each of which,
when closely examined, is found to be double (fig. 2 _b_), and near
the basis, where not abraded, to be ornamented with minute beads on
each side: these beads are largest on those ridges which border the
sutures. The parietes are not permeated by pores; on their inner
surfaces there are a few rather prominent but short ridges, instead
of the many finer ridges, as in _P. bissexlobata_. The sheath is of
great thickness; in young specimens layers scale off its summit, all
round the shell, as in Tubicinella; it is only slightly hollowed out
at its lower margin; it descends more than half-way down the walls.
The midribs, formed by the folded walls, are more prominent than in
the last species, and descend lower down; hence the basal membrane is
more convex than in the last species, for it projects downwards as
much as the shell projects upwards. In medium-sized specimens, the
midribs project inwards to a distance exceeding half the breadth of a
compartment. A less proportional length of the midrib, in an inward
direction, is formed by the inward folding of the wall, and a greater
length by the thickening of its inner longitudinal margin, than in
the last species. The basal edge of the inner end of the midrib is
smooth.
_Dimensions._--The average size of the numerous specimens which
I obtained at the Galapagos Islands was about .2 of an inch in
diameter, and I found none larger; but Mr. Cuming's specimens from
Lord Hood's Island are half an inch in diameter.
_Opercular Valves._--These closely resemble those of the former
species, with the exception that the carinal end of the growing or
under surface of the Tergum is much squarer.
_Cirri._--The only difference which I could perceive was, that the
rami of the first pair were not quite so unequal in length.
_Species Dubiæ._
3. PLATYLEPAS ----?
_Hab._--Imbedded in the skin of a sea-snake, taken off Borneo.
I am indebted to Dr. Gray for a single specimen of this supposed
species, but as it is very young and imperfect, wanting the opercular
valves and cirri, I do not choose to name it. The shell presents all
the usual characters of the genus; the rostrum, I may remark, being
pushed to the left side. The parietes are permeated by pores of
considerable size, which shows that the species is distinct from _P.
decorata_. On the inner basal surfaces of the walls, there are two or
three very distinct ridges on each side of the midrib; and this fact,
together with the size of the parietal pores, makes me suspect that it
is not an immature specimen of _P. bissexlobata_.
10. _Genus_--TUBICINELLA. Pl. 17, fig. 3 _a_-3 _c_.
TUBICINELLA. _Lamarck._ Annales du Museum, tom. 1 (1802), Tab. 30,
fig. 1.
CORONULA. _De Blainville._ Dict. des Sciences Naturelles, (1824).
_Compartments six, of equal sizes; shell sub-cylindrical, wider at the
top than at the basis, belted by several large transverse ridges._
_Hab._--Southern Pacific Ocean, Western South America, New South
Wales, Cape of Good Hope; imbedded in whales, and often associated
with _Coronula balænaris_.
This genus is closely allied to Coronula, and perhaps De Blainville was
right in uniting them. But Coronula, as it now stands, is so natural
a genus, that it seems a pity that a form so entirely different in
general aspect, as Tubicinella, should be forced into it. The main
difference between these genera consists in the walls being here not
folded, in the simpler radii, and in the general shape of the shell and
of the included animal's body; but there are many other minor points
of difference. The most novel character in Tubicinella consists in
the shell being lined almost close down to the basis by the opercular
membrane. In the opercular membrane thus forming a long tube, and in
the general shape of the animal's body, we shall presently see that
Tubicinella is closely related to Xenobalanus. In numerous other
respects Tubicinella is almost equally allied to the latter genus,
to Platylepas, and to Coronula. Finally, several points of structure
indicate that Tubicinella may be considered as a Coronula, with the
shell much simplified in structure.
1. TUBICINELLA TRACHEALIS. Pl. 17, fig. 3 _a_-3 _c_.
LEPAS TRACHEALIS. _Shaw._ Nat. Miscell. (1789-1813), vol. 17 (1806?)
tab. 726.
---- TRACHEÆFORMIS. _Wood._ General Conch. (1815), tab. 4, fig. 1-3.
TUBICINELLA TRACHEALIS. _J. E. Gray._ Annals of Philosophy, (new
series), vol. 10, (1825.)
---- MAJOR ET MINUS. _Lamarck._[127] Annales du Mus. Nat., tom. 1
(1802), Tab. 30, fig. 1-2.
---- BALÆNARUM. _Lamarck._ Animaux sans Vertèbres, (1818).
---- ---- _Chenu._ Illust. Conch. (Plate).
---- ---- _Sowerby._ Genera of Recent and Fossil Shells (Plate).
---- LAMARCKII. _Leach._ Encyclop. Brit. Suppl., vol. 3 (1824), Pl.
57.
CORONULA TUBICINELLA. _De Blainville._ Dict. des Sciences Nat., tom.
32, Pl. 117, fig. 5 (1824).
[127] It may be observed that I have here broken through the great
law of priority; for it appears to me too grossly incorrect to
retain the specific name either of _major_ or _minus_ in a genus
including a single species. Lamarck himself seems to have been
of this opinion, by giving, in 1818, the new specific name of
_balænarum_; but Shaw's name of _trachealis_ has the clear right to
priority, if _major_ or _minus_ be rejected.
_General Appearance._--Shell elongated, sub-cylindrical, with the
upper end rather wider than the lower, and therefore widening in
a direction the reverse of that usual with sessile cirripedes.
The shell is often a little bent to one or the other side: it is
surrounded by from two or three to about ten very prominent, strong,
blunt ridges or belts, placed at rather irregular distances from
each other. The surface is finely striated longitudinally. The six
compartments are of nearly equal sizes and shapes. In full-grown
specimens the parietes are not wider at the base, and often they are
even a little narrower, than at the summit of the shell: in young
specimens the parietes do widen a little downwards. The radii are
narrow; but in young specimens they are proportionably much broader
(Pl. 17, fig. 3 _b_) than in old specimens. The whole compartment,
including the radius and wall, is always a little wider at the summit
than at the base, in accordance with the shape of the whole shell.
The operculum consists of four nearly equal-sized, similar valves,
projecting above the upper end of the shell, which is always broken
and jagged: the valves are united to the sheath by a very thick,
much folded membrane. The aperture leading into the sack is bordered
by very prominent lips, projecting above the opercular valves; the
latter have their upper layers always scaled off. The shell is
imbedded in the whale's skin up to the level of its operculum. The
largest specimen which I have seen was barely one inch (.95) in
diameter at the summit, and 1.5 in length; the longest specimen which
I have seen scarcely attained a length of one inch and three quarters.
_Structure of the Shell._--The parietes are thin, and if the sheath
(which extends to near the basis) be removed, they are rendered
extremely thin. They are formed by an outer and inner lamina, united
by fine longitudinal septa, projecting at the basis beyond the
laminæ. The pores thus produced (which in a transverse section are
oblong in outline), run up to the summit of the shell, and are not
filled up by shelly matter; but I presume that the included tubular
threads of corium are protected, at the broken upper end of the
shell, by transverse membranous septa. The outer lamina of shell,
as in Coronula, is formed, though obscurely, by the union of ledges
projecting from the longitudinal septa. The circular prominent belts,
surrounding the shell, are formed by the longitudinal septa, at
certain, irregular and rather distant periods, growing outwards; the
wall at each belt being increased to nearly twice its thickness in
other parts. At each belt the threads of corium within the parietal
pores lend off minute branches to supply the thickened wall. These
belts, which continuously surround the shell, correspond (as is best
seen in young specimens), with the little knobs or beads, which, in
Coronula (Pl. 16, fig. 4), rise separately, and not quite regularly,
on the longitudinal parietal septa, and which, I believe, are formed
at every successive period of growth; here they are much larger,
stand in straight transverse rows, become confluent, and are formed
only at occasional intervals. The whole external surface of the shell
is covered by membrane, stronger and more persistent than is usual
with most cirripedes.
Internally the sheath extends almost to the basis of the much
elongated shell, and terminates in a slight shoulder: it is divided
as in common Balanidæ, and differently from in Coronula, into zones
of growth, but these are very broad; at the upper end of the shell,
which, as will hereafter be explained, is always breaking away,
the sheath readily yields along the oblique planes, which separate
the zones of growth and dip outwards: a similar but less strongly
marked structure occurs in Platylepas, and in no other genus. We
shall presently see that the sheath presents a much more anomalous
character, in being lined down to its basal edge by the innermost and
last-formed layers of the opercular membrane.
_Radii._--The radii are narrow. The belts which surround the shell
are prolonged, with slightly diminished prominence, across the radii,
their formation being simply due to the radii being here thicker than
in other parts. It can be seen more plainly in Tubicinella, than in
other Balanidæ, that the membrane externally investing the shell,
splits along the radii during the diametric growth of the shell,
and is continually repaired and added to along these lines by new
longitudinal slips of membrane. The radius consists (as usual) of an
inner and outer lamina, which latter does not extend quite to the
line of suture--a slightly gaping fissure being thus left. The two
laminæ are connected by septa, which are not denticulated, but near
the outer lamina bi- or tri-furcate, and the ends of the branches
thus formed spread out, forming a sort of outer scalloped lamina, in
advance of the true outer lamina. The fine threads of corium running
between these septa, do not spring, as in all common cirripedes, from
a fillet of corium occupying the actual suture, but from two nearly
circular threads of corium occupying two tubes, which run along the
line of junction between the radius and the compartment whence it
springs. In Coronula alone we have a nearly similar structure; for
the fine threads of corium occupying the _proper_ radius, spring from
a _single_ very minute tube (Pl. 16, fig. 7, _d′_), occupying the
same position with the _two_ tubes in Tubicinella. I may further add,
that the structure of the proper radius in Coronula is precisely the
same as here just described, but being on so very minute a scale, I
did not there describe it so carefully as I have here done.
_Alæ_: these are only remarkable from their extreme thinness; for
they are not thicker than the inner lamina of the radius. Their
sutural edges are quite smooth. Forming part of the sheath, they
extend down close to the basis of the shell; where, instead of, as in
general, ending abruptly in a rectangular shoulder, they slope off
into their own compartment.
_Basis._--The basal membrane is complicated, owing to the shell, when
full-grown, barely, or not at all, increasing in diameter, and, in
consequence, membrane after membrane, each with its own cement-ducts
attached to it, are thrown down one nearly over the other. In the
Introduction (p. 143--Pl. 28, fig. 3), I have fully described the
cementing apparatus, which is very curious from one of the ducts
always having a loop with two spurs projecting from it. The basal
membrane does not equal in diameter the base of the shell, for the
membrane externally covering the walls is inflected inwards all round
for a considerable width, and is then united to the basal membrane:
in Coronula, the basal membrane extends only under the internal
cavity of the shell, and not under the folded walls, and therefore
presents a somewhat analogous structure.
_Opercular Valves._--The scuta and terga are nearly of the same size
and shape: they are mitre-formed, and higher than broad. They do not
fill up the orifice of the shell. The scutum is a little larger than
the tergum, and rather less symmetrical, the rostral corner of the
valve being a little produced. There is no hollow or crest for the
adductor muscle, which is small. In the tergum there is no trace of
a spur. The two valves are not articulated together, but standing
close to each other are united, as well as the scutum to the scutum,
and tergum to the tergum, by thick, brown, tough, yet soft membrane,
in layers continuous with, but differing in appearance from, the
surrounding opercular membrane. The layers of shell, forming the
valves, are thick, and only the three or four lower layers are
usually preserved, the upper ones having symmetrically scaled off,
leaving snow-white surfaces. Owing to the thickness of the successive
shelly layers, and to the circumstance of each new layer being but
very little larger than the last, the scaling off of the old upper
layers is a quite necessary process; for otherwise the orifice into
the sack would have been encumbered and almost closed by four long,
slightly tapering points, prolonged upwards from the basal layers
that form the four existing valves. The same scaling off process
takes place in Platylepas, and amongst pedunculated cirripedes in
Lithotrya. Microscopical examination does not exhibit any fine
spines on the membrane investing the valves, or any tubuli in the
shelly layers after their dissolution in acid: in this respect the
valves resemble those of Coronula. The summits of the valves project
freely for about a third of their own height, above the level of
the membrane by which they are surrounded. The orifice leading into
the sack is bordered by very protuberant lips, standing up even
considerably above the upper freely projecting portions of the valves.
The _Opercular Membrane_, connecting the valves and the top of the
shell, is thick and tough, and deeply folded in concentric wrinkles.
As in Coronula, it consists of two or three separate membranes (each
composed of many laminæ) one over the other, united to successive
shelly layers of the opercular valves. As the upper shelly layer
scales off, the membrane attached to it is likewise thrown off. The
innermost laminæ of the last-formed opercular membrane extend down,
closely attached to the sheath, to its basal edge, and therefore
nearly to the basis of the shell; the outer and older laminæ, all
closely attached one within the other and to the sheath, extend to
a less and less distance downwards; consequently, the animal's body
is enclosed in a tube, thinning out downwards, formed by the laminæ
of the successive opercular membranes, surrounded outside by the
shell: only in the following genus, Xenobalanus, shall we meet with a
nearly analogous structure. As the shell of Tubicinella increases in
diameter, from the growth of the radii, the opercular membrane lining
the sheath is necessarily split along the six lines of suture, in the
same manner as is the membrane externally investing the shell; in a
like manner, also, it is repaired and added to by new longitudinal
slips of membrane. Of this structure, in the opercular membrane, I
have seen no other instance; for in most genera the old opercular
membrane is moulted, and a new and larger one formed at each period
of growth; in Coronula, in which the opercular membrane is likewise
for a time persistent, it does not run far down the inside of the
shell, and each new membrane is formed large and extensible, so as
to allow, without splitting, of some increase in the diameter of
the shell. The opercular membrane at the summit of the shell, in
Tubicinella, is folded in concentric lines, and so deeply, that
the basal edges of the opercular valves are generally hidden: this
folding arises partly from each last deposited and innermost membrane
being originally formed slightly folded, but chiefly from the rapid
downward growth of the shell, and the consequent downward movement of
the whole animal's body, together with the opercular valves to which
the body is attached, and this necessarily tends to wrinkle and fold
the opercular membrane. Owing to the opercular membrane extending far
down inside the shell, and being firmly attached to the sheath, as
the upper part of the shell breaks away and disintegrates (which we
shall presently see is constantly taking place), small particles of
shell are left adherent to the circumferential and folded parts of
the opercular membrane; and this at first much perplexed me.
_Muscles of the Sack_: these extend down almost to the base of the
shell, but in the lower part they spread out and become thin and very
irregular, not even corresponding on the opposite sides of the body.
The fasciæ in the upper part show very distinct transverse striæ, but
lower down these become either obscure or entirely deficient. In all
these characters the muscles of Tubicinella and Coronula resemble
each other. The rostral depressor muscles of the scuta consist each
of four small bundles of fasciæ; the lateral depressores run not
quite straight down, but in a curved course towards the carinal end
of the sack: the tergal depressores are proportionally smaller than
in ordinary sessile cirripedes, but they project and form two crests
(with some fasciæ between them), which support the Branchiæ. The
membrane lining the sack, I may here mention, is unusually strong.
_Branchiæ._[128]--These are enormously developed; the two together
covering two thirds of the area of the sack. Each consists of two
folds, both deeply plicated. They are attached longitudinally to
the two crests, including and formed by the muscles running from
the terga to near the basis of the shell. The branchiæ are likewise
attached transversely to the sack, under the basal margins of the
terga.
[128] These have been described by Professor Owen in the second
volume of the 'Descriptive Catalogue of the Museum of the Royal
College of Surgeons.'
_Mouth._--The labrum is very finely hirsute, without teeth; the palpi
have a short row of moderately long spines along their exterior
basal margins. The mandibles have four rather narrow, sharp teeth,
which (excepting the first) have double points: between the second
and third, and again between the third and fourth teeth, there is a
single small intermediate tooth: the inferior angle is irregularly
pectinated. The maxillæ are small; there is a small notch beneath the
two upper great spines, and a second notch near the inferior angle.
Between the outer maxillæ, there is a square-topped mentum. Hence we
see that the mouth in all its few peculiarities, resembles that of
Coronula.
_Cirri._--These are short, with short and broad segments protuberant
in front. The pedicel of the first cirrus is very broad, and
exteriorly clothed with fine hairs: its rami are slightly unequal
in length. The second and third cirri are very short. The three
posterior pairs are remarkable from the pairs of main spines being
placed so close one under the other, and in an oblique direction,
that at first they appear to form a single crowded transverse row:
the dorsal tufts are rather large.
_Body._--The body is remarkable from its nearly vertical position,
and from the much elongated pyramidal form of the prosoma, extending
down nearly to the bottom of the sack. The membrane investing the
prosoma, presents a few circular folds, falsely appearing like
articulations. The œsophagus enters the stomach rather obliquely.
With respect to the generative system, I have only to remark, that
the vesiculæ seminales are of great length, and convoluted to a
remarkable degree. The ovarian cæca, form a thick layer at the
bottom of the sack; they do not appear to extend up the shell round
the sack. The only other point, which I shall here mention, is that
beneath the basal articulation of the first cirrus, there is a
longitudinal swelling, ending in a freely projecting point, .06 of
an inch in length; at first, I thought, that we here had a rudiment
of a filamentary appendage like those found in several Lepadidæ;
but closer examination showed an orifice at the apex, leading into
the acoustic meatus, in which the singular, wrinkled, heart-shaped
acoustic vesicle, mentioned in the Introduction, hangs suspended.
Alongside the freely depending point, with an orifice at its end,
there is a smaller upward projecting point, without any orifice, but
hollow within and lined by corium; I believe it opens internally into
the acoustic meatus.
_Attachment and general Growth of Shell._--All the specimens which
I have seen have been attached in groups. They are buried up to
the level of the operculum in the whale's skin; and their summits,
I suspect, lie even beneath the general surface of the body of
the whale. It is certain that the shells grow much at their basal
ends. As in the case of Coronula, the flat membranous basis does
not actually penetrate the skin; but the general pressure of the
whole group of shells seems to push inwards the skin of the whale,
and directly beneath each shell the formation of new epidermis
is apparently checked. Between the shells, however, though close
together, the epidermis continues to be formed, and is pushed upwards
between them, in the same manner as it is forced into the flattened
cavities on the under side of the shell of Coronula.
The manner in which a full-grown shell assumes its ordinary shape,
at first appears very perplexing:[129] it has to change from a
cylinder, at first probably not much above the 1/50th of an inch in
diameter, to a cylinder nearly one inch in diameter: and this is
not effected by the growth of the radii, for the radii never reach
the basis, and the basis of course has to increase in diameter
like the rest of the cylindrical shell. The radii serve only to
keep the summit of the shell wider than the basis, which is the
natural shape of this species; and in large-sized specimens, this
purpose is sometimes aided by the parietes during their downward
growth decreasing slightly in width. In ordinary nearly full-sized
specimens, the parietes are of the same width at the top and bottom,
but in some large-sized specimens, as just stated, they even become
narrower towards the bottom; as they grow only at the bottom, one
does not at first see how they can ever increase in width, or how
the older shells can have acquired their present diameter. But an
examination of young specimens, from .1 to .3 of an inch in diameter,
at once serves to show how the shell attains its full size and shape:
for here the parietes are all found to increase downwards sensibly
in width, though at a much slower ratio than in other sessile
cirripedes; in larger, but not full-grown specimens, a similar
increase can by care be detected: hence by long-continued growth at
the base of the shell, with the removal of the upper part, a young
Tubicinella of small diameter will be converted into an old one of
large diameter, retaining during all the time its sub-cylindrical
form, with its summit rather broader than its base. With respect
to the removal of the upper part of the shell, this seems almost
constantly going on, for the summit of every specimen invariably had
a freshly broken aspect. The peculiar structure of the sheath, which
is the strongest part of the shell, namely, its division into oblique
layers, separable by a slight force, doubtless is subservient to the
repeated breakage of the summit. In some species of Tetraclita and
of Balanus, gradual disintegration of the upper part of the shell is
a necessary element in the growth of the animal, in order that the
orifice may increase in size, and here we have mechanical breakage
equally necessary.
[129] I am indebted to Dr. J. E. Gray for calling my attention to
this subject, and explaining to me several points.
Some curious results follow from the peculiar growth of Tubicinella
just described. At Plate 17, fig. 3 _b_, we have a careful drawing of
a lateral compartment, together with its radius, (which latter does
not here concern us), taken from a shell .2 of an inch in diameter.
The two protracted dotted lines show the form which this compartment
would have assumed, if it had continued growing downwards at the
same rate of increase in width as hitherto. But the increase in
width always seems to become less and less as the shell grows older;
hence the dotted lines, representing the wall after long-continued
growth, ought to have been drawn diverging or widening still more
slowly than they do. The lateral compartment in fig. 3 _a_ is the
exact size of the compartment of a large specimen nearly one inch in
diameter; in this specimen the parietes, far from increasing in width
downwards, had commenced, as is represented, decreasing. Compartments
of all intermediate sizes between those figured at 3 _a_ and 3 _b_
can easily be shown in different specimens. From these facts we may
safely infer, that if the whole growth of the compartment 3 _a_ had
been preserved, instead of its upper end having been continually
chipped away, it would have had even a more tapering form than that
represented by the whole and dotted lines in the two figures, and
would have exceeded six inches in length! this of course being also
the length of the whole shell. The young Tubicinella, of which 3
_b_ is a compartment, was imbedded in the whale's skin nearly up to
the level of its operculum; if it had lived, it would no doubt have
grown to the length just specified, viz., above six inches, but as
all the growth is at the lower end, the bottom of the shell, it might
be thought, would necessarily have become buried in the whale's skin
to this same depth; and the summit of the shell, on this same view,
would have been buried to a depth by as much less as the height or
length of the old shell itself, namely, by about one inch and a half
less than the six inches. As far as I can judge from an examination
of several large groups of full-grown specimens, preserved in their
imbedded condition, the summits of the shells seem always to lie a
little beneath the surrounding level of the whale's skin, but not
nearly to the extent here just inferred. Nor can I believe that the
epidermis of the whale had ceased being formed under these specimens,
whilst it had gone on being formed all round them, to the thickness
of between four and five inches, and that it had subsequently
disintegrated to this same thickness,--which processes would account
for the summit of the shell being still on nearly a level with the
surface of the whale. The view which seems to me most probable, is,
that the rapid downward growth of the shell, besides indenting the
whale's skin, at the same time slowly pushes the whole shell out
of the skin, and thus continually exposes the summit to the wear
and breakage which seems to be necessary for its existence. On this
view, the very peculiar form of Tubicinella, which is retained during
life, namely, the slightly greater width at top than at bottom,
is beautifully explained, viz., for the sake of facilitating the
protrusion of the shell; for the ordinary conical shape of sessile
cirripedes, with the apex upwards, would have rendered the pushing
out of an imbedded shell almost impossible; on the other hand, we can
see that the likewise very peculiar, concentric, prominent belts may
be necessary to prevent too easy protrusion.
_Fossil Species._--I do not believe that this genus has hitherto
been found fossil. The _Tubicinella maxima_ of Ch. Morren, said to
have been found (see Bronn, Index Palæontologicus) in the chalk of
Belgium, I have good reason to believe does not really belong to this
genus.
11. _Genus_--XENOBALANUS. Pl. 17, fig. 4 _a_-4 _c_.
XENOBALANUS. _Steenstrup._ Videnskabelige Meddelelser. Aaret, 1851.
SIPHONICELLA (sine descript.) _Darwin._ Monograph on the Lepadidæ, p.
156 (1852).
_Shell almost rudimentary, star-formed, composed of six compartments,
with a long peduncle-formed body rising from the middle: opercular
valves none._
_Hab._--North Atlantic Ocean, attached to Porpoises; Mus. R. T. Lowe,
Steenstrup.
This Cirripede, in appearance the most anomalous of its family, has
affinities distinctly pronounced. Four years ago the Rev. R. T.
Lowe sent me some specimens, which he had obtained from a porpoise
between Madeira and England; and I named them in MS. Siphonicella,
from their relationship to Tubicinella,--a fact which I mention only
because Sir C. Lyell has alluded to this genus under the above name
(without any description), in his anniversary address to the Geological
Society, as have I, in my volume (p. 156) on the Lepadidæ. Since that
time Professor Steenstrup has described and named the genus, fully
recognising its place and affinities, and has most kindly sent me a
magnificent group of specimens.
This genus singularly resembles, in general appearance, some of
the pedunculated Cirripedes, so much so that in the specimens sent
me by Mr. Lowe, in which the almost rudimentary shell was, from
disintegration and its deep imbedment, not plainly visible, I did not
in the least doubt that I was examining a new genus of Lepadidæ. I may
mention, as a proof how truly all the parts and organs are correlated
in Cirripedes, that I was at first in despair when I found a species
to all appearance pedunculated, with its labrum not bullate, its palpi
of large size, its third pair of cirri totally unlike the fourth and
succeeding pairs, and with only a single layer of muscles round the
peduncle; but when, in addition, I found that there were branchiæ,
and that these were double, I felt convinced that I was dissecting a
disguised sessile cirripede, and that its true place was near Coronula:
soon, I found the imbedded and almost rudimentary shell, of which a
mere fragment would equally well have declared the true position and
relationship of the whole animal. Though Xenobalanus, in external
aspect, is so completely masked, yet in its habits, namely, in living
attached on Cetaceans, as in its essential structure, it displays its
real affinities. In the course of the following description, it will
be seen that in the shell, the affinity is almost equally close to
Coronula and Platylepas, but that, considering the whole animal, the
affinity is somewhat closer to Tubicinella. Xenobalanus may indeed
be described as a Tubicinella without opercular valves,--with the
opercular membrane thickened down to the basis,--and with the shell,
excepting the few last-formed basal zones of growth, almost wholly
removed by the breakage of its upper end; this remnant of a shell,
however, presenting some strong points of resemblance to Coronula.
1. XENOBALANUS GLOBICIPITIS. Pl. 17, fig. 4 _a_-4 _c_.
XENOBALANUS GLOBICIPITIS. _Steenstrup._ Videnskabelige Meddelelser
fra den Naturhist. Forening i Kjöbenhavn, for Aaret, 1851. Tab.
3, fig. 11-15.
_General Appearance._--The shell is in an almost rudimentary
condition, and appears like a small white irregular star, imbedded up
to its top in the skin of the porpoise. Out of this thin, star-shaped
shell, a cylindrical, flexible, peduncle-formed body springs, which
forms the main part of the animal; it is narrow where coming out of
the central cavity of the star, but soon acquires its full diameter;
at the upper end it has a reflexed hood, and hence is broader,
and this has the appearance of forming a capitulum, like that of
a pedunculated Cirripede. This pseudo-capitulum is formed by a
membranous reflexed collar or hood, which is very narrow at the lower
end of the orifice, close under the mouth, and becomes wider and
wider towards the upper and carinal or posterior end of the orifice;
hence the lower reflexed edge of the hood is only slightly oblique or
even nearly transverse. The orifice leading into the sack is large,
and nearly in the same straight line with the peduncle; it is a
little hollowed out in the middle at the upper end, and on each side
of this medial hollow, there is a small rounded projection or horn,
not perforated, but hollow, as may be seen by turning up the hood and
looking at its under side. These two little horns curiously bring to
mind the ear-like appendages in _Conchoderma aurita_ (_Otion_), but
these latter are perforated, open into the sack, and point outwards.
The peduncle-formed body answers, as we shall presently see, to the
main part of the shell in Tubicinella, and the hood, as it would
appear, to the lips of the sack-aperture, which project between its
scuta and terga; of these valves there is not here a trace. The
whole surface is smooth, and is formed by rather thin membrane, of
an orange colour; but from the colour of the underlying corium, the
whole appears of a dark chocolate red, the reflexed hood being rather
lighter coloured. It is singular how closely the colour resembles
that of some dark varieties of the above-mentioned _Conchoderma
aurita_, and likewise of _Anelasma squalicola_, both pedunculated
Cirripedes, having oceanic habits, and destitute, to a remarkable
degree, like Xenobalanus, of shelly valves.
The largest specimen which I have seen was very nearly two inches in
length: in this specimen the star-shaped shell measured, from extreme
point to point, nearly a quarter of an inch in diameter, but the
internal cavity only about one eighth of an inch. This latter measure
gives also the diameter of the peduncle, where coming out of the
shell; the diameter just beneath the hood, was in this same specimen
rather more than a quarter of an inch, and therefore greater than the
diameter of the points of the shell. The depth of the shell from the
upper rim to the basal membrane, in one specimen which I measured,
was only one twentieth of an inch, and this specimen had its
pseudo-peduncle one inch and three quarters in length, consequently
thirty-five times as long as the shell was deep.
_Structure of Shell._--The almost rudimentary shell (fig. 4
_b_) consists of a small, thin, six-rayed disc, formed of six
compartments, each of which, instead of being outwardly convex, as
in ordinary Cirripedes, is deeply bowed inwards. The narrow sutures
(_s s_) separating the six compartments, run along the middle of
the six rays, each ray being composed of the bowed ends of the
walls of the adjoining compartments. The rays are a little curved
towards the carinal end of the shell. It is remarkable that the
rostrum is smaller and less deeply folded inwards than the other
compartments, and the lateral compartments are a little smaller
than the carino-lateral compartments, which is exactly the reverse
of what is usually the case. Only about four zones of growth have
been preserved in any specimen, and consequently the shell is very
nearly of the same diameter at the top and bottom; for the upper end
of the shell is rapidly removed, as in Tubicinella, by the scaling
off of the upper rims of the sheath, and by the disintegration of
the walls. The zones of growth are commonly not piled exactly over
each other, but rather obliquely, as represented in fig. 4 _b_. Each
zone projects, forming a prominent, sharp, toothed ridge round the
shell. In Coronula (Pl. 16, fig. 6) and its allies, the outer lamina
of the wall is formed by the union, a little above the basal margin,
of ledges running along the sides of the longitudinal septa. In
Xenobalanus (Pl. 17, fig. 4 _c_) similar ledges are less perfectly
joined, and apertures seem always to be left in transverse rows
under the transverse toothed ridges, which latter are best seen
in fig. 4 _b_. The apertures, of course, are covered by membrane.
The transverse ridges are surmounted by knobs arising from the
longitudinal septa; and the knobs themselves are capped by other
little heads, which are not represented in the drawing. Owing to
these projections, and to the prominence of the transverse ridges
and of the longitudinal septa, the external membrane is attached so
firmly to the shell that even with the aid of caustic potash it can
hardly be separated.
The internal cavity of the shell is small: it approaches a hexagon
in shape, with the rostral side very short, and the lateral sides
curved inwards. It is lined by a rather thick sheath, which descends
very near to the basal membrane; the sheath is divided into very
distinct, successive convex zones of growth. The external membrane
of the pseudo-peduncle is attached with remarkable strength to
these rib-like zones of the sheath. The alæ (_a_ in fig. 4 _b_) are
represented by mere angular shoulders, received into very slight
notches, and placed at the inner ends or entrances of the double
walls, or rays as I have called them. With respect to the radii, they
also are in an extremely rudimentary condition; but a thin layer of
shell, apparently continuous and homologous with the sheath, extends
from the sheath along both sides of each ray, and on the rostral side
(whence the radius ought to arise), about half-way from the end of
the ray, gives rise to a projection or ridge (_d_, in fig. 4 _b_)
which runs from the top to the base of the shallow shell. From this
longitudinal ridge, septa, parallel to the basis, extend to nearly
the extremity of the ray or double wall. These represent the radii;
but they never grow, so as to increase the diameter of the shell.
These radii evidently correspond to the additional or pseudo-radii
in Coronula, which in that genus lie between the parallel, as here,
and folded parietes. Of the true radius, having the same thickness
as the paries, I here saw only traces in an internal, very slight,
longitudinal ridge running up the shell, close to the outer extremity
of each ray or double wall.
The membranous _Basis_ is united all round to a rather wide flap of
membrane which is inflected from the outer surface of the shell. The
cement-glands appear to be mere enlargements of the cement-trunks,
which latter extend in two nearly straight and parallel lines. From
each gland two cement-ducts proceed, one of which runs parallel to
the cement-trunk.
The pseudo-peduncle, forming the main part of the animal, has, as far
as external appearance is concerned, been sufficiently described.
The part forming the hood apparently answers to the protuberant lips
of the operculum, and the lower part to the sub-cylindrical shell of
Tubicinella; both shell and peduncle in the two genera being wider
at top than at bottom. If in imagination we chip away (an action
always in progress) the whole upper part of the shell of Tubicinella,
leaving only two or three zones of growth at the base, we shall
convert it into a Xenobalanus, with every internal part and organ
occupying the same relative position: for it should be borne in mind
that the shell of Tubicinella is lined close down to the basis by
the opercular membrane, and this is strictly comparable with the
outer membrane of the pseudo-peduncle of Xenobalanus. The body, as
in Tubicinella, is attached in a vertical position, with the longer
axis of the thorax and of the much elongated prosoma extending in
the direction of the longitudinal axis of the pseudo-peduncle. The
point of attachment of the body to the lower part of the hood is
much elongated, but presents the usual muscles running to near the
base of the labrum and embracing the prosoma. The adductor scutorum
muscle is well developed, and is placed close beneath the mouth,
where the collar or hood commences folding over: hence, no doubt, the
lower end of the large and long orifice into the sack can be closed
by the adductor; but the upper end probably can only be blocked up
by the outer surfaces of the curled cirri. The scuta, I may add,
if such had existed, would certainly have covered the point where
the adductor muscle is attached to the exterior membrane. The sack
extends down to within the almost rudimentary shell: the tunic lining
it is unusually strong; indeed, in the reflexed hood-like portion,
it is as strong as the properly external membrane. The latter seems
to be moulted in large strips, and not in a single piece, like the
opercular membrane in Balanus: just above the shell, fragments of
three or four of these outer coats are retained by their very firm
attachment to the sheath. The two layers of corium, lining the outer
membrane of the pseudo-peduncle and the inner tunic of the sack,
instead of being, as usual, united almost continuously together,
stand some little way apart, and are connected by longitudinal septa;
hence, in a transverse section, especially of the lower part, the
sack is surrounded by an irregular ring of square tubes of corium.
The muscles surrounding the sack and imbedded in the inner fold of
corium, are very thinly spread out; they branch, and even sometimes
cross each other; they are more numerous at the carinal and rostral
ends, but certainly cannot be said to form six (or five) bundles, as
in all other sessile cirripedes. Some of the fasciæ extend down to
the very basis, and some up to the summit, to near the two little
horn-like projections. I could not perceive any transverse striæ on
these muscles. Altogether, they are very weak, and cannot have much
power in moving the whole peduncle-like body.
_Branchiæ._--These are largely developed: they are attached to two
approximate, longitudinal, fleshy crests, which extend more than
half-way down the sack, along the carinal margin. Each branchia is
double, the two folds being united where attached in a transverse
line across the sack, on a level with the attachment of the body.
The inner fold is much smaller than the outer; not extending half so
far down the sack, and not extending so far transversely; it is also
hardly at all plicated. The larger and plicated fold extends down
considerably below the lower end of the prosoma, and altogether fully
equals one third of the entire length of the animal, measured from
the shell to the summit of the orifice. Both folds are formed of very
delicate membrane.
_Mouth._--Labrum unusually prominent, as measured from its basal
margin to the crest, which is but slightly notched, hairy, and
without teeth. Palpi broad, heart-shaped, clothed on their inner
sides by a thick brush of spines, which here, as on the other
gnathites and cirri, are almost all doubly serrated. On the outer
margin of the palpi there are a few longer spines. Mandibles villose,
with five teeth, of which the fifth is very small and of irregular
shape: the inferior angle is broad and pectinated. There are no
intermediate teeth between the second, third, and fourth teeth, as
in the three foregoing genera. Had I not known that the lower main
mandibular teeth were always laterally double in the Balaninæ, and
had I not observed how obscure this structure was in Coronula and
Tubicinella, I should have overlooked the merest vestiges of double
teeth in the present genus; indeed, in some specimens the teeth
seemed to be absolutely single. The maxillæ are villose: their edge
exhibits a trace of being notched under the two great upper spines.
The outer maxillæ are bilobed, but not very plainly: between these
organs there is no little prominent mentum, as in the three previous
genera.
_Cirri._--The cirri are short, particularly the three anterior pairs.
The segments in all are singular, from being so much compressed, so
short, and of such great breadth; they are protuberant in front.
In the second and third cirri, the broad _lateral_ faces of the
segments, with the exception of the posterior face of the posterior
ramus, are almost bare of spines. In the three posterior pairs of
cirri (Pl. 29, fig. 6), the segments are protuberant in front, and
support three pairs of short thick spines, with an intermediate tuft;
the dorsal tufts are unusually small: their pedicels are remarkable
from the upper segment, and the upper part of the lower segment,
being produced into a rounded protuberance, dotted with spines: I
have met with a similar structure only in _Scalpellum vulgare_. In
the third pair (fig. 5) there is only a trace of this structure; and
in the second pair the anterior margin of the pedicel is straight,
and clothed with three tufts of bristles. The pedicel of the first
cirrus is very broad, and clasps the mouth.
The cirri and mouth are dark chocolate red, like the outside of the
animal and the upper part of the sack. The thorax is redder and
paler. The four posterior articulations of the thorax are straight
and transverse; the next segment, or that corresponding with the
second pair of cirri, is slightly inflected, in the usual way,
towards the prosoma. The prosoma is pale coloured, extraordinarily
elongated, and bluntly pointed; it extends down (see the dotted
outline in Pl. 17, fig. 4 _a_) about one third of the length of the
whole animal. The orifice leading into the acoustic sack forms a
freely depending little point beneath the basal articulation of the
first cirrus. The stomach in the uppermost part is deeply and closely
plaited longitudinally, but has no cæca; it runs down (externally
coated, as usual, by the testes) to the lower point of the prosoma,
and is then doubled back on itself, so that it is very long.
_Generative System._--The probosciformed penis is short and thick,
and covered with very minute tufts of bristles: there is no
knife-edged projection at its dorsal base. The vesiculæ seminales are
much convoluted and of great length. The ovarian tubes form a small
sheet within the rudimentary shell, in the normal position, over
the basal membrane; and likewise higher up between the two folds of
corium surrounding the pseudo-peduncle; they do not, however, appear
to occur round the lower part of the peduncle: they extend highest
on the rostral and carinal sides, and lowest on the two lateral
faces. The ova are wonderfully numerous; they are 15/2000ths of an
inch in length: they form, instead of two thin lamellæ, two almost
cylindrical packets, which are held together by most feeble membrane.
Each packet, in the upper part lies between the two folds of the
branchiæ; and in the lower part, is embraced only by the larger outer
fold. The two packets of eggs sometimes cohere together at their
lower ends.
_Imbedment._--The shell is imbedded up to its summit, but the shell
is very shallow. The imbedment seems due either to the compression
of the epidermis of the porpoise, or to its formation beneath the
shell having been checked; the outline of the true skin under the
dark-coloured epidermis is not in the least affected. The epidermis
fills up the bay-like spaces formed by the inwardly folded walls, and
firmly adheres to them.
_Affinities._--This genus presents very varied affinities to
Tubicinella, Coronula, and Platylepas. To the latter it is more
especially allied in the compartments, being singly folded inwards,
though the sides of the folds are not here closely pressed together,
as in Platylepas: in both these genera the fold is less deep in the
rostrum, of which fact, in Platylepas, the final cause is evident,
but here there seems no cause, excepting the simple one of affinity.
Xenobalanus is further allied to Platylepas, in the lesser size of
the inner fold of each branchia, compared with the outer, and in
the structure of the cement-glands, and to a certain extent in that
of the sheath. To Coronula the special alliance is shown by the
remarkable character of the pseudo-radii lying between the parallel
and adjoining walls, and in the general character of the cirri: in
Coronula we have the terga sometimes quite aborted, and the scuta
of small size, thus exhibiting a tendency to the entire absence of
opercular valves, so remarkable in Xenobalanus. To Tubicinella, the
alliance is still more plainly shown in the external shape of the
whole animal, wider at top than at bottom,--in the opercular membrane
descending almost to the very base--in the relative positions of the
different parts and organs--in the upper end of the shell continually
scaling off--in the prominence of the transverse external ridges--in
the sheath being divided into successive zones of growth, and being
prolonged nearly to the basal membrane--and in the edges of the alæ
being smooth. Internally, the resemblance is also plainly shown,
in the strength of the internal tunic of the sack--in the branchiæ
springing from two approximate fleshy crests--in the freely depending
acoustic orifices--in the form of the thorax and prosoma--and
consequently of the alimentary canal.
Seeing the state of the almost rudimentary shell, it is not difficult
to imagine its total disappearance. Thinking of this, it occurred
to me to doubt for a few minutes, whether Anelasma, described in my
volume on the Lepadidæ, which presents many points of resemblance with
Xenobalanus,--viz., in colour, in the strength of the internal membrane
of the sack, in the size of the orifice with its thin membranous
margin, in the entire absence of opercular valves, and in epizoic
habits,--might not in truth be a sessile cirripede, in an extremely
altered condition; for the cirri of Anelasma are in so rudimentary a
state, and the mouth in so modified a condition, that the internal
characters by which pedunculated cirripedes can be distinguished from
the Balaninæ, though not from the Chthamalinæ, are almost lost. But
if Anelasma had belonged to either of the two sub-families of the
Balanidæ, the sack would have penetrated almost to the bottom of the
peduncle; there would have been only a single layer of longitudinal
muscles round the peduncle; and there would not have existed ovigerous
fræna. I believe that Anelasma and Xenobalanus are only analogically
connected, being no more related together by true affinity, than any
other two genera in the Lepadidæ and Balanidæ.
_Sub-Family_--CHTHAMALINÆ.
_Shell with the rostrum having alæ, but without radii: rostro-lateral
compartments without alæ on either side: parietes not porose._
_Mouth with the labrum bullate; palpi hardly touching each other: third
pair of cirri with the segments resembling those of the fourth pair._
The shell in this sub-family consists of four, six, or eight
compartments, with the addition, in Catophragmus, of several whorls
of supplemental compartments or scales, like those forming the lower
part of the capitulum in Pollicipes. The rostrum has alæ and no
radii; in shape and size it resembles the carina. The rostro-lateral
compartments are destitute of alæ; in all cases they overlap the
adjoining compartments, and have radii (when such are developed) on
both sides. In the genus, Pachylasma, however, the shell must be
looked to very young, in order to detect this normal structure, for
soon the true rostrum and rostro-lateral compartments become blended
together, making a compound rostrum, destitute, as in the Balaninæ,
of alæ, but furnished with radii. The parietes are never porose, nor
furnished with regular symmetrical ribs (representing the longitudinal
parietal septa) on their inner surfaces: sometimes, however, on the
basal internal edges, there are some irregular depending points. In
conformity with the simplicity of the parietes, the Radii (which are
seldom much developed) have likewise a simple structure; and often are
merely formed by the simple lateral growth of the parietes. The scuta
and terga are articulated together more deeply than is usual in the
Balaninæ; and the terga never have a long spur.
Looking to the animal's body. The Labrum is always swollen and
bullate, and though hollowed out, is never notched in the middle. The
Palpi are rather small, and have not their tips nearly touching each
other. The mandibles generally have their lower main teeth laterally
single, though sometimes they are double: the inferior angle is always
pectinated. The maxillæ are always notched under the upper pair
of great spines. Of the _Cirri_, the third pair much more closely
resembles, in external structure and powers of movement, the fourth
than the second pair, though sometimes a few of the basal segments,
especially on the anterior ramus, are thickly covered with bristles, as
on the segments of the second pair. In three species belonging to two
genera, there are caudal appendages, resembling those generally present
amongst the Lepadidæ, but never found in the Balaninæ. The Branchiæ
are generally smaller than in the Balaninæ; and are sometimes quite
rudimentary.
In the introductory description (p. 152 and 176) of the Family
Balanidæ, I have already discussed the relations of the Chthamalinæ to
the Balaninæ, and of the several genera to each other, so that I need
not here re-enter on the subject.
12. _Genus_--CHTHAMALUS. Pl. 18, 19.
CHTHAMALUS. _Ranzani._ Memoire di Storia Naturale, 1820.
EURAPHIA. _Conrad._ Journal Acad. Nat. Sc. Philadelphia, vol. 7, 1834.
_Compartments six: basis membranous, but sometimes in appearance
calcareous, owing to the inflected parietes._
Distribution mundane; attached generally to littoral rocks and shells.
This, the typical genus, is the largest and widest distributed group of
the sub-family Chthamalinæ. The founder of the genus apparently did not
perceive its essential character; Savigny, however, as is evident from
the excellent figure in the great work on Ægypt, perfectly understood
the difference between Chthamalus and Balanus. I was first indebted
to Dr. J. E. Gray for explaining to me this difference; but the only
published account which I have met with is in a paper by the Rev. R. T.
Lowe,[130] in which he states, on the authority of Mr. Clark of Bath,
that in Chthamalus the anterior compartment or rostrum has alæ like
the posterior compartment or carina, the anterior or rostro-lateral
compartments being destitute of alæ. These characters being exactly
reversed in Balanus, as I have already explained (p. 176) under the
sub-family of Balaninæ.
[130] 'Zoological Journal,' vol. 3, p. 76, 1828.
The shell, owing apparently to its containing much animal matter, is
particularly subject to disintegration; and when thus much affected it
is quite impossible to distinguish the species by external characters.
It is, in fact, best to cast on one side external appearance, though
when the shells happen to be well preserved, each species has its own
peculiar aspect. We have in this genus smooth and plicated, cylindrical
and depressed varieties of most of the species. The development of the
radii is very apt to vary, and even the compartments often become so
completely united and calcified together that the sutures are almost
or quite obliterated. A more serious difficulty in discriminating
the species, arises from the fact of the opercular valves, not only
varying extremely in external appearance in consequence of the greater
or less disintegration of their apices, and consequent exposure of
their articular ridges and furrows (compare fig. 1 _a_, 1 _b_, 1 _c_,
in Pl. 18), but from their truly varying in outline with the varying
shape of the shell: this latter circumstance is probably due to the
opercular membrane which unites the valves to the shell being very
narrow, and in consequence, differences in the shape of the shell
affect the opercular valves, in a manner and to a degree to which the
Balaninæ are not subject (compare fig. 1 _e_, 1 _f_, 1 _h_, in Pl. 18).
The scuta, on the other hand, differ to an unusually slight degree in
the different species. In the common _Chthamalus stellatus_, which
abounds on the southern British shores, the whole external aspect
of the shell is often so completely masked, owing to its varying
shape, its obliterated sutures, its deeply disintegrated and punctured
surface, and by the corroded condition of its opercular valves, that
I have found this species, in the collections of naturalists who have
attended to cirripedes, arranged actually on the same tablet, mingled
with specimens of _Balanus balanoides_. I have myself several times
found it less troublesome to discriminate these two genera by the
included animal's body than by the shell,[131] though the latter, when
well preserved and developed, possesses such obviously well-marked
characters: the same thing has occurred to me with some of the other
species.
[131] It will be found ultimately to save time and trouble, to soak
for half an hour in hot caustic potash, a specimen out of each
group of shells to be examined, and then well wash and brush the
separated valves and compartments; this process has been followed
by me with all the species here to be described.
_General Appearance._--The shell is generally depressed, but sometimes,
when growing in groups, cylindrical. The surface is either smooth or
longitudinally folded; and, as already stated, very apt to be deeply
disintegrated. The radii, when developed, are narrow, with their
summits oblique and rounded; but they are often quite absent, and
sometimes even the sutures are almost obliterated. The rostrum and
carina are of the same shape and size, and the two lateral compartments
on each side are of nearly equal breadths. The orifice is generally
sub-rhomboidal, being widest towards the carinal, instead of towards
the rostral end, as is usual in Balanus: but in _Chthamalus fissus_
the orifice is narrow and elongated. The opercular valves have their
apices generally disintegrated and worn away, and are then seen to be
deeply locked together. The colour of the shell is dirty white or dull
purplish-red or brown; but in _C. intertextus_ rich violet-purple. The
species are small, not often exceeding half an inch in basal diameter,
with the exception of _C. Hembeli_, of which I have seen a specimen two
and three quarters of an inch in diameter.
_Scuta._--These present no particular character, excepting in most
cases the large development of the articular ridge, and sometimes the
presence of a furrow above the articular ridge, of which only traces
can rarely be detected in Balanus; hence in these cases, as in _C.
Hembeli_ (Pl. 18, fig. 5 _a_, 5 _c_) and _scabrosus_ (Pl. 19, fig. 2
_a_, 2 _c_), the line of articulation between the scutum and tergum
is more complicated than is usual. There is generally a slight pit,
sometimes even furnished with small crests, for the lateral scutal
depressor muscle: in _C. scabrosus_ (fig. 2 _d_, _q_) a part of this
muscle is attached to a small pit at the basi-scutal corner of the
tergum,--a fact of which I have observed no instance in any other
genus. In _C. intertextus_ the terga and scuta are calcified together,
without even a suture being visible on their internal faces. In _C.
Hembeli_ the valve is externally marked by a few longitudinal furrows.
The _Terga_, like the scuta, have a prominent articular ridge and deep
furrow. In many specimens of _C. stellatus_ and in _C. scabrosus_
the valve is narrow: in _C. fissus_ it is triangular and nearly
equilateral. The crests for the depressores muscles are well developed:
in _C. scabrosus_ (Pl. 19, fig. 2 _d_, _p_) these crests are united
into a plate, which, together with the outer lamina of the valve, forms
a deep narrow pit: in _C. Hembeli_ the crests are furnished with small
sub-crests. The opercular membrane is narrow; it is sometimes furnished
with a few minute spines.
_Structure of Parietes._--The parietes are solid, and composed of
successive layers of shell; the inner surface varies in condition
in the same species, being either smooth or marked with branching
impressed lines, or mamillated, or often irregularly punctured for the
entrance of tubuli. Owing to these tubuli, the walls, when externally
disintegrated, often become punctured. In certain depressed varieties
of both _C. stellatus_ and _scabrosus_ the walls are supported by
irregular depending columns, placed along either one side or both sides
of the sutures. In _C. intertextus_ we have the remarkable character of
the wall of the shell growing (I presume after a certain age) almost
rectangularly inwards, thus forming a rather wide, flat, calcareous rim
round the central basal membrane. In _C. Hembeli_ the internal basal
edges of the parietes, in one moderately young specimen, were rugged
with irregular points, but presented no other remarkable appearance;
but in five old and very large specimens, the whole basis was
calcareous, being absolutely continuous with the inner lamina of the
parietes, showing that the latter had grown flatly inwards all round,
and had then become confluent in the middle, so that there was no
longer any basal membrane; excepting, no doubt, that which had existed
in the younger stage, and which would be preserved in a functionless
condition between the surface of attachment and the inflected parietes.
When an opercular valve or compartment is dissolved in acid, layers of
tissue are left, and these are seen to be penetrated by tubuli, which
enter at the punctures before mentioned on the inside of the shell:
these tubuli often stand in groups of three or four together; they
are about 1/10000th of an inch in diameter. Besides these irregularly
scattered tubuli, there are in the opercular valves of _C. antennatus_,
innumerable smaller parallel tubuli, running to the external investing
membrane.
_Structure of the Radii and Alæ._--The radii, when developed, are
always rather narrow. Their recipient furrows are generally nearly as
broad as the radii themselves. Their edges are either quite smooth, as
in _C. antennatus_; or very finely crenated; or, as in _C. dentatus_
and _Hembeli_ (Pl. 18, fig. 3 _a_, 5 _a_), so strongly crenated as to
make the suture, both externally and internally, toothed: in these
two species, the radii are ribbed in transverse lines parallel to
the basis, each rib corresponding with one of the projecting and
interlocking teeth on the sutures. In _C. intertextus_, and much less
plainly in most specimens of _C. scabrosus_ (Pl. 19, fig. 1 _a_, 2
_a_), we have a structure in appearance very different, for the radii
here consist of several very oblique plates, (_i. e._ nearly parallel
to the parietes) on both sides of the sutures, which are interfolded or
locked together: I believe that this structure is a mere modification
of that in _C. dentatus_ and _Hembeli_, the transverse ridges on the
radii of those species being here developed into oblique plates. We
shall hereafter meet with a similar structure in the genus Verruca;
to which genus, until meeting with these two species of Chthamalus, I
had thought that the interfolding sutures had been confined. The alæ
have their edges generally finely crenated: during diametric growth
(when such takes place), they are rarely added to above the level
of the opercular membrane, and hence their summits are oblique: in
_C. intertextus_, however, the alæ are laterally added to above the
opercular membrane, and their recipient furrows are likewise added to,
of which fact I have seen no other distinct instance in any genus;
hence on both sides of the sutures, in the sheath of this species, the
lines of growth are upturned. In some much disintegrated specimens,
both of _C. stellatus_ (var. _depressus_) and of _C. antennatus_, the
radii have been corroded away, and the diametric growth is effected
exclusively by the growth of the alæ, which are moreover much exposed,
and rendered conspicuous. The sheath descends a moderate distance down
the shell. When a shell is boiled in potash, the sutures (excepting
when abnormally calcified together, as very often happens with some
species) always fall apart, showing that the union is simply by
animalised matter.
_Basis._--The basis is always membranous; but we have seen, in _C.
intertextus_, that the walls form a flat ledge all round the base,
and that in _old_ specimens of _C. Hembeli_, they grow so far inwards
and become so completely confluent, that they might most easily be
mistaken for a true calcareous basis. I may add, that in one elongated
specimen of _C. stellatus_ from La Plata, the walls had likewise
grown rectangularly inwards, forming a flat base, and had then turned
upwards in the middle, forming a medial crest, with the edges not quite
calcified together. The true basal membrane is very obscurely divided
into concentric slips. I observed in several species, attached to the
lower surface, an excessively fine network, quadrangular or hexagonal,
of yellow vessels, which seemed insensibly to pass into the sheets,
discs, and globules of cement, by which the membranous basis adheres to
the supporting surface. I saw, in _C. antennatus_, numerous irregular,
bifurcating, and inosculating cement-ducts, of unequal diameter, often
crossing each other, and sending off branches ending in points: the
older ducts, instead of being solidly filled up with cement, were only
divided by septa. I did not succeed, in any species, in discovering the
cement-glands.
_Mouth._--The _labrum_ is slightly bullate, with the middle portion
depressed, but not forming a notch; in some species it is hairy, and
in some pectinated with short spines. The _palpi_ are of moderate size.
The mandibles have from three to five main teeth, the number sometimes
varying even in the same species: the lower teeth are either plainly
double laterally, or very obscurely double, or to all appearance
quite single: a rather large lower portion of the mandible is finely
pectinated. The maxillæ are always notched under the two or three large
upper spines: the notch bears some fine spines: beneath the notch there
are some large spines, and at the inferior angle some smaller ones.
_Cirri._--The first and second pairs are always very short compared
with the four posterior pairs. The rami of the first pair are slightly
unequal. The third pair, in length and arrangement of the spines, very
closely resembles the three posterior pairs; in _C. intertextus_,
however, the few basal segments, chiefly on the anterior ramus, are
thickly clothed with bristles, like the segments of the second cirrus.
In _C. antennatus_ (Pl. 29, fig. 2), the anterior ramus of this same
third pair is usually (one single specimen being excepted) much
elongated, having at least twice as many segments as the posterior
ramus, but the number is variable; and these segments, either all,
or only the upper ones, instead of having their spines regularly
arranged in pairs, in a double row, are surmounted each by a circle of
spines: I suspect that these elongated rami of the third cirrus act
as antennæ. It can hardly be an accidental coincidence, that certain
genera, as Lysmata and Pandalus, amongst the Macrourous Crustaceans,
have the same leg (homologically the second thoracic limb) elongated
and antenniformed. Certain varieties of _C. stellatus_ and _cirratus_,
also, have the anterior ramus of this same third cirrus considerably
elongated. We are thus reminded of the remarkable variability in the
numbers of the segments, and in the arrangement of their spines, in the
cirri of _Tetraclita porosa_; in that species, however, it was chiefly
the terminal segments of the _posterior_ ramus of the third cirrus
which were so highly variable. The three posterior pairs of cirri in
Chthamalus support from three to five pairs of main spines on each
segment, the number often varying in the same species, with some minute
intermediate bristles. The dorsal surfaces of these segments, in some
of the species, are serrated in an upward direction.
_Body, &c._--The body does not present any particular character: in
_C. scabrosus_ there is a slight ridge running from the base of the
first cirrus towards the adductor scutorum muscle: this ridge is
clothed with a few hairs; there are also some hairs at the carinal
end of the sack. In _C. dentatus_, also, there are hairs on the outer
tunic of the prosoma. In two species which I opened, there were no
cæca to the stomach. The ova vary in length from 13 to 14/2000th of
an inch in length; they are packed in two lamellæ lying on each side
of the animal's body. The larva just escaped out of the egg, in _C.
stellatus_, _scabrosus_ and _dentatus_, had a large probosciformed
mouth.
_Branchiæ._--These present a very singular amount of difference within
the limits of the same genus. In _C. stellatus_ and _antennatus_ we
have a simple fillet, tapering a little, barely plicated, and about
half, or more than half, as long as the sack; in _C. scabrosus_ the
branchiæ are entirely aborted, or are perhaps represented by the slight
hairy ridge at the carinal end of the sack: in _C. dentatus_, on the
other hand, each branchia consists of two large folds, barely plicated,
almost covering the whole side of the sack, so that here the branchiæ
are developed to an unusual degree, more than in Balanus, and as in
Coronula and its allies: in the same manner as in these latter genera,
the outer fold is considerably larger than the inner fold.
_Affinities._--_C. intertextus_ is the most distinct species of the
genus, as shown by the peculiar radii and alæ, by the scuta and terga
being calcified together, by the character of the third pair of cirri,
and by the inflected parietes forming a ledge round the membranous
basis; but in this latter respect _C. intertextus_ resembles _C.
Hembeli_. _C. Hembeli_, in its serrated radii, is closely related to
_C. dentatus_; and this latter species differs in the structure of
its radii only in degree from certain varieties of _C. stellatus_.
Lastly, _C. intertextus_, in its peculiar radii, closely resembles _C.
scabrosus_, and this latter species does not differ much from the other
species. Hence the genus Chthamalus has no claims to be subdivided into
smaller genera.
_Range._--The species are found all round the world, from (as far as
I have seen) 54°-55° north, to Cape Horn, in 55°-56° south. All the
species, of the habits of which I know anything, are littoral; and
in many parts of the world are excessively numerous, quite covering
large spaces of the coast-rocks, and often coating the coast-shells.
_Chthamalus dentatus_ is littoral, like the other species; but it
often lives attached on _Balanus tintinnabulum_ and _amphitrite_, on
the bottoms of ships arriving in British ports from the west coast
of Africa. I do not know of any instance of more than two species
occurring in the same region. Some of the species have large ranges:
_C. scabrosus_ extends from the Falkland Islands and Tierra del Fuego
to Peru; and _C. stellatus_ has an enormous extension over almost the
whole world, excepting the west coast of South America and Australia.
I do not believe any species of the genus, owing probably to their
littoral habits, have hitherto been found fossil.
1. CHTHAMALUS STELLATUS. Pl. 18, fig. 1 _a_-1 _h_.
LEPAS STELLATA. _Poli._ Testacea Utriusque Siciliæ (1795), Tab. 5,
fig. 18-20.
---- DEPRESSA (_var._) Ib., Tab. 5, fig. 12-16.
CHTHAMALUS STELLATUS. _Ranzani._ Memoire di Storia Naturale (1820),
Tab. 3, fig. 21-24.
---- GLABER (_var._) _Ranzani._ Ib.
---- STELLATUS. _Philippi_ (!). Enumeratio Mollusc. Siciliæ.
LEPAS PUNCTATUS. _Montagu_ (!). Testacea Britannica (1803).
_Shell white or gray, generally much corroded and punctured: radii
(when present) narrow, with their sutural edges most finely crenated;
tergum with the crests for the depressor muscle depending barely
beneath the basal margin._
_Var._ (_a_, communis) fig. 1 _a_, 1 _f_: _Shell conical depressed,
upper part corroded, walls folded, sutures moderately plain or
obliterated; radii not developed; orifice broadly oval_.
_Var. (b): Shell elongated, sub-cylindrical; sutures obliterated;
surface much corroded; orifice almost circular._
_Var._ (_c_, communis) fig. 1 _c_, 1 _e_, 1 _h_: _Shell conical,
folded, sometimes covered by membrane; radii developed, narrow;
orifice sub-hexagonal, toothed._
_Var._ (_d_, fragilis) fig. 1 _d_: _Shell conical, smooth, thin;
compartments easily separable, pale-coloured; radii developed,
narrow; orifice large, toothed, sub-hexagonal._
_Var._ (_e_, depressus) fig. 1 _b_, 1 _g_, 1 _h_: _Shell much
depressed, surface much corroded, smooth: alæ largely exposed, marked
by lines of growth: radii not present; parietes on the under side
often supported by pillars; orifice sub-hexagonal._
_Hab._--Southern shores of England, Ireland, Isle of Man,
Mediterranean, Madeira, Cape de Verde Islands, Southern United States
(Charlestown), West Indies, Brazil (Bahia), Rio Plata (Guritti
Island), Red Sea, Philippine Archipelago, Coast of China, Gulf of
Corea, Oregon or Northern California.
This species is very widely distributed and extremely common. On
the coast-rocks of the southern shores of England it is, in parts,
even more numerous than the _Balanus balanoides_, with which it
often grows mingled. As already stated, it is often confounded in
British collections with this species of Balanus, under the name
of _Balanus punctatus_; that Montagu had this Chthamalus in view,
when describing his _Lepas punctatus_, is certain, from his original
specimen in the British Museum, but whether this was the case with
his predecessor, Pulteney, in the Dorset Catalogue, I do not feel so
sure. _Chthamalus stellatus_ varies, as we shall immediately see,
extremely in appearance. Some of the varieties, as _var. depressus_,
which in external aspect are especially distinct, I have no doubt are
really varieties, but whether this is the case with some of the forms
from the more distant localities, is a little more doubtful; but I beg
that it may be observed, that I have, in the case of every one of the
varieties, and of all the specimens from distant localities, cleaned
with potash and most carefully examined the disarticulated valves, and
likewise dissected the included animal's body.
_General Appearance._--British specimens are usually conical, and
have their walls folded, corroded in their upper parts, with the
radii not developed, and the sutures more or less obliterated; in
this state the orifice of the shell is entire, and very broadly oval:
I have seen specimens in this condition from Madeira, Brazil, and the
Gulf of Corea. The shells when crowded, are rendered cylindrical,
and more or less elongated, with the sutures as viewed externally
quite obliterated; the surface rugged and much disintegrated; and
the orifice nearly circular: in this extreme condition I have seen
specimens from England, from near Genoa, and from the mouth of
the Plata: the specimens from these last two localities were of
remarkable size, being half an inch in diameter, and rather more than
half an inch in height. Again, other British specimens (fig. 1 _c_),
though not nearly so common, are rather steeply conical, and have not
only their sutures distinct, but narrow radii are plainly developed
on apparently both sides of the sutures; in this case the orifice
is slightly toothed, and is rather elongated: I have seen specimens
in this condition, but with their walls rather more deeply folded,
from the Cape de Verde and Philippine Archipelago. Other specimens,
from some unknown tropical sea, differed only in the walls being but
slightly folded, and being so perfectly preserved as to be externally
covered with membrane: this latter circumstance gave the specimens
a peculiar appearance. In this condition were some specimens (fig.
1 _d_) attached to oysters sent to me by Professor Agassiz, from
Charlestown; and which differed from all the others that I have seen,
in the thinness and smoothness of their compartments, and in the
facility with which the compartments separated from each other and
from the surface of attachment: hence I have called this form, which,
until finding more important differences, I must rank as a variety,
_fragilis_: it has stronger claims than the other varieties to be
specifically separated; but I suspect that it has been exposed to
some peculiar modifying conditions.
Lastly, we have the variety (fig. 1 _b_) excellently figured and
described by Poli, under the name of _L. depressa_: this is much
depressed, with the walls thin, not folded, with the surface much
corroded, with the sutures very distinct, with the radii not at all
or barely developed, but with the alæ largely developed, and marked
with lines of growth, resulting from the diametric growth of the
shell: the orifice is hexagonal, but broadest towards the carinal
end: most of the specimens, _but not all_, have on the under sides
of their compartments rugged pillars depending from beneath the alæ,
for the purpose, apparently, of supporting the much depressed shell.
This much depressed variety attains a larger basal diameter (but not
a greater bulk or internal capacity) than any other variety, namely,
sometimes three quarters of an inch. The great peculiarities of this
variety result, apparently, from its much depressed form, deeply
corroded not folded walls, and considerable diametric growth; from
the latter cause the alæ are largely developed; as I can find neither
internally nor externally any fixed diagnostic character I have not
hesitated to rank this form as a variety. Poli found his specimens
mingled with the ordinary _C. stellatus_, on the shores of Sicily;
and I collected at St. Jago, in the Cape de Verde Archipelago, some
specimens nearly as well characterised, also associated with the
common variety.
The shell itself is dirty-white or gray, or brown: in some varieties,
however, the white is nearly pure: internally the parietes are
generally tinted dull purple. The corium of the sack is dark
greenish-black, with a white edge to the lips lining the aperture
between the opercular valves.
_Structure of Shell and Radii, &c._--The under side of the parietes
is either quite smooth, or marked with slight, branching, depressed
lines; or mamillated; or irregularly studded with large pores. I have
already alluded to the pillars, depending from the under sides of the
alæ in most specimens of _var. depressus_: these pillars tend to form
ridges, parallel to the sides of the compartment, like those we shall
presently see in certain specimens of _C. scabrosus_. The radii are
very narrow, when best developed; their edges, when disarticulated,
can be seen, when examined by a strong lens, to be finely crenated.
The edges of the alæ are likewise very finely crenated.
_Scuta._--The outline of these valves varies considerably in
specimens taken out of the same group: we have either a nearly
equilateral triangle (fig. 1 _f_) or the tergal margin (1 _e_)
is much shorter than the other margins. There is always a deep
depression for the adductor muscle, and a small pit of very variable
depth for the lateral depressor. But the tergal margin offers
the greatest variability; here we see a very prominent articular
ridge or fold, having either a straight edge or a single or double
prominence (fig. 1 _e_-1 _h_). In specimens in the same group we
find considerable variation in these points; but the amount of
difference is sometimes so great, that I long hesitated whether
to rank some of the varieties as species. The _Terga_, likewise,
vary greatly in shape and width: in some of the commonest varieties
(and in _var. depressus_) the valve is very narrow, with the under
surface channelled or concave: in other varieties the valve is much
broader and flatter. The spur is but slightly developed. The crests
for the depressor muscles barely descend beneath the basal margin of
the valve. The articular ridge, in some varieties (as in specimens
from Madeira and the Cape de Verde Islands), is extremely prominent
and straight (1 _f_); in others, it is little prominent and deeply
sinuous (1 _h_). This great variability in the articular margins
of the scutum and tergum seems to be mainly due to the corrosion
to which these valves have been subjected, and their consequently
modified growth: in some specimens the articular ridge of the scutum,
and in others that of the tergum, has been largely developed, in
either case their mutual outlines being greatly affected.
_Branchiæ._--These are narrow, hardly at all plicated, elongated,
being about half as long as the sack. In a specimen from La Plata,
this organ ended rather more abruptly in a point than it did in other
specimens.
_Mouth._--The crest of the labrum is usually hairy, but in a specimen
from Bahia (Brazil) there were some very fine teeth. The palpi vary
somewhat in shape, and sometimes have a row of bristles along their
basal exterior margins. The mandibles usually have four main teeth,
the lowest one being confluent with the inferior pectinated angle: in
_var. depressus_, in the same individual, there were only three teeth
on one side of the mouth and five on the other; the lower main teeth
are laterally double, but generally one tooth of each pair is so
small and obscure as to be perceived with difficulty. In the maxillæ,
there are some large spines above the notch, and in the notch some
fine ones: in _var. depressus_, and in a cylindrical _var._ from La
Plata, there was quite a tuft of small spines above the notch.
_Cirri._--The outer surface of the pedicel of the second cirrus
bears a tuft of long, fine, plumose hairs: the terminal segments of
the rami of this cirrus sometimes (as in the La Plata specimens)
support a clump of coarsely pectinated spines. In specimens having
six segments in the shorter ramus of the second cirrus, the shorter
ramus of the third cirrus had fifteen segments. The two rami of the
third cirrus are usually equal in length and in the number of their
segments; but in the Brazilian specimen there were fifteen segments
in the posterior, and twenty-six in the anterior ramus; in another
specimen, fixed on a tropical Perna, there were in the two rami of
this third cirrus eighteen and twenty-four segments. In the three
posterior pairs of cirri each segment carries either four or five
pairs of main spines: the segments vary a little in the degree to
which they are elongated, being most elongated in the var. from La
Plata, with an elongated shell.
_Varieties._--It will have been observed, that the shell, in the
specimens from several distant quarters of the world over which this
species, as I believe, ranges, differs considerably in external
aspect: so do the opercular valves; and so do the parts of the
mouth and cirri: but I cannot make out that these differences are
coordinated. Thus, _var. depressus_, which is so entirely different
from the others in appearance, differs only internally in the
presence of a tuft of fine spines above the notch of the maxillæ;
and this character is found in the La Plata variety, which, as far
as the shell and opercular valves are concerned, is at the other end
of the scale of variation. Again, _var. fragilis_, from Charlestown,
presents, in the animal's body, hardly any difference. The Brazilian
specimens, which in the shell and operculum offer only quite common
characters, have the remarkable peculiarity of a considerable
difference in the length and number of the segments in the rami of
the third cirrus; they, also, have the segments of the sixth cirrus
considerably elongated, and the labrum finely toothed. Of these
peculiarities one alone, namely, the inequality in the rami of the
third cirrus, but in a lesser degree, is common to the specimens
adhering to the tropical Perna, which had a shell very unlike the
Brazilian variety, but which, on the other hand, differed scarcely in
a single character from some other specimens from an unknown tropical
sea, in which the rami of the third cirrus were quite equal. The La
Plata specimens differ most in internal characters, viz., in the
tuft of fine spines above the notch of the maxillæ, in the coarsely
pectinated spines on the tips of the second pair of cirri, in having
the segments of the sixth cirrus much elongated, and in the apex of
the branchiæ being abruptly pointed; yet in the shell and operculum
they were identical with certain Mediterranean varieties. From these
several facts, I must believe that all the widely distributed forms
here grouped together, do really belong to the same species.
2. CHTHAMALUS ANTENNATUS. Pl. 18, fig. 2.
_Shell conical, generally smooth: when not deeply corroded of a pale
dirty flesh-colour: sutures always distinct: radii, when present, with
their sutural edges quite smooth._
_Hab._--New South Wales, (Moreton Bay, 27° S.; Sydney; Twofold Bay),
Van Diemen's Land (Hobart Town). Attached to littoral rocks and
shells; Mus. Brit., Cuming, Darwin.
_General Appearance and Structure of Shell._--Shell conical, rather
smooth; when not much corroded, of a pale dirty flesh-colour; often
covered by membrane; sometimes deeply corroded, extremely rugged,
and then of a brown colour; in this condition not much punctured,
as generally is the case with _C. stellatus_. Sutures almost always
quite distinct; rarely the shell becomes cylindrical with the sutures
obliterated. Orifice moderately elongated, sub-hexagonal. Radii
rather narrow, but not so narrow as in _C. stellatus_, smooth, with
their upper margins very oblique: when disarticulated their edges
are quite smooth. The edges of the alæ are sometimes crenated, and
sometimes not so, being only marked by lines of growth; they are
often rather thick. The parietes are usually rather thick, with their
internal surfaces smooth, and not mamillated, as is so often the case
with _C. stellatus_. The largest specimens which I have seen, were .6
of an inch in basal diameter.
_Opercular Valves._--These are hardly distinguishable from those of
_C. stellatus_. The only very slight difference which I can point out
is, that the crests for the tergal depressores are less spread out,
and depend rather more beneath the basal margin of the valve; and
lastly, that the surface of the tergum, just above these crests, is
rather prominent.
_Branchiæ_: these are oblong; taper but little, and have a broad
rounded end: they are scarcely plicated.
_Mouth._--The crest of labrum is hairy: the palpi are square, and
have no bristles along their basal exterior margins, but long ones
at their truncated ends. The mandibles have three or four main teeth
apparently single: the inferior coarsely pectinated portion is short.
The maxillæ are deeply notched.
_Cirri._--The first and second pairs, and portions of the third, are
darker coloured than the three posterior pairs. The rami in both of
the first two pairs are slightly unequal in length. The third cirrus
is much longer than the second: in a specimen in which there were
six segments in the shorter ramus of the first and second pairs,
there were twenty segments in the posterior and shorter ramus of
the third cirrus; and in this same individual there were no less
than forty-nine segments in the anterior ramus. In another specimen
(Pl. 29, fig. 2) the number of segments in the two rami of the
third cirrus, was 20 and 41; in another, the numbers were 18 and
53; in several other specimens the numbers varied in about these
proportions; but in one single specimen the numbers were equal.
Not only did the number of segments thus vary in the two rami,
but likewise the arrangement of the spines on the segments in the
anterior and longer ramus; in some specimens the spines on all the
segments were arranged in a single circle, and then the organ had a
specially antenniformed appearance: in other specimens, some of the
lower segments (in one case thirteen in number) had the spines placed
in regular pairs precisely as on the posterior ramus, and as on the
three posterior pairs of cirri. Under the genus I have pointed out
the resemblance between this structure and that occurring in certain
Macrourous Crustaceans. The pedicel of the third cirrus had its
spines more crowded and irregular than on the three posterior pairs
of cirri. The segments in the latter vary in bearing either three or
four pairs of main spines. The whole dorsal surfaces of the lower
segments of the several posterior cirri are serrated in an upward
direction by short spines, but to a variable degree.
3. CHTHAMALUS CIRRATUS. Pl. 18, fig. 4 _a_, 4 _b_.
_Shell white or gray: sheath and opercular valves generally clothed by
fimbriated membrane: tergum, with its basi-carinal angle depending and
pointed._
_Hab._--Peru, Chile, Chiloe, Northern Chonos Islands. Attached to
littoral rocks, and sometimes to littoral shells, often mingled with
_Chthamalus scabrosus_; Mus. Brit., Cuming, Darwin.
_General Appearance and Structure of Shell._--Shell dirty white or
gray: sometimes tinted pale purple within; irregularly conical, or
much depressed, or cylindrical and much elongated. Generally much
corroded, sometimes well preserved and covered by membrane. Orifice
rather large, of variable shape. Sutures often quite obliterated.
Radii when developed narrow, with their sutural edges, as well as
those of the alæ, generally very finely crenated, but to a variable
degree. The membrane lining the sheath and covering the opercular
valves, is remarkable from each zone being fimbriated; for this
expression is more correct than to say that the membrane bears a row
of spines, though the fimbriæ do approach in character to spines;
sometimes, though rarely, the fimbriæ are branched. The largest
specimens which I have seen (from Coquimbo and Valparaiso) were half
an inch in basal diameter, and some of these were so much elongated
as to be one inch in height.
_Scuta._--The scuta are rather narrow: they have a somewhat peculiar
appearance, from the articular furrow being wide, and from the
articular ridge projecting with a uniform curvature: the pit for the
lateral depressor muscle has some minute crests, of which I have seen
traces in the foregoing species. The _Terga_ vary somewhat in shape:
they have the basi-carinal angle of the valve, where the narrow
crests for the depressores are placed, pointed and dependent, and
the surface of the valve above these crests is prominent. Altogether
the opercular valves have a sufficiently distinct character to be
recognised without much difficulty.
_Mouth._--The crest of the labrum is not toothed; the palpi have long
hairs along the exterior basal margin. The lower main teeth of the
mandibles are plainly laterally double. _Cirri._--The pedicel of the
second cirrus is extremely broad, and on the exterior margin supports
a tuft of very long, finely plumose spines: in some specimens each of
the lower segments of the anterior ramus of this cirrus bore one or
two very large spines, doubly and extremely coarsely pectinated. In
two specimens the rami of the third cirrus were of equal length: but
in one specimen (from Iquique, Peru), having seven segments in the
shorter ramus of the first and second pairs of cirri, the posterior
ramus of the third pair had fourteen segments, and the anterior ramus
twenty-two segments. The posterior cirri have segments carrying five
pairs of main spines: the dorsal surfaces of the lower segments are
serrated.
4. CHTHAMALUS FISSUS. Pl. 18, fig. 6 _a_, 6 _b_.
_Shell brownish, plicated; orifice twice as long as broad: tergum
triangular, equilateral, with the basal and carinal margins slightly
protuberant._
_Hab._--California, attached to _Lottia grandis_. Peru(?); Mus. Brit.
_General Appearance and Structure of Shell._--Shell globulo-conical,
irregular, with the walls much folded. Colour brownish. Sutures
generally distinct in young specimens, and nearly obliterated in
full-grown individuals. Radii, when present, very narrow. The orifice
of the shell, in full-grown specimens which have their summits a
little worn, is narrow, much elongated, about twice as long as wide,
oval or ovate, with the rostral end the narrowest: the elongation of
the orifice offers almost the only character by which this species
can be externally recognised. The lateral compartments are rather
wider than usual in proportion to the rostro-lateral compartments.
Basal diameter of largest specimen .2 of an inch.
_Scuta_, elongated transversely, with the pit for the adductor muscle
bordered on the under side by an adductor ridge rather more prominent
than usual in this genus. _Terga_, triangular, equilateral, with
the margins slightly curved and protuberant: the basal margin is
regularly and equably curved from one end to the other.
Neither the _Mouth_ nor _Cirri_ offer any peculiar characters
distinct from the genus. I may mention, however, that the crest of
the labrum is toothed, and that the segments of the posterior pairs
of cirri support five pairs of main spines.
The characters by which this species differs from _C. stellatus_ and
from the other species, consist almost exclusively in the triangular
and equilateral terga, and in the much elongated orifice of the shell;
and these differences I believe to be of specific value. I must,
however, confess that I have examined one young specimen attached to
a _Pollicipes elegans_, in which the orifice was not nearly so much
elongated, and in which the terga were not so equilateral, with the
basal margin not quite equably curved, but more protuberant on the
scutal than on the carinal side: from an examination, however, of only
one specimen, and that a young one, I cannot decide on its specific
nature.
5. CHTHAMALUS DENTATUS. Pl. 18, fig. 3 _a_-3 _c_.
CHTHAMALUS DENTATUS. _Krauss_ (!). Die Südafrikanischen Mollusken,
1848, tab. 6, fig. 27.
_Shell dirty white or brownish: sutures formed by interlocking teeth:
tergum with the carinal margin protuberant._
_Hab._--South Africa, Natal; West Africa, Loanda and the Gold Coast;
West Indies (?). Attached to ships' bottoms and to littoral shells,
and to _Tetraclita serrata_, _Balanus perforatus_, and _amphitrite_;
often attached to _Balanus tintinnabulum_ and _amphitrite_ on ships'
bottoms.
_General Appearance and Structure of Shell._--Shell dirty white,
pale brown, or gray: conical, moderately depressed: walls either
broadly and irregularly folded (fig. 3 _a_), with the surface
corroded, or (when attached to ships' bottoms and sometimes to other
Cirripedes) narrowly and regularly folded (3 _b_), with the surface
well preserved, smooth, and generally covered by thin brown membrane.
These latter specimens generally have the shell more steeply conical,
with the orifice rather smaller, and the radii broader, than in
the first-mentioned specimens, which are attached to coast-rocks
and shells, and have had their summits worn down. The sutures in
all cases are tolerably distinct, and have their edges toothed and
interlocked: the crenations are visible before the compartments are
disarticulated, when viewed either internally or externally, but
occasionally they are obscure. The radii, when best developed, are
rather narrow, and of equal width on both sides of the sutures, with
their summits rounded: their surfaces are finely ribbed transversely,
each rib corresponding with, or rather forming a point of, the
toothed edge. On the under side these teeth usually are a little
hollow or are pitted: the radii, in fact, may be said to be folded,
like the parietes, only more symmetrically and narrowly, so that the
points in the opposed edges interlock. The edges of alæ are serrated,
but more finely than the radii. The parietes are rather thin, with
their under surfaces generally smooth. The sheath does not descend
far down the shell. The colour of the corium lining the sack and the
animal's body varies considerably, being either almost black or pale
purple; and the specimens adhering to ships' bottoms are internally
almost white. Of these latter specimens, I have seen some .6 of an
inch in basal diameter: of corroded specimens attached to littoral
shells, I have not seen one quite .4 of an inch in basal diameter.
_Scuta_, with the articular ridge very prominent: the pit for the
adductor is deep, and there are generally some distinct, though
minute, pits for the lateral depressores. The _Terga_ (fig. 3 _c_)
have the articular ridge very prominent; and the carinal margin is
rather more arched and protuberant than in the other species.
_Mouth._--The crest of the _labrum_ is toothed: the palpi are short,
truncated, with long spines arising from their ends, and along the
basal exterior margin. In the mandibles the first tooth is seated
rather far from the succeeding teeth: the inferior part is coarsely
pectinated: the maxillæ are deeply notched. The tips of second pair
of _Cirri_ have many coarsely pectinated spines: the shorter ramus
had six segments, whilst the shorter ramus of the third pair had
eighteen segments: the segments on the posterior cirri carry five
pairs of main spines.
_Branchiæ, &c._--These consist of a double fold, the outer one being
the largest, and the inner semi-circular, as has been described under
the genus. In a young specimen, only one tenth of an inch in basal
diameter, the branchiæ consisted of a single fold; in a specimen a
little larger, there were two folds, but these were of equal size:
ultimately the outer fold increases in size so as to become nearly
double the inner fold. The _Ovarian tubes_ are remarkable from their
large diameter. I was surprised to observe in the specimen only one
tenth of an inch in basal diameter, that the larvæ were ready to
escape. On the prosoma, there are some longish _hairs_ arranged in
short rows, and some few on the membrane lining the sack, and some on
the opercular membrane and valves.
It may be seen in Pl. 18, fig. 3 _a_, and 3 _b_, that I have here
united two varieties considerably different in external aspect: I have
done this without hesitation, inasmuch as there are intermediate forms,
and as the differences are analogous with those so commonly occurring
in sessile cirripedes; dependent on whether or not the specimens
have been exposed to corrosion. I have seen both varieties from
Natal, and both from the west coast of Africa; although the smooth,
well-preserved, narrowly plicated varieties seem more common in western
than in southern Africa. With respect to the range of the species, I
have seen a specimen from the West Indies, but it was the variety
which so commonly adheres to ships' bottoms. This variety often arrives
alive in British ports; and I have seen a specimen picked up dead on
the beach near Dublin.
6. CHTHAMALUS HEMBELI. Pl. 18, fig. 5 _a_-5 _d_, _e_.
EURAPHIA HEMBELI. _Conrad._ Journal Acad. Nat. Sc. Philadelphia, vol.
7, 1831, Pl. 20, fig. 6.
_Shell dull reddish purple: sutures, when not obliterated, formed
by interlocking teeth: basis sometimes rendered calcareous by the
inflexion of the parietes: scutum with two or three furrows extending
down the middle of the valve._
_Hab._--California, near S. Diego, according to Conrad. Mus. Brit.,
Cuming.
I have seen five, old, large specimens, from an unknown locality, with
their whole surfaces deeply corroded, and with most of the sutures
obliterated; and three separated valves of a young specimen. From these
materials, imperfect as they are, I feel no hesitation in identifying
this species with the _Euraphia Hembeli_ of Conrad, which is remarkable
in several respects, and especially from being gigantic in size,
compared to other members of its sub-family.
_General Appearance and Structure of Shell._--The young specimen
(fig. 5 _b_, about .7 of an inch in basal diameter) consists only of
the carina, and the two lateral compartments; but these, as far as
I can judge, resemble the specimen figured and described by Conrad,
which was two inches in diameter. Shell depressed, spreading, surface
moderately smooth, covered by brownish membrane: shell itself pale
dull reddish-purple. The radii are not very narrow, with their
summits rounded and very oblique: their edges are toothed, and their
external surfaces are transversely ribbed, in correspondence with
the interlocking points of the sutures. On the internal surface, the
toothed suture is not visible, except near the base of the shell,
owing to the overlapping of the alæ. The alæ have oblique summits,
which are slightly notched owing to the upturned prominent lines of
growth. The parietes are thick; their basal internal surfaces are
rugged, with slightly branching ridges.
The old specimens (fig. 5 _a_) are so much corroded that not a
particle of the external surface is left: one of them which was
2-3/4 of an inch in basal diameter. Shell much depressed, spreading;
colour pale purple; orifice large, rhomboidal, with a slight hollow
on each side for the corners of the scuta: sutures generally in
part or wholly obliterated; where still preserved, the interlocking
toothed structure is distinct; the sheath is strongly marked by lines
of growth, and is of a dark brown colour. The carina and rostrum
are of unusually large size compared with the lateral compartments;
and this, as far as I can judge, must have been the case with the
younger specimens. That portion of the rostro-lateral compartment
which forms part of the sheath, is reduced to a mere ridge. The most
remarkable character is, that all these old specimens (of which the
smallest measured nearly one inch and a half in diameter) had a flat,
wide, calcareous basis, which is absolutely continuous with the inner
lamina of the parietes, whereas in the younger specimen there was no
appearance of any tendency in the parietes thus to grow inflected.
There can be hardly any doubt that in a series of specimens some
would be found with the parietes first forming a flat narrow ledge
round the true basal membrane (as in the following species); and that
in others, this ledge would be wider and wider, till its edges met in
the middle, and coalesced into a continuous plate.
_Opercular Valves._--I have seen these only in the old corroded
specimens (fig. 5 _c_, 5 _d_): they are locked together by remarkably
strong articular ridges and furrows.
The _Scuta_ have externally two or three impressed lines or narrow
furrows, proceeding from the apex to the middle of the basal margin;
these can be seen only in one of my specimens, owing to the degree
to which the valves have suffered disintegration; but they are
mentioned by Conrad. The basal margin is rather short compared with
the other two margins. The tergal margin is remarkable from the depth
of the upper furrow above the articular ridge, and from the size of
the prominence (appearing like the true apex of the valve) above
this upper furrow. There is a hollow for the adductor muscle, and
traces of crests for both the rostral and lateral depressores. The
_Terga_ are generally but little corroded, and hence the dark brown
membrane with which they are covered is well preserved; the shelly
matter itself is also brown: there is only a trace of this colour in
the more corroded scuta: the external surface of the terga is very
smooth. The spur is pretty well developed, and is half as wide as
the whole valve. The lines of growth are upturned along the carinal
margin. The articular ridges and furrows are much developed. The
crests for the depressor muscles are extremely strong; they depend
beneath the basal margin, and are remarkable (fig. 5 _c_) from being
furnished each with fine sub-crests.
_Animal's body_, unknown.
7. CHTHAMALUS INTERTEXTUS. Pl. 19, fig. 1 _a_, 1 _b_.
_Shell, when well preserved, violet-purple: sutures, when not
obliterated, formed by oblique interfolding laminæ: basis membranous,
but surrounded by a ledge formed by the inflected basal edges of the
parietes: scutum and tergum completely calcified together._
_Hab._--Philippine Archipelago; Mus. Cuming.
_General Appearance and Structure._--Shell depressed, with a
large diamond-shaped orifice. Colour beautiful violet-purple, but
externally much obscured by disintegration, causing the shell to
be ashy gray. Walls smooth or slightly folded. Sutures generally
quite, or almost quite obliterated; but when well preserved, they
differ remarkably in appearance from those in the foregoing species;
for the radii externally here consist of oblique plates or laminæ
arising on both sides of the sutures, standing nearly parallel to the
parietes, and interfolding with each other. These laminæ are rather
plainly marked by lines of growth. Essentially the radii do not
differ much from those in _C. dentatus_ and _Hembeli_; we have but
to produce obliquely upwards the transverse and interlocking ribs on
their radii, and so convert them into laminæ. During the diametric
growth of the shell, the sutural edges of the alæ are added to, in
the usual manner, by upturned lines of growth; and, in addition, the
recipient furrows of the alæ are similarly added to, so that the
lines of growth are upturned, and alæ appear to have been developed
on both sides of the sutures in the same way as the radii appear to
have been developed on both sides in many Chthamalinæ, though rarely
in the Balaninæ. The inside of the shell is beautifully coloured rich
violet; it is punctured with small holes as so often is the case with
_C. stellatus_. In every specimen (all full-grown) which I opened,
the inner basal edges of the parietes were inflected rectangularly
inwards, forming a smooth-edged ledge all round the basal membrane,
which, in proportion to the width of this ledge, was by so much
reduced in diameter. The largest specimen which I have seen was .35
of an inch in basal diameter.
_Opercular valves._--The scuta and terga, in all the specimens which
I have seen, were firmly calcified together; in some, a trace of a
suture could be seen externally, but hardly a trace internally. In
one specimen, there were vestiges of some impressed lines on the
scutum, in exactly the same position in which such occur in _C.
Hembeli_. The scutum is rather narrow. The basal margin of the tergum
is either straight, or depends a little on the scutal side, thus
producing a small spur: the crests for the depressor muscles are
strongly marked, and depend beneath the basal margin.
_Mouth._--The labrum is strongly toothed: the palpi have long hairs
along the exterior basal margin: the mandibles have only three main
teeth, and the inferior coarsely pectinated portion is short; the
maxillæ deeply notched. _Cirri_: the first and second pairs have
their rami slightly unequal in length; the third pair differs from
the same pair in the other species of the genus, in having some few
of the basal segments on the anterior ramus thickly clothed with
spines, so as to be brush-like: there is even a trace of a similar
structure in the lowest segments of the posterior ramus. In the three
posterior pairs of cirri the segments are much elongated, and support
four pairs of spines.
_Branchiæ._--Unknown.
8. CHTHAMALUS SCABROSUS. Pl. 19, fig. 2 _a_-2 _d_.
_Shell (when well preserved) dull purplish-brown: sutures formed by
oblique interfolding though rarely well developed: tergum with a deep
narrow pit, at the basi-carinal angle, for the depressor muscle._
_Hab._--Peru, Chile, Tierra del Fuego, Falkland Islands. Very common;
attached to littoral rocks and shells, and often associated with
_Balanus flosculus_, and sometimes with _Chthamalus cirratus_; Mus.
Brit., Cuming, W. Dunker, Darwin.
_General Appearance and Structure._--Shell generally depressed; when
growing crowded, sometimes cylindrical: colour dingy purplish-brown,
but when much corroded, dirty gray; very young shells are very dark
green, owing to the corium, which is of this colour, being seen
through the valves. Surface generally rugged, from irregular slight
longitudinal folds, and from the transverse overlapping tile-like
lines of growth; but sometimes the surface is nearly smooth, with
very slight longitudinal folds, these being gray coloured, the
intermediate parts being pale dingy purple, the shell thus becoming
striped. Orifice rhomboidal, passing into trigonal, owing to the
great width of the carinal end. Sutures generally very distinct,
rarely obliterated in the cylindrical varieties. Radii narrow,
generally exposing much of the alæ, which are plainly marked by
lines of growth: the radii themselves, when well developed, which
is not often the case, consist of small laminæ or ridges, placed
on both sides of the sutures, and interfolded or interlocked
together: usually only a trace of this structure is exhibited,
but occasionally, along some or along all the sutures (as in the
specimen figured), the laminæ of the radii interfold, as plainly as
in _C. intertextus_. The alæ differ slightly from the alæ of the
other species, in not forming so much of a rectangular projection,
the lower margin running with a gentle curve into the parietes. The
internal surface of the parietes is either smooth, or near the basal
margin is roughened with depending points: in some specimens from the
Falkland Islands, both edges of each suture were inflected, forming
a double ridge, with roughened edges, resting on the basal membrane,
and supporting the shell. I must mention that in my notes made at
these Islands, I remark that the basal membrane seemed sometimes to
be surrounded by a calcareous rim; none of the specimens brought home
are thus characterised; but bearing in mind the affinity of this
species to _C. intertextus_, no doubt we have here an indication
of the shelly ledge surrounding the basis, as described under that
species. The largest specimens which I have seen are a quarter of an
inch in basal diameter.
The _Opercular Valves_ generally have their summits much worn down.
The _scuta_ are elongated in the line of the longer axis of the
orifice; the articular ridge is very prominent, and is placed in
the middle of the tergal margin. The _terga_ are very narrow, as
in some varieties of _C. stellatus_: they are remarkable in two
respects, namely, in the depressor muscle being attached to a plate,
formed apparently by the union of the usual crests, parallel to the
outer lamina of the valve itself, a deep narrow cavity (fig. 2 _d_,
_p_) being thus formed; and secondly, in the far more extraordinary
circumstance of the existence of a small pit (_q_) at the extreme
basi-scutal corner of the valve, in which about half of the scutal
lateral depressor muscle is attached: I have observed no other
instance in any cirripede of the partial attachment of a muscle
properly belonging to one valve to another valve. The figures of
the valves 2 _b_, 2 _c_, 2 _d_, are from specimens most unusually
perfect, with the upper portion not worn away; the ordinary
appearance of the valves as seen from above, is given in fig. 2 _a_;
at fig. 2 _d_, a view is given of the tergum seen from vertically
beneath, showing the cavity for its own depressor muscle, and for
part of the lateral depressor muscle of the scutum.
_Mouth._--The crest of the labrum is hairy, without teeth: the palpi
have long spines at the end, but none along the inferior margin: the
mandibles have either four or five graduated teeth, the lower ones of
which are plainly double laterally: the maxillæ have a very sinuous
edge. _Cirri_: the first and second pairs are very short: on the four
posterior pairs the segments support either four or five pairs of
main spines, with the small intermediate spines rather larger than
usual.
_Branchiæ._--None; but where they ought to occur, there are two very
slight ridges clothed with hairs, about the 1/100th of an inch in
length. On the prosoma, there is a slight ridge, extending from the
base of the first cirrus towards the adductor scutorum muscle, also
clothed with hairs; this unusual character of the prosoma being hairy
is common to _C. dentatus_.
This species is the commonest cirripede on the shores of the Falkland
Islands: many of the specimens are there crowded together, and rendered
elongated and cylindrical, with the walls very thin, and the sutures
often obliterated; as the opercular membrane is very narrow, the
opercular valves are much influenced both in their outline and in
their crests and articulations, by the varying form of the shell: I
have even seen specimens with the scutum and tergum on one side twice
as large as on the other side.
13. CHAMÆSIPHO--_Nov. Genus_.[132] Pl. 19.
[132] Χαμαι [Greek: Chamai], on the ground, and σιφων [Greek:
siphôn], a tube.
_Compartments four, with the sutures often much obliterated: basis
membranous._
_Distribution_, Australia, China (?). Attached to littoral shells and
rocks.
The two species united under this genus agree in having only four
compartments, and in these having a strong tendency to become
confluent; but they resemble each other hardly in any other respect,
more than do all the species of the present sub-family. _Chamæsipho
columna_ differs from all the other Chthamalinæ in the structure of
its second pair of cirri, and _C. scutelliformis_ differs from all in
its shell,--namely, in the small size of the rostrum, with its alæ
but little developed, and in the very peculiar apertures in the three
other compartments. Hence this genus can hardly be considered a very
natural one, though I could not introduce the two present species into
Chthamalus, or into any other genus, without doing still more violence
to the principles of classification followed throughout this work.
Chamæsipho bears nearly the same relation to Chthamalus, as Tetraclita
and Elminius do to Balanus.
1. CHAMÆSIPHO COLUMNA. Pl. 19, fig. 3 _a_-3 _c_.
LEPAS COLUMNA. _Spengler._ Skrifter Naturhist. Selbskabet, b. 1,
(1790), Tab. 6, fig. 6.
_Sutures, excepting during early youth, generally obliterated both
externally and internally: tergum with small pits for the attachment of
the depressor muscle._
_Hab._--New South Wales, Tasmania, New Zealand; extremely common;
attached to littoral shells and rocks; often associated with
_Chthamalus antennatus_ and _Elminius modestus_; and in New Zealand,
often thickly coating _Elminius plicatus_.
I have identified, with some doubt, the present species with the _Lepas
columna_ of Spengler, obtained from Otaheite, as no description is
given by him of the opercular valves, and more especially as Spengler's
specimens were an inch in height and seven lines in width, which is
much larger than any of the many specimens seen by me: from Spengler's
clear description of the structure of his shell, it evidently belongs
to the present genus.
_General Appearance and Structure of Shell._--The sutures separating
the four compartments are generally, excepting at an early age, quite
obliterated, both internally and externally, the shell in this case
consisting of a single piece, with its summit and opercular valves
always much worn. Occasionally the sutures are preserved, and then
the four compartments are seen to be of nearly equal sizes. The
orifice is always broadly oval, with the carinal end the broadest;
and it often approaches closely to circular. The upper part of the
shell is frequently steeply conical, with the lower part spreading
and folded; sometimes deeply folded. Very young shells are apt to
be remarkably smooth. The radii appear never to be developed: the
inside of the shell is smooth. The colour is either blackish-green
(this being the tint of the corium lining the sack), with the upper
part gray from disintegration; or the lower part is brown, from
the investing membrane, the shell itself being pale coloured. This
species seems particularly liable to grow crowded together; often
covering rocks and shells with a honey-combed layer. The basal
diameter of some of the largest specimens was .3 of a inch. In Mr.
Cuming's collection, however, there is an Australian specimen .55
in diameter and .3 in height, which is, moreover, remarkable from
the projecting, extremely rugged, overlapping, dark-coloured layers
of growth, which surround the lower part of the shell: we have seen
that Spengler's specimens, said to have come from Otaheite, are even
broader and considerably higher.
_Scuta_: from the disintegration which the valves have undergone, the
scuta and terga are externally seen to be locked together by a deeply
sinuous articulation. The _Scuta_ have a wide articular furrow and a
very prominent articular ridge; but the exact outline of these parts
varies greatly, very much as in the genus Chthamalus.
The _Terga_ are very narrow, with the under surface channelled: the
attachment of the depressor muscle offers the only peculiarity,--the
muscle being attached to four or five little pits, placed
transversely to the longer axis of the valve; the septa between these
little pits evidently answering to the crests as usually developed.
_Mouth._--The crest of the labrum is hairy, and slightly bullate.
The palpi are rather small, with long spines at their ends. The
mandibles have four (sometimes five) teeth, with the inferior portion
pectinated. The maxillæ are notched.
_Cirri._--The first pair presents no remarkable character. The second
and third pairs are subject to extreme variation, as in _Chthamalus
antennatus_ and _Tetraclita porosa_. In all the specimens, the
anterior ramus of the second cirrus is short, with all the segments
thickly covered with bristles; it is the posterior ramus which varies
so much in relative length and in the arrangement of the bristles;
but in no case are all the segments clothed with bristles as on
the anterior ramus, and as is normally the case with all sessile
cirripedes. In some specimens from New Zealand, the anterior ramus
having only five segments, the posterior ramus was twice as long,
having sixteen segments, with the bristles arranged in circles,
but standing rather thicker together on the basal segments. In a
Tasmanian specimen, the posterior ramus was only a little longer than
the anterior ramus, and the spines were arranged in regular pairs (as
on the three posterior pairs of cirri) on all the segments, excepting
the few basal ones, on which they were more crowded: a nearly similar
arrangement occurred in some other specimens from unknown localities,
excepting that the rami were of nearly equal length. In the third
cirrus, in all the specimens, the anterior ramus has three or four of
its basal segments much broader than the upper segments, and thickly
clothed with spines (as is the case with one species of Chthamalus,
viz. _C. intertextus_) all the other segments having regular pairs
of spines. The posterior ramus of the third cirrus varies in being
either much longer than, or only equal in length to, the anterior
ramus; in the former case (in the New Zealand specimen) the spines
were arranged in circles, giving an antenniformed structure to the
ramus; and in the latter case they were arranged in regular pairs.
In Mr. Cuming's great Australian specimen, there was a further
peculiarity, in the presence on the posterior cirri of a tuft of
intermediate spines between the main pairs; and, in there being on
those segments, which are thickly covered with spines, certain very
large spines, doubly pectinated, with the pectinations elbowed,
closely like the spines met with on the cirri in certain species of
Pollicipes. Finally, the segments in the three posterior pairs of
cirri support five or six pairs of main spines; the dorsal surfaces
of the segment are rough and hairy.
_Branchiæ_, rudimentary; consisting of a small, simple, tongue-formed
fold, projecting about 1/100th of an inch. Ova, 16/2000ths of an inch
in length.
2. CHAMÆSIPHO SCUTELLIFORMIS. Pl. 19, fig. 4 _a_-4 _d_.
_Rostrum very small, elongated, triangular: lateral compartments, each
with an aperture, and carina with two similar apertures, all four
leading into shelly tubular columns._
_Hab._--Attached to _Pollicipes mitella_, probably from the seas of
China; Mus. Brit.
_General Appearance._--This very singular shell would not, without
some examination, be thought to be a cirripede. From the symmetrical
position of the four apertures, with the diamond-shaped orifice in
the middle, with the sutures on each side of the rostrum, and from
its depressed and circular form, this shell bears some resemblance to
the perforated species of Scutella. Shell much depressed, generally
nearly circular, with the basal margin highly sinuous and even
sometimes almost branched. Surface slightly irregular, marked by
fine lines of growth, and covered by brown membrane. Of the four
compartments, the _Rostrum_ is very narrow, triangular, and comes up
to the orifice almost in a point: it is rather depressed, that is, it
lies rather below the level of the other compartments: the straight
sutures separating it from the lateral compartments are distinct in
the upper part, though always obliterated in old shells in the lower
part: these sutures are generally far plainer than those separating
the lateral compartment from the carina, which in most cases are
obliterated and calcified together, excepting close to the orifice.
The alæ of the rostrum are not externally visible, and there are no
radii to any of the compartments. The _Carina_ is twice as broad as
the rostrum, and is furnished with alæ of the usual shape, which are,
to a certain extent, externally visible. The _Lateral Compartments_
are broad, being broader than the carina; they are both penetrated,
down to the surface of attachment, by a hole or rather tube,--the
two holes standing opposite the rostral end of the operculum: the
carina is penetrated by two rather smaller but similar holes. It
is these four holes which give to this cirripede its very singular
aspect: they are rather smaller than the orifice of the shell; they
are oval, with their longer axes placed in the direction of the ray
of the circular shell: their manner of formation will be immediately
explained. The orifice is neatly diamond-shaped and broad: it is
rather small compared with the whole shell, and is closed by the
operculum, which is seated near the summit of the shell. Basal
diameter of largest specimen 2/10 of an inch; few, however, attained
this size, and perfect larvæ were included in much smaller specimens.
_Structure of Shell._--The rostrum (fig. 4 _c_) is remarkable from
its small size, and from the plainness of its sutures, in comparison
with those separating the other compartments, and this is exactly
the reverse of what I should have expected in a compartment tending
to become rudimentary. Not only is the rostrum small, but the alæ
project to an unusually small degree, and gradually slope away into
the lower part of the parietes. These peculiarities are even more
strongly marked in very young shells: thus in one of the size of a
pin's head, the rostrum consisted of a minute parallelogram, without,
as far as I could see, any alæ, and was only one fourth of the size
of the carina,--this latter compartment being only half as wide as
the lateral compartments. The carina, at this early period, had quite
distinct alæ.
The tubular prolongations from the four external holes are of course
very conspicuous on the under side of the shell (4 _b_); and their
structure is there plainly seen. In extremely young shells the holes
are not present; but very soon, at four points of the circumference,
namely, two in the carina and one in each lateral compartment, the
basal edge becomes indented, and during growth more and more deeply
indented; at last the horns or points of the bays, thus formed in
the circumference of the shell, grow inwards and meet, the four
indentations being thus converted into four rings or holes; as
the shell is added to, at its circumference, these come to stand
further and further from the exterior margin; and as the shell at
the same time rises above the surface of attachment, the holes are
added to at their basal edges, and are thus converted into shelly
tubes, generally freely open at the bottom as well as at the top.
Sometimes these tubes are closed at the bottom, and this is usually
caused by their sides having been added to in a spiral direction. A
somewhat sinuous double ridge or fissure, leading from the tubes or
holes to the exterior border, can always be perceived on the under
side of the shell. Occasionally, though rarely, in very old shells,
a second series of holes is formed outside the first four holes,
and often a tendency to this may be perceived in the just-mentioned
fissures expanding a little at their outer ends, thus forming four
new circumferential indentations. The purpose of this peculiar
structure, apparently, is to give support to the much depressed and
thin shell. In _Chthamalus stellatus_ and _scabrosus_, we have seen
a slight indication of a similar structure, in the formation on the
under side of the shell, but confined to the lines of sutures, of
obscurely tubular pillars: we have also something analogous in the
singular midribs, in Platylepas, causing the membranous basis to be
convex. I need only further add, that the parietes, in Chamæsipho,
are rather thin, and are composed of translucent shell, punctured for
the entrance of tubuli, with the punctures often arranged in straight
lines.
_Opercular Valves_ (fig. 4 _d_).--These are attached by a narrow
opercular membrane to the sheath, but little beneath the summit
of the shell. The _Scuta_ are considerably arched or convex: the
articular ridge is very prominent, and there is a thick strong
adductor ridge. The _Terga_ have a short, rather broad, rounded spur,
placed very nearly in the middle of the valve: the crests for the
tergal depressores are moderately developed, and are simple.
_Mouth._--The crest of the labrum is hairy, and is much hollowed
out. The palpi are small and narrow, with long bristles at their
apices. The mandibles have four or five teeth, with the lower part
pectinated. The maxillæ are notched. Of the _Cirri_, the second pair
is short, and all the segments are thickly clothed with bristles.
The third pair in the arrangement of the bristles resembles the four
posterior pairs. The segments in these pairs are elongated, and
support four pairs of main spines.
I was not able to observe any _Branchiæ_.
14. PACHYLASMA--_Nov. Genus._[133] Pl. 19, 20.
[133] Παχυς [Greek: Pachus] thick, and ελασμα [Greek: elasma], a
valve.
CHTHAMALUS. _Philippi._ Enumeratio Mollusc. Siciliæ.
_Compartments, when the shell is very young, eight; when mature, either
six, or in appearance only four owing to the close union of the lateral
compartments: basis calcareous._
_Distribution_, Mediterranean, and New South Wales; deep water.
The two species here included form a very natural genus, though,
as far as the shell alone is concerned, at first sight there is an
unusual amount of difference between them. This genus offers an
instance of a case, far from uncommon in nature, though so unfortunate
for the systematist, in which the most obvious and useful characters
of a group are completely masked. When I first examined _Pachylasma
giganteum_,[134] I did not doubt that it was a _Balanus_; and when
I first looked at _P. aurantiacum_, I thought, from there being in
appearance only four compartments, that it was an Elminius; in neither
case, from the absence of alæ to the rostrum, did I even suspect
that the species belonged to the sub-family of the Chthamalinæ.
But when I examined the included animal's body, I found, in both
species, the labrum bullate, not notched, with the palpi small, and
the mandibles with their lower teeth not laterally double. Again I
found in the third pair of cirri only the basal segments thickly
clothed with spines; and lastly, there were caudal appendages. Now
these characters are pre-eminently those of the Chthamalinæ; in fact,
they are those met with in the typical genus Octomeris, with the
exception of the presence of caudal appendages, and these occur in
Catophragmus,--a genus standing next to Octomeris, and in no other
genus of sessile cirripedes. Moreover, if we look to the shell of
Pachylasma, the absence of pores in the parietes, or at least of
symmetrical longitudinal ribs on their inner surfaces, and the peculiar
character of the narrow radii, hardly differing in structure from the
parietes, are characters which are rare in the Balaninæ, but universal
in the Chthamalinæ. Hence, taking the whole organisation of the two
species of Pachylasma, it is certain that they must be ranked amongst
the Chthamalinæ, though the leading character of the group, namely,
the rostrum being furnished, like the carina, with alæ, here fails.
Owing to this conviction, I examined very young individuals of _P.
giganteum_, and in specimens only the 1/100th of an inch in height, I
was interested by finding eight separate compartments, with the rostrum
having distinct alæ; hence, at this early age, as far as the shell is
concerned, this species may be said to be an Octomeris; and we have
seen that this likewise holds good with the included animal's body; as
the young shell increases in size the minute rostrum and rostro-lateral
compartments blend together (Pl. 19, fig. 5 _b_), without even traces
of sutures being left.
[134] Dr. Philippi called this species a Chthamalus; giving this
generic name from an examination only of the separated valves in a
fossil condition.
With respect to _Pachylasma aurantiacum_, I have no doubt that at an
early age it would possess a perfectly distinct rostrum with alæ; for
in the one specimen which I have seen, the compound rostrum is divided
by sutures, faintly visible, both externally and internally (Pl. 20,
fig. 1 _a_, 1 _b_), into three compartments (B, A, B), of which the
middle one, or true rostrum, still shows, in the manner in which it
underlaps the little rostro-lateral compartments, vestiges of alæ. I
may remark, that we have here the same structure as in Chelonobia,
formerly described, with the following differences, that here the
sutures pass through the outer lamina of the parietes, so that, as
seen externally, the separation of the three compartments is much
more perfect than in Chelonobia; on the other hand, internally, the
separation is less distinct, as the two rudimentary rostro-lateral
compartments do not form part of the sheath. _Pachylasma aurantiacum_
is further remarkable, from the two lateral compartments (C, D) on
each side, tending to blend together, being only separated by sutures
not more distinct, externally, than those separating the compound
rostrum, but more distinct internally, for they run up the sheath.
From these facts it follows, that this species, viewed outside,
without particular care, would be said to consist, like an Elminius or
Chamæsipho, of only four compartments; if the sheath alone were looked
at, there would be said to be six compartments; but when the walls,
especially their basal edges, are carefully examined, either internally
or externally, the eight compartments can be plainly distinguished.
With respect to the affinities of this genus, we have seen that both
in the included animal's body, and, at an early age, in the shell, it
is extremely close to Octomeris: in the presence of caudal appendages,
and in the basis being calcareous, we have a clear affinity with
Catophragmus; in the two lateral compartments of _P. aurantiacum_,
tending to become blended together, we have some relationship exhibited
to Chamæsipho, as is likewise shown in the structure of the second and
third pairs of cirri. If the genera of the Chthamalinæ were ranged in a
circle, Pachylasma would be the point of contact with the Balaninæ. I
must repeat, that it is extremely unfortunate that when the shell alone
of _P. giganteum_ is examined, it is hardly possible to separate this
genus from Balanus.
1. PACHYLASMA GIGANTEUM. Pl. 19, fig. 5 _a_-5 _d_.
CHTHAMALUS GIGANTEUS. _Philippi_ (!). Enum. Mollusc. Siciliæ, 1836.
_Shell and operculum dirty white: carino-lateral and lateral
compartments furnished with similar alæ._
_Hab._--Mediterranean; Sicily; deep water; often attached to the
_Millepora aspera_, and sometimes associated with _B. tulipiformis_.
_Fossil_ in the tertiary beds, near Messina. Mus. Brit., Cuming,
Stutchbury, Lyell, Philippi.
I owe to the kindness of Dr. Philippi, authentic specimens of his
_Chthamalus giganteus_, from the tertiary beds of Messina: had this
distinguished naturalist seen recent specimens, or a fossil one with
all the valves united, he would no doubt have perceived that this
species cannot be classed with Chthamalus. I am also indebted to
Sir Charles Lyell for some magnificent specimens, which he himself
collected near Messina.
_General Appearance._--Shell conical, rugged, irregular, with the
lines of growth plain; colour dead dirty white. Orifice large,
diamond-shaped, narrow towards the carinal end; notched. Radii
narrow, barely distinct from the parietes; in the same individual
sometimes absent, and sometimes forming a mere ribbon, confined to
the lower edge of a compartment. The diametric growth is effected
by the alæ, which seen externally are broad, and strongly marked by
lines of growth. Basal diameter of largest recent specimen 1.15;
height of highest compartment (a carina) 1.4; and width at base of
widest rostrum .85 of an inch. Amongst the fossil specimens, height
of highest compartment (a carina) 1.8, width of the same one inch;
width of widest rostrum 1.1 of a inch. One of these carinæ, a little
below the middle point, was actually .3 of an inch in thickness, from
which circumstance I have given the generic name of Pachylasma.
_Scuta_, triangular, but the width varies a little: growth-ridges
prominent, sinuous, with a few slight furrows radiating from the apex
of the valve. Internally, the articular ridge is not very prominent;
nor is there a deep depression for the adductor muscle, and none for
the lateral depressor muscles. The upper part of the valve projects
freely.
_Terga_, broad, broader than the scuta: the growth-ridges are
prominent, and angularly upturned close along the scutal margin; the
carinal half of the valve is smooth, with the faintest traces of
longitudinal striæ. A portion of the valve, nearly half of its entire
width on the scutal side, is slightly depressed below the general
level, and depends slightly beneath the basal margin on the carinal
side: this evidently forms the spur. The carinal margin is nearly
straight, with the lines of growth upturned along it. Internally, the
articular ridge in the upper part is extremely prominent: the crests
for the depressor muscles are very prominent, and depend beneath the
basal margin like a comb: they extend over nearly half the basal
margin, and the muscle, in a corresponding manner, is unusually
spread out.
_Structure of Shell and Radii_: the compartments are attached to
each other less strongly than in any other cirripede which I have
examined, so that when dried specimens are soaked in spirits of wine
they generally fall to pieces with a touch. In full and half-grown
specimens the carino-lateral compartments are nearly as broad as the
lateral compartments; in very young specimens, about 1/20th of an
inch in basal diameter, they are proportionally much narrower. The
_walls_ are strong even in young specimens; in old ones they attain a
thickness I have scarcely seen equalled except in Chelonobia. Their
internal surfaces are smooth, as is the basal internal margin in
young specimens, but in old specimens it is roughened with short,
blunt ridges and little points. The _Radii_ are often absent; when
present they are very narrow, and consist merely of a ribbon-like
portion, formed by obliquely upturned layers of growth, more
prominent than on the parietes. The _Alæ_ are very largely developed;
they are added to, during the diametric growth of the shell, in a
regular sweep all the way down to the basal margin, and consequently
they do not form a rectangular shoulder as is usual; externally they
are plainly marked by lines of growth: they are added to a little
above the line of attachment of the opercular membrane; their summits
are very oblique. The sheath has its basal edge slightly hollowed out.
_Basis_, solid, calcareous, very irregular, and of variable thickness.
I have, under the Genus, alluded to the structure of the rostrum:
in one shell, the basal diameter of which barely exceeded 1/20th
of an inch; the compound rostrum (being 5/200ths of an inch in
height), had its basal margin (being 9/200ths of an inch in width)
rendered deeply sinuous (see Pl. 19, fig. 5 _b_) by two indentations,
corresponding with and caused by two notches at the top of the valve.
These two notches extended down barely 1/100th of an inch (strictly
4/1000ths) from the summit and then disappeared; so that when the
shell was under 1/100th of an inch in height, (only one distinct zone
of growth having been formed), the now compound rostrum consisted
of three separate compartments; and there were eight compartments
altogether. Of the above three little compartments, the middle one,
or true rostrum, had large alæ, which could be most distinctly seen,
extending on both sides, under the little rudimentary rostro-lateral
compartments. These latter overlapped the compartments on both sides
of them, as in all the Chthamalinæ.
_Mouth_: labrum bullate, with no central notch; nor is the inner fold
of the labrum, forming the supra-œsophageal cavity, thickened, as in
Balanus: minute muscles run from this inner fold straight back to the
cavity formed by the outer bullate fold: the crest of the labrum is
hairy, with a row of the minutest bead-like points or teeth. _Palpi_,
small, broad, placed almost parallel to the sides of the mouth, with
their apices not nearly touching each other. _Mandibles_, with three
large nearly equal-sized _single_ teeth; the whole inferior angle
strongly pectinated. _Maxillæ_, small, with a broad, square notch
beneath the two or three great upper spines.
_Cirri_: first pair short, with the rami equal in length. Second
pair, with the anterior ramus having broader segments than those
of the posterior ramus, and with all the segments, except the few
uppermost, thickly covered with spines; the posterior ramus has
rather less than half the segments thickly covered. Third pair, very
slightly shorter than the sixth pair; anterior ramus with the lower
segments, less than half of the whole in number, thickly covered
with spines; posterior ramus with only the lowest segments, about
one fifth of the entire number, thickly covered; the other segments
of these two rami, and the upper segments of the posterior ramus of
the second pair, closely resemble in the regular arrangement of their
spines in pairs, the three posterior pairs of cirri. The pedicel of
the third pair supports numerous, irregularly scattered bristles. The
segments of the sixth pair bear four or five pairs of main spines,
with a few intermediate spines.
_Caudal Appendages._--Multiarticulate, narrow, tapering, situated on
each side of the anus: each segment has two little tufts of spines
on each side of its upper edge. These appendages are about one third
longer than the pedicel of the sixth pair of cirri: in a specimen,
in which the rami of the sixth pair had twenty-three segments, the
caudal appendages had nineteen segments.
_Penis_ short, hairy, finely-ringed, with no projecting point at its
dorsal basis. _Branchiæ_ moderately large, nearly circular, not much
plicated.
_Fossil Specimens._--With respect to the Sicilian specimens sent me
by Dr. Philippi and Sir C. Lyell, I can see no difference whatever
from the recent specimens, excepting in their greater size and
thickness; it must, however, be borne in mind that I have seen only
half-a-dozen recent shells. The one fossil scutum which I have seen
is rather broader than is usual with the recent, but I have seen one
nearly as broad. This species seems to have been extremely common
when the beds at Messina were deposited, and probably it attained
a larger size than it does at present. The compartments are always
found separated, which is easily understood, by the facility with
which, as above stated, recent specimens fall to pieces.
2. PACHYLASMA AURANTIACUM. Pl. 20, fig. 1 _a_-1 _d_.
_Shell tinged with orange; viewed externally seems formed of only four
compartments, owing to the carino-lateral and lateral compartments on
each side being separated only by an obscure fissure._
_Hab._--New South Wales; apparently from deep water, attached to
sandstone.[135]
[135] I am indebted to Mr. Bowerbank for this unique and
interesting species, which I have deposited in the British Museum.
_General Appearance._--Shell conical, smooth, with a tinge of orange
colour; orifice large, deeply notched, sub-triangular. Viewed
externally, the compartments in appearance are only four; but on
close examination, the lateral compartments are seen to be divided
by a very fine fissure into two nearly equal compartments. The
rostrum is broad and flat, and when carefully examined, it also is
seen to be divided by two fine fissures into three compartments;
of these the middle one, or true rostrum, is a very little broader
than the rostro-lateral compartment on each side; hence, on careful
examination, the shell is found to consist of eight compartments. The
carina is much compressed. The radii (in my one specimen) are not
developed. The carina, and the two lateral compartments alone, have
alæ; for the carino-lateral compartments are too closely joined to
the lateral compartments, and the true rostrum is too closely joined
to the rostro-lateral compartments, to have their alæ developed. The
four alæ which are developed, are very broad, widely exposed, and
marked externally by lines of growth. The basal diameter of my one,
apparently old specimen, is one inch.
_Structure of Shell, &c._--The inner surface (fig. 1 _b_) of the
rather thick parietes is smooth, except close to the base, where
it is roughened by a few irregular points. The sheath descends
low down, and has its lower edge slightly free. The sutures are
much plainer on the internal than on the external surface, and
can be here plainly seen to be eight in number. That portion of
the carino-lateral compartment, which helps to form the sheath,
is narrow, though the whole compartment is of very nearly equal
width with the lateral compartment. It is a singular fact, that no
portion of the rostro-lateral compartment helps to form the sheath;
for the alæ of the lateral compartments, overlap the whole upper
part of the rostro-lateral compartments, and abut against the true
rostrum. Hence, when the sheath alone is examined, the number of
the compartments appears only six. In a section the true rostrum
can be seen to underlie the rostro-lateral compartments, and thus
exhibits vestiges of alæ. The fact of the rudimentary rostro-lateral
compartments not forming a part of the sheath offers a marked
difference from Chelonobia, which otherwise has this part of the
shell very similarly constructed. The diametric growth of the shell,
which seems to be considerable, is effected by the four large alæ of
the carina and of the lateral compartment on each side. The sutural
edges of the alæ are added to in a regularly inclined line down to
the basis. The _basis_ is calcareous, and not very thin.
Both opercular valves (fig. 1 _c_, 1 _d_) closely resemble those of
the last species.
The _Scuta_ are remarkably narrow and elongated: the external
surface is slightly furrowed longitudinally, the prominent lines
of growth are much wrinkled. The articular ridge is blunt, and not
very prominent. The upper part of the valve is reflexed, and a
considerable portion must have projected freely. The _Terga_ are
nearly twice as broad as the scuta: the carinal half of the valve is
very smooth, and is coloured beautiful reddish-orange. The spur is
tolerably distinct. The articular ridge is not very prominent.
The _Mouth_ resembles that of the last species. The mandibles have
three sharp teeth, with the inferior part narrow and pectinated
with long spines: the edge, also, between the upper main teeth, is
pectinated with short spines, which latter often have their summits
crenated. The _maxillæ_ are notched; the inferior corner is produced
into a small step-formed projection. The outer maxillæ are more
pointed than is usual.
The _Cirri_ most closely resemble in every detail those of _P.
giganteum_; I can point out only one slight difference, namely, that
the lower segments, in the posterior ramus of the third cirrus, which
are thickly clothed with spines, are more numerous in proportion to
the upper segments with the spines arranged in regular pairs (being
as 4 to 14), than in _P. giganteum_, in which only one fifth (or 4 to
20) are so clothed.
_Caudal Appendages._--These are very small, not being more than
one fifth of the length of the pedicel of the sixth cirrus: their
segments are indistinct, and they support a very few coarse spines.
The rami of the sixth cirrus, in the one specimen, had twenty-five
segments, whilst each caudal appendage had only five. Hence the
caudal appendages are far less developed than in _P. giganteum_.
15. _Genus_--OCTOMERIS. Pl. 20.
OCTOMERIS. _G. B. Sowerby._ Zoological Journal, vol. 2, p. 244, July,
1825.
_Compartments eight: radii with their edges crenated: basis membranous._
_Distribution_, Cape of Good Hope; Philippine Archipelago.
The two species of this genus differ considerably in external
appearance, though not in essential character. Both, as the name
expresses, have eight compartments: the carino-lateral pair are
rather narrower than the lateral. The basis is membranous. The radii
are narrow, and are distinctly crenated on both sides of the sutures
with the teeth neatly interlocking; but these teeth can hardly be
distinguished in the large, corroded specimens of _O. angulosa_. The
crenated structure of the radii is identical with that described
under _Chthamalus dentatus_ and _Hembeli_; if, indeed, we were to add
carino-lateral compartments to the shells of these two species, they
would belong to Octomeris. I have seen only a few specimens of either
species of Octomeris, and none preserved in spirits; and therefore I
know nothing of the anatomy of the softer parts: I was not able to make
out distinctly any branchiæ. The cirri differ considerably in the two
species, in nearly the same way as in the two species of Chamæsipho. In
the structure of the second pair of cirri, and in the tendency of the
basal margin of the parietes to form bay-like indentations, _Octomeris
angulosa_ shows some special affinity to _Chamæsipho scutelliformis_.
Under Pachylasma, I stated that that genus was closely related to
Octomeris; and I have just alluded to the close affinity of the latter
to the division of the genus Chthamalus, which has crenated radii.
1. OCTOMERIS ANGULOSA. Pl. 20, fig. 2 _a_-2 _b_.
OCTOMERIS ANGULOSA. _G. B. Sowerby._ Zoological Journal, vol. 2,
July, 1825. And Genera of Recent and Fossil Shells, Plate.
---- STUTCHBURII. _J. E. Gray._ Annals of Philosophy, new series,
vol. 10, August, 1825.
---- AUGUBRA (?) _Chenu._ Illust. Conch., Tab. 4, fig. 2.
_Shell dirty white, rugged and massive: alæ thick, with their sutural
edges coarsely crenated._
_Hab._--Algoa Bay, Cape of Good Hope. Attached to littoral rocks;
often associated with _Balanus Capensis_ and _Chthamalus dentatus_;
Mus. Brit., Cuming, Stutchbury, Bowerbank.
_General Appearance and Structure of Shell._--Shell extremely rugged,
irregular, massive, generally much corroded, steeply conical or even
sub-cylindrical: orifice large, broad, rhomboidal, of nearly equal
breadth at both ends. Colour dirty white, often slightly tinted
yellow from the investing membrane, and from thin layers of punctured
membrane alternating with the laminæ of shell. The parietes, in old
specimens, have very irregular longitudinal ridges, or rather plates
projecting out, sometimes much branched, and generally curved inwards
so as to meet each other, thus forming round the basal margin a
circle of cylindrical apertures. In old large specimens the radii are
not developed, and till the compartments are disarticulated there
is no trace of the toothed structure of their sutural edges: in
this condition the sutures exist as deep, rugged, narrow fissures.
In younger shells, the radii, though narrow, are distinct, and have
their surfaces transversely ribbed, and their edges toothed and
interlocked with the teeth of the recipient furrow. Some of the
specimens present a curiously deceptive resemblance to _Elminius
plicatus_. Basal diameter of largest specimen, one inch and a
quarter; height one inch.
The parietes are remarkably thick, but the compartments separate
easily: I have, however, seen one instance in which they were
partially calcified together. Their internal surfaces are
very smooth. The sheath does not descend low. The alæ project
rectangularly; they have thick edges, and these are coarsely crenated
in transverse lines. Of the radii sufficient has been said.
_Scuta_ (fig. 2 _b_): these, in all the specimens seen by me but
one, have been deeply corroded, and their outline, as in Chthamalus,
considerably modified: in the one specimen well preserved, the
exterior growth-ridges were extremely prominent. The articular ridge
does not project much, nor is the articular furrow very deep. There
are more or less distinct crests for the lateral depressor muscles.
The _Terga_ are rather narrow, with a small rounded spur, moderately
distinct. The tips of the crests for the depressor muscle barely
depend beneath the basal margin.
_Mouth_: labrum considerably bullate, with the crest hairy and
furnished with a few most minute teeth. Palpi small, with their
tips not nearly touching each other. Mandibles with four teeth,
of which the lower ones are laterally double: inferior pectinated
portion small. The maxillæ are notched, but their outline differed on
opposite sides of the individual examined. Outer maxillæ bilobed on
their inner face.
_Cirri._--The first pair has the rami unequal in length by about
three segments, having six segments on one and nine on the other
ramus. In the second pair the anterior ramus is remarkably short and
extremely broad; the five segments of which it is composed being
thickly covered with bristles, some of which are very coarsely and
doubly pectinated: the posterior ramus is nearly twice as long as
the anterior ramus, and has eleven segments; of these, excepting the
two basal segments which are rather thickly covered with spines, the
others resemble in the arrangement of their spines the four posterior
pairs of cirri, with the exception that there is a single transverse
row of rather long spines along the upper and inner lateral edge
of each segment. The segments in all the four posterior pairs of
cirri resemble each other in bearing each four pairs of main spines
(of which the two lower pairs are short), with a tuft of small
intermediate spines.
The penis is remarkable from its very small size and shortness, not
being more than once and a half as long as the pedicel of the sixth
cirrus.
2. OCTOMERIS BRUNNEA. Pl. 20, fig. 3 _a_, 3 _b_.
_Shell reddish-brown, depressed, thin, finely furrowed longitudinally:
tergum with the basal margin having a slight angular bend._
_Hab._--Philippine Archipelago; rare; Mus. Cuming.
_General Appearance and Structure._--Shell circular, much depressed;
coloured brown, with an orange-red tint; surface regularly and
narrowly furrowed in longitudinal lines, the intermediate rounded
ridges projecting at the basal margin in finger-like points. The
eight compartments, when disarticulated, are of nearly equal sizes;
the carino-lateral and rostro-lateral compartments being rather
smaller than the others. The radii are neatly toothed; but with the
exception of these teeth, which are equally developed on both sides
of the sutures, and which closely interlock in the lower part of the
shell, but stand a little apart in the upper part, the radii can
hardly be said to be developed. The alæ do not form a rectangular
shoulder, the lower margin being regularly curved into the parietes;
the upper margin is only slightly oblique; the external surface
is furrowed by lines of growth. The shell increases a little in
diameter, chiefly by the growth of the alæ. The parietes are not very
thick, with the inner surface smooth, but with punctures placed in
rows for the entrance of the tubuli: the laminæ of shell alternate
with layers of yellow finely punctured membrane. Basis formed of thin
membrane, in concentric slips. Basal diameter of largest specimen, .6
of an inch.
_Scuta_ broad, with the lines of growth few in number, but extremely
prominent, so as to form folds: basal margin with a very slight
angular bend: articular ridge not very prominent. _Terga_, with a
narrow slip along the scutal margin, having the lines or folds of
growth so prominent as almost to form a series of small transverse
pits: the basal margin of this portion forms an angle with the rest
of the basal margin: no spur can be said to exist: the crests for the
depressor muscle depend a little beneath the basal margin: articular
ridge very prominent.
_Mouth_: labrum and palpi as in the last species. Mandibles with
three main teeth, apparently single, of which the lower tooth has
its upper edge pectinated: the inferior part of the mandible is
pectinated as usual. The maxillæ have two notches, one beneath the
two upper large spines, and the other in the middle, separating some
thicker and thinner spines.
_Cirri_: first and second pairs short, with the anterior rami in
each longer by about two segments than the posterior rami; all the
segments thickly covered with bristles. The third cirrus is much
longer than the second cirrus, with the posterior ramus longer than
the anterior ramus; on the latter the three basal segments, and on
the posterior ramus the two basal segments are thickly covered with
bristles; the other segments have bristles arranged as on the three
posterior pairs of cirri, namely, each segment has four pairs of main
spines, of which the two lower pairs are short.
The unarticulated support, whence the articulated portion of the
penis arises, is unusually long, equalling the pedicel of the sixth
cirrus.
16. _Genus_--CATOPHRAGMUS. Pl. 20.
CATOPHRAGMUS. _G. B. Sowerby._ Genera of Recent and Fossil Shells.
Plate.
_Interior compartments eight, with several exterior whorls of small
supplemental compartments: basis either membranous or calcareous._
_Distribution_, West Indies and Australia. Attached to littoral
shells and rocks.
This genus is very remarkable amongst sessile cirripedes, from the
eight normal compartments of the shell being surrounded by several
whorls of supplemental compartments or scales: these are arranged
symmetrically, and decrease in size but increase in number towards the
circumference and basal margin. A well preserved specimen has a very
elegant appearance, like certain compound flowers, which when half open
are surrounded by imbricated and graduated scales. The Chthamalinæ, in
the structure of the mouth and cirri, and to a certain extent in that
of the shell, fill up the interval between the Balaninæ and Lepadidæ;
and Catophragmus forms, in a very remarkable manner, the transitional
link, for it is impossible not to be struck with the resemblance of
its shell with the capitulum of Pollicipes. In Pollicipes, at least
in certain species, the scuta and terga are articulated together--the
carina, rostrum, and three pairs of latera, making altogether eight
inner valves, are considerably larger than those in the outer
whorls--the arrangement of the latter, their manner of growth and
union,--all are as in Catophragmus. If we, in imagination, unite some
of the characters found in the different species of Pollicipes, and
then make the peduncle so short (and it sometimes is very short in _P.
mitella_) that the valves of the capitulum should touch the surface of
attachment, it would be impossible to point out a single _external_
character by which the two genera in these two distinct families could
be distinguished: but the more important differences in the arrangement
and nature of the muscles which are attached either to the opercular
valves or surround the inside of the peduncle, would yet remain.
Although all the valves of the shell, even the eight in the innermost
whorl, are very thin, yet from their number in the successive whorls,
and from each being concave inwards, so as to form a cavity or tube
into which the corium enters, the total thickness of the sides of
the shell is very considerable. Both of the species of Catophragmus
occurred mingled, in the one case with _Tetraclita porosa_ and in the
other with _T. purpurascens_; now the walls of these shells, we know,
are very thick, and are permeated by several rows of pores, occupied by
threads of corium; seeing this, we may be permitted to believe, that
the several exterior whorls of valves in Catophragmus, between which
the corium is prolonged for some way upwards, are of service to the
animal, by thickening its shell, in an analogous, but not homologous,
manner, as in Tetraclita.
Considering the whole structure, external and internal, of
Catophragmus, with the one great exception of the exterior whorls of
valves, there is hardly a single generic character by which it can be
separated from Octomeris and Pachylasma; indeed, I am not quite sure
that it would not have been better to have run these three genera
together.
Of the two species, I will first describe _C. polymerus_, and
not the _C. imbricatus_ of Sowerby, inasmuch as I have plenty of
excellent specimens of the former, whereas the original specimens of
_C. imbricatus_, in the British Museum, consist of one old and not
perfect shell, without the opercular valves or the included animal's
body; and the other, though quite perfect, far from mature. As far as
these materials allow of minute comparison, the whole shell, with the
exception of the basis, and the opercular valves agree very closely in
the two species, whereas the included animal's body differs more than
is usual in nearly related species;--thus, _C. imbricatus_ has caudal
appendages, of which there is no trace in _C. polymerus_, and I have
seen only one other instance in which this organ was absent in one
species (_Scalpellum villosum_) and present in the other species of the
same genus. Under these circumstances it will be most convenient first
to describe in detail _C. polymerus_, and then only indicate the points
of difference in _C. imbricatus_.
1. CATOPHRAGMUS POLYMERUS. Pl. 20, fig. 4 _a_-4 _e_.
_Basis membranous: caudal appendages none._
_Hab._--New South Wales (Twofold Bay), Mus. Darwin; Swan River (?),
Mus. Cuming. Attached to littoral rocks and shells, and associated
with _Tetraclita purpurascens_, _Balanus nigrescens_, _Chthamalus
antennatus_, _Chamæsipho columna_.
_General Appearance._--Shell nearly circular, moderately or slightly
depressed; colour gray. The eight normal compartments of the inner
whorl are two or three times as large as those in the second whorl;
the other smaller compartments or scales graduate very regularly in
size, to mere beads, at the extreme basal edge, and are arranged
symmetrically. The general aspect of the shell depends chiefly on
the degree to which the surface has been disintegrated, and differs
greatly in the two extreme states. When well preserved, the general
appearance is very elegant; the scales are all neatly imbricated;
they terminate upwards in points, but with their extreme tips
generally broken; from laterally overlapping each other, their
external surfaces (as seen, when corroded, in fig. 4 _d_, or when
slightly exposed after the continued growth of their basal margins,
in fig. 4 _a_) become longitudinally keeled, often with a secondary
ridge or shoulder on one side; they are, also, crossed by rather
conspicuous and regular lines of growth, or more strictly, former
lines of union between the several valves: the orifice of the shell
in this perfect condition is sub-rhomboidal and notched, and the
scuta are united to the terga by nearly straight sutures. On the
other hand, when the shell has been considerably corroded, and this
seems to be the more common condition, the appearance is not elegant:
the scales in the successive whorls are not imbricated, but owing to
their upper parts having been worn down, they present a tesselated
surface (4 _d_), with the tesseræ graduated in size, and of a
peculiar shape, namely, a rectangle, with a more or less broad square
projection on the exterior side, together often with a large square
notch on one or both corners, caused by their laterally overlapping
each other. Some of the shells are so deeply corroded, that no
portion of the original surface is preserved, excepting the lowermost
bead-like scales; and the whole shell has so rugged an aspect,
that the successive whorls of the worn-down valves might easily be
overlooked. In the corroded specimens, the orifice approaches to
circular in outline, and is large and nearly entire: the scuta and
terga are deeply interlocked together. The largest specimen which I
have seen was one inch and a quarter in basal diameter.
_Structure of the Shell._--The eight inner normal compartments are
not thicker than the outer valves, and are far thinner than in
ordinary sessile cirripedes. They are arranged as in Octomeris.
The lateral compartments are broader than the rostro-lateral and
carino-lateral compartments. The shelly laminæ, of which these
compartments and the opercular valves are composed, alternate with
yellow membranous layers, exactly as is the case with Octomeris.
The lateral edges of the compartments, beneath the alæ, in the six
compartments having alæ, bend inwards, especially just above the
basis, so that the compartments in their lower parts (fig. 4 _c_)
stand much more separate than is usual. The basal edge of each is
irregularly toothed. The sheath presents no particular character.
The eight compartments have longitudinal shoulders and ribs on their
external surfaces, caused by the lateral overlapping and pressure of
the exterior scales. They are also marked by slight, transverse or
oblique calcareous ridges, caused by the attachment of the membrane,
by which they are united to the smaller compartments or scales
outside them. A new line of attachment, and consequently a new ridge
is formed, lower and lower down at each period of growth, as the
shell is added to at the basis,--in the same manner as new ridges are
added to the lower edge of the sheath at each period of growth. The
shell, excepting in old worn-down specimens, increases largely by
diametric growth: during the diametric growth, the outer scales must
be laterally separated a little from each other, and probably they
are laterally added to; but there are no distinct lines of suture,
or rows of smaller scales, corresponding with the sutures between
the eight inner compartments. The alæ do not project much; their
edges, as well as the shoulders into which they fit, are generally
irregularly crenated: they are added to during diametric growth
above the line of attachment of the opercular membrane. There is no
appearance of radii; but as the eight inner compartments are added to
laterally, and are often crenated, on the edges which correspond with
the radii in other Cirripedes, such edges must be considered as radii.
With respect to the scales in the several outer whorls, they resemble
each other except in size, and the outermost scales are reduced to
mere transversely elongated beads. Their basal edges are concave
inwards, being bent like the ridge of a house; hence sub-triangular
spaces or tubes, lined by the corium, run up between the scales. When
perfectly preserved, the outline of each scale is a much elongated
triangle, but usually, from their summits having been worn off, the
outline is nearly that of a parallelogram. Their basal edges are
dentated, and their upper parts, both on the outside and inside, are
marked, where joined to the other compartments, by slight calcareous
ridges: outside, there are longitudinal shoulders (Pl. 20, fig. 4
_b_), caused by the lateral overlapping of the adjoining scales;
these are best seen in section in corroded specimens (fig. 4 _d_):
on the inside there are, also, in the upper part, slight medial
longitudinal ridges, caused by the sutures, which the scales have
covered.
In large old specimens there are ten, or even more, whorls of
compartments, but it is scarcely possible to count them with
any accuracy. The first whorl consists of the eight large inner
compartments, though, homologically, it is doubtful whether the
rostro-lateral compartments in any sessile cirripede really belong
to the same whorl with the others. The second whorl consists of
eight smaller pieces, covering the eight sutures in the first whorl
(see the tracing of the basal edges of all the compartments and
valves in a very perfect specimen, Pl. 20, fig. 4 _c_). The third
whorl, in large and perfect specimens, consists of twice the number,
or sixteen, still smaller scales, corresponding with the sixteen
sutures of the second whorl; but sometimes there are less than
sixteen pieces, owing to some of the scales being large enough to
cover two adjoining sutures as well as the intermediate portion of
the compartments of the first whorl. In the fourth whorl, instead
of there being, even in the most perfect specimens, twice sixteen,
or thirty-two pieces, there are only twenty-four; this being caused
by single pieces (placed alternately with two pieces) being broad
enough to cover two sutures as well as the intermediate portion of
the compartment of the third whorl. In the succeeding whorls this
same arrangement seems to be the usual one, so that in the fifth
whorl, instead of there being, in the most perfect specimens, twice
twenty-four, or forty-eight pieces, that is, twice the number in the
last whorl, there are only thirty-six scales, or once and a half as
many scales.
The _Basis_ is thin and membranous; it firmly adheres to the surface
of attachment.
_Scuta_, nearly flat: the articular ridge is very prominent, and
there is a deep articular furrow both above and below; but the
precise outline of the ridge and furrows varies: there is a pit
for the adductor muscle, but no crests or marks for the other
muscles. The _Terga_ are remarkable from the extreme prominence of
the articular ridge and depth of the articular furrow. The basal
margin viewed internally seems straight, so that there appears to be
no spur; but viewed externally, when the crests for the depressor
muscles are seen to depend considerably beneath the true basal
margin, a slight, very broad spur may be perceived to exist. These
depending crests for the muscles are rather thin, but they extend
over half the basal margin of the valve.
_Mouth._--The labrum is very bullate, being as long in its
longitudinal axis as the rest of the mouth: crest hairy, with some
very minute teeth. Palpi truncated, with their apices not nearly
touching each other; thickly clothed with spines. Mandibles with
three large single teeth, of which the lower one has a single
fine tooth at its upper basal edge, showing a tendency to become
pectinated; inferior part short, coarsely pectinated. Maxillæ
notched, with a slight second notch and slight double prominence in
the lower part.
_Cirri._--First and second pairs short, with the rami in each
unequal in length by about four segments; on both rami in the second
cirrus, and in the shorter ramus of the first cirrus, there are some
coarsely pectinated spines. All four posterior cirri are alike; the
segments bear five pairs of strong spines, with a large intermediate
tuft of fine spines: the dorsal tuft is also large, consisting of
short thick, and long finer spines. There is no vestige of _caudal
appendages_, though present in the succeeding species.
Branchiæ moderately large, in area equalling the prosoma; surface not
plicated.
2. CATOPHRAGMUS IMBRICATUS.
CATOPHRAGMUS IMBRICATUS. _G. B. Sowerby._ Genera of Recent and Fossil
Shells, Plate.
_Basis calcareous: caudal appendages present._
_Hab._--Antigua, West Indies, attached to a _Tetraclita porosa_.
As stated under the genus, this species is known from two specimens
in the British Museum, one of which is full-sized, being three
quarters of an inch in diameter, but is destitute of the outermost
whorls, of the basis, opercular valves, and animal's body; the other
is perfect, but very young, being barely two tenths of an inch in
basal diameter. As far as the characters can be made out from these
materials, I can perceive no difference from _C. polymerus_ in the
shell, excepting that in the small specimen of _C. imbricatus_, there
would appear to exist fewer whorls. The opercular valves are likewise
closely similar: in the scuta, however, of the young specimen in the
present species, the articular ridge seems to be a little broader,
but this is so variable a character that no confidence can be placed
in it: these valves, moreover, externally have a broad furrow along
the middle, running from the apex to the basal margin, which is not
the case with the scutum of _C. polymerus_; but then I have often
seen, in young specimens of Balanus, a similar furrow, which is quite
absent in full-grown specimens. When we come to the basis we find a
good diagnostic character, for here it is calcareous: it is rather
thin, solid, and white; towards the outside it is pitted with small
cavities, corresponding with the small teeth on the basal edges of
the compartments. The latter adhere firmly to the basis. The central
internal surface is covered by an irregular network of imbedded
cement-ducts, some of which bifurcate. From the description here
given, it will be seen that any figure would have been superfluous,
the last species having been so well illustrated.
_Mouth._--I can point out no difference, excepting that the palpi are
here more oval or less truncated at their ends; and that the lower
corner of the maxillæ seems to be more prominent. In the _Cirri_, the
rami of the first and second pairs are nearly equal in length: none
of the spines are coarsely pectinated. In the four posterior pairs
of cirri, instead of a tuft of small spines on each segment between
the pairs of main spines, there are only a few minute intermediate
spines: the dorsal tufts are also here smaller, but are, as in the
last species, composed of short thick, and longer thinner spines.
_Caudal Appendages._--This is the only species of sessile cirripede,
with the exception of the two species of Pachylasma, which possesses
these organs: they are situated on each side of the anus in the usual
position: they are minute, equalling in length only the lower segment
of the pedicel of the sixth cirrus: in a specimen in which the rami
of the sixth cirrus had eighteen or twenty segments, these appendages
consisted of only three tapering segments, supporting a few thick
spines.
_Branchiæ._--I believe I discovered these, consisting of two minute
pouches, placed at the carinal end of the sack: if this observation
be correct, this species differs from _C. polymerus_ in the much
smaller size of these organs.
REMARKS ON BRONN'S LIST OF FOSSIL BALANINÆ AND CHTHAMALINÆ.
The following species of fossil Balanidæ are given in that most
useful work, the 'Enumerator Palæontologicus' in Bronn's 'Gesichte
der Natur:' it has appeared to me that a few words on each species,
might hereafter save others the trouble of searching through several
works.
_Tubicinella maxima_ of Morren, said to have been found in the Chalk:
this would have been a wonderful fact, considering that no true
sessile cirripede has hitherto been found in this formation, and that
it implies the existence of Cetacea at this period; but I have been
informed that the fossil in question is not a Cirripede.
_Diadema bifidum_ = Coronula bifida of Bronn, in his 'Italiens
Tertiär-Gebilde' (1831), p. 126 (no Plate). Without a much fuller
description I can form no judgment on this species.
_Diadema vulgare_ is probably the _Coronula barbara_, a Crag fossil
described by me.
_Pyrgoma undata_, Michelotti, in 'Bull. Soc. Geolog.' tom. x, p. 141,
a mere name without any description: probably it is a synonym of
_Pyrgoma sulcatum_, Philippi, 'Enum. Mollusc. Siciliæ,' which is a
synonym of _Pyrgoma Anglicum_ of the present work; found recent and
fossil.
_Acasta Montagui_ is probably the extinct _Acasta undulata_ described
by me.
_Chthamalus giganteus_ of Philippi, is the _Pachylasma giganteum_ of
this work; found recent and fossil.
_Chthamalus stellatus_, said by Philippi to be found fossil in
Sicily; such may be the case, but the littoral habits of the species
do not render it very probable.
_Balanus carbonarius_ of Petzholdt, found in the Carboniferous
formation! but I have given my reasons, in my 'Monograph on the
Fossil Lepadidæ,' p. 5, for disbelieving that this is a Balanus, or
even a Cirripede.
_Balanus ostrearum_ appears to be a mere name by Conrad, published
by Morton in his 'Synopsis of the Organic Remains of the Cretaceous
Group,' 1834, Appendix, p. 8.
_Balanus peregrinus_ is briefly described and poorly figured, without
the opercular valves, by Morton, in his 'Synopsis,' _ut supra_, p.
72, Pl. 10, fig. 5: this Eocene species apparently resembles the
Eocene _B. unguiformis_ of Sowerby, described in this work; but quite
indispensable details of structure for identification are not given.
Another figure of this shell is given in the 'American Journal of
Science' (N. S.), vol. i, Pl. 2, fig. 6.
_Balanus circinnatus_, _communis_, and _pustula_, of Defrance,
as well as all the other species named by him, are described so
imperfectly, that the descriptions are of no value whatever, every
description being applicable to every species: I must add, that the
_B. communis_ is not the _B. communis_ of British authors, a name
applied to several forms.
_Balanus Finchii_ is briefly described and figured, but without the
opercular valves, by Isaac Lea, in his 'Contributions to Geology,'
1833, p. 211, Pl. 6, fig. 222. I do not think that I have seen this
species.
_Balanus Holgeri_, Geinitz, 'Grundriss der Versteinerungen,' tab. ix,
fig. 19. No opercular valves are given; this species cannot be even
approximately recognised.
_Balanus proteus_, Conrad, 'Fossil Shells of Miocene Formation of U.
States,' p. 77, Pl. 44 (in 'Journal Acad. Nat. Sc.,' Phil., vol. vii,
p. 134). I cannot recognise this species; it resembles _B. porcatus_;
but as the radii are rather narrow, and apparently with slightly
oblique summits, it may be _B. concavus_; the opercular valves are
not figured.
_Balanus sagittata_ is merely a provisional name without any
description, given in a paper on the Crag by S. Woodward, in the
'London and Edin. Philosoph. Magazine, Brewster, Taylor, and
Phillips,' vol. vii, July-December, 1835, p. 354.
_Balanus sublævis_, J. de C. Sowerby, in 'Geolog. Trans.,' 2d series,
vol. v, Pl. 25, fig. 3. Plate extremely imperfect; description
extremely short and useless; a species from India not to be
recognised.
_Balanus balanoides_; the species thus named by Ranzani, and found
by Philippi in Sicily, certainly is not the true _Lepas balanoides_
of Linn., but may be _B. amphitrite_ of this work. In British
collections of Crag specimens I have found the _B. dolosus_ (nov.
spec.) thus named: I much doubt whether the truly littoral _B.
balanoides_ of Linnæus has been found fossil.
_B. costatus_ of Montagu, a synonym of _B. sulcatus_ of Bruguière,
and of _B. porcatus_ of this work; found fossil and recent. This
species was originally described by Linnæus under the name of _Lepas
balanus_. But the fossil _Lepas balanus_ of Brocchi is a different
species; if it be the same with the recent _Lepas balanus_ of Poli,
then it is the _B. perforatus_ of the present work. Again, the _B.
sulcatus_ of Bronn, in his 'Lethæa Geognostica' (tab. 36, fig. 14),
is quite different from the _B. sulcatus_ of Bruguière (_i. e._,
_B. porcatus_ of the present work), as is at once obvious from the
oblique summits of the radii. Lastly, _B. tesselatus_ of Sowerby is a
synonym of _B. porcatus_, _sulcatus_, _costatus_, and _Lepas balanus_.
_B. punctatus_ of Montagu is a synonym of _Chthamalus stellatus_, see
remarks on that species: the name of _B. punctatus_ is often applied
by British authors to varieties of _B. balanoides_, see remarks on
that species.
_B. rugosus_ of Morris's 'Catalogue' is a synonym of _B. crenatus_;
found recent and fossil.
_B. sulcatus_ of Bruguière, a synonym of _B. porcatus_; found recent
and fossil: see remarks under _B. costatus_.
_B. tintinnabulum_ of Linn., found recent and fossil; but no trust
whatever ought to be placed in the identifications of this species
given in several works; thus the _Lepas tintinnabulum_ of Brocchi is
distinct. The _B. crassus_ of Sowerby is the true _B. tintinnabulum_.
The _B. fasciatus_ of Dujardin (perhaps only a MS. name) probably
is also this species. The _B. crispatus_ is only a variety of _B.
tintinnabulum_.
_B. dentiformis_, Defrance: see _B. circinnatus_.
_B. ornatus_, Münster, 'Beiträge zur Petrifact.,' B. 3, p. 29, tab.
vi (1840). No opercular valves or details of structure of the several
species of Balanus named by Münster, are given, and consequently none
can be recognised with certainty.
_B. concavus_ of Bronn, fully described in this work, with the
synonyms given; recent and fossil.
_B. crassus_ of Sowerby, a synonym for _B. tintinnabulum_.
_B. latiradiatus_, Münster, probably a synonym of _B. tintinnabulum_,
see remarks under _B. ornatus_.
_B. pectinarius_, Bronn ('Italiens Tertiär-Gebilde,' p. 128), does
not appear to me fully enough described to be recognised.
_B. pictus_ of Münster, possibly a synonym of _B. amphitrite_: see
remarks under _B. ornatus_ and _balanoides_.
_B. plicarius_, Bronn: see remarks under _B. pectinarius_.
_B. porosus_ of Hausman (according to Münster of Blumenbach). I
cannot recognise this species in Münster's 'Beiträge.'
_B. pustularis_ of Lamarck, 'Animaux sans Vertèbres.' Scarcely one
of the fossil species of Balanus, described by Lamarck, can be
recognised; the descriptions are extremely imperfect. There is a
figure of _B. pustularis_ in Münster's 'Beiträge,' which makes me
think that this may be a synonym of _B. concavus_ of Bronn.
_B. pyramidalis_, Münster: see remarks under _B. ornatus_.
_B. rhomboicus_, Bronn: see remarks under _B. plicarius_.
_B. squamosus_, Defrance: see remarks under _B. circinnatus_.
_B. stellaris_, Bronn ('Lethæa Geognostica,' tab. 36, fig. 13). To
this species, _Lepas stellaris_ (I presume a misprint for _stellata_)
of Poli, is given as a synonym; but the _Lepas stellata_ of Poli is a
Chthamalus, and this certainly is not the case with _B. stellaris_. I
have received two specimens from the Continent named _B. stellaris_,
but they certainly differed from the form so called by Bronn, for
in that, the parietes are said to be porose and the radii very
narrow: these foreign specimens I have named _B. inclusus_. A species
described by me as _B. corrugatus_, resembles in external appearance
the _B. stellaris_ of Bronn, but it is mere labour in vain to attempt
identifying Balani by their external characters.
_B. striatus_, Defrance: see remarks under _B. circinnatus_.
_B. tertiarius_, Risso, 'Hist. Nat. de l'Europe Merid.,' vol. iv. I
cannot recognise this species.
_B. tesselatus_, Sowerby, a synonym of _B. porcatus_; recent and
fossil: see remarks under _B. costatus_.
_B. zonarius_, Münster: see remarks under _B. ornatus_; possibly this
is a synonym of _B. concavus_.
_B. amphimorphus_, Lamarck: see remarks under _B. pustularis_.
_B. crispatus_, _var._ of _B. tintinnabulum_.
_B. cylindraceus_, Lamarck. From Chenu's 'Illust. Conch.,' in which
work Lamarck's original specimens are figured; it appears that this
is the _B. psittacus_ of South America, where it is also found
fossil, but assuredly Lamarck is quite in error when he states that
a variety of this species occurs fossil near Turin. Bronn (in his
'Italiens Tertiär-Gebilde,' p. 127) gives as a synonym to the Turin
fossil the _Lepas tintinnabulum_ of Brocchi (in the 'Conchologia
Fossile Subapennina,' t. 2, p. 597), and this probably is correct;
and I have hardly any doubt that the _Lepas tintinnabulum_ of Brocchi
is the _B. concavus_ of Bronn described in the present work.
_B. perforatus_ of Bruguière is said by Philippi to be found in
Sicily: see remarks under _B. tulipa_.
_B. semiplicatus_, Lamarck: see remarks under _B. pustularis_.
_B. tulipa_ of Müller is the _B. Hameri_ of this work, under which
name full information on its geological history has been given.
The _B. tulipa_ of Poli and Ranzani is the _B. tulipiformis_ of
the present work; and this latter species is said by Philippi to
occur fossil in Sicily. But there has been so much confusion in
the identification of _B. tulipiformis_, _B. Hameri_, and _B.
tintinnabulum_, and likewise of _B. perforatus_ (which by Poli was
wrongly considered to be _L. balanus_, Linn.), that doubts must be
entertained about which have been really found, until the Sicilian
fossils are all carefully examined.
_B. ovularis_, Lamarck: see remarks under _B. pustularis_.
_B. Uddevallensis_, Linnæus, is probably a synonym of the _B. Hameri_
of this work and the _B. tulipa_ of Müller.
[A few other references may be added to those given by Bronn.]
_B. miser_, stated by Lamarck to be found fossil: see remarks under
_B. pustularis_.
_B. patellaris_ of Lamarck, is stated by Marcel de Serres to be found
fossil with several other species of Balanus; thus named without any
description in the 'Annales des Scien. Phys. et Nat. de Lyon,' tom.
i, p. 417.
_B. virgatus_, _delphinus_, and _crispus_, are names given by
Defrance, with absolutely worthless descriptions.
_B. radiatus_ is too briefly described by Risso ('Hist. Nat. de
l'Europe Merid.,' tom. iv, 1826), without a figure, to be recognised;
it probably is not the _B. radiatus_ of Spengler, Wood, and other
authors.
_B. goissopomo_, _lævis_, and _radiatus_, are mere names without
any description, published in a Catalogue by F. Hoeninghaus, in the
'Jahrbuch für Mineral. Geog.,' &c., 1831, p. 155.
_B. humilis_. Conrad, in the 'American Journal of Science,' vol. ii
(N. S.), p. 400, 1846, has given a short description, with a woodcut,
of this species, from the upper Eocene of Florida. The opercular
valves are not described, and I doubt whether the species could be
recognised.
2. Family--VERRUCIDÆ.
_Cirripedia without a peduncle: scuta and terga, not furnished with
depressor muscles, moveable only on one side, on the other side united
immoveably with the rostrum and carina into an asymmetrical shell._
The one genus herein contained differs so considerably from all the
others in the Order, in the extraordinary unequal development of
the two sides of the shell, that I have instituted a Family for its
reception. If compelled to place it in one of the foregoing families,
I should with much hesitation rank it in the sub-family Chthamalinæ,
rather than amongst the Lepadidæ; for it is destitute of a peduncle,
and has a shell, though a very different one from that of any true
sessile cirripede. In the interfolding sutures which may be considered
as representing radii or alæ, in the basis being divided into
concentric slips, and in the whole of the basis being attached to the
supporting object, this same line of affinity is clearly manifested.
On the other hand, in the general shape, manner of growth, and kind
of articulation of the scutum and tergum, there is so close an
approach to the Lepadidæ, that had I seen these very important valves
separately, I should certainly have concluded that they had come from a
Pollicipes, allied to certain Cretacean fossil species, as _P. fallax_
and _elegans_; it likewise, perhaps, deserves notice, that the upward
growth of the rostrum, in _Verruca nexa_, is a peculiarity found only
in the valves of the Lepadidæ. Verruca differs both from the Lepadidæ
and Balanidæ in the whole shell or external covering, having no other
muscle besides the adductor scutorum. In the characters derived from
the animal's body, Verruca approaches both families; but in the absence
of branchiæ, and in the great development of the caudal appendages,
perhaps it comes rather the nearest to the Lepadidæ. Whatever affinity
there is to the Balanidæ, it is much stronger to the sub-family
Chthamalinæ than to the Balaninæ; though the non-bullate labrum, in
three of the species, and the great dissimilarity of the third cirrus
from the three posterior pairs, at first seems to indicate a closer
relationship to the Balaninæ; but the labrum is never notched, as in
the latter sub-family, and in _V. nexa_ it is bullate, and supports
palpi of only small size. The dissimilarity, also, of the third pair
of cirri, compared with the posterior pairs, is hardly greater than
in _Chthamalus intertextus_ and _Chamæsipho columna_, members of the
Chthamalinæ, though abnormal in this one respect. Perhaps even a
special affinity is evinced between certain species of Chthamalus, as
_C. intertextus_, and certain species of Verruca, as _V. nexa_, namely,
in the interfolding sutures and in the very peculiar, inflected basal
margin of the walls. Upon the whole, the affinities of the Verrucidæ
are complex, and nearly equally divided between the two great families
of Balanidæ and Lepadidæ, or sessile and pedunculated cirripedes.
_Genus_--VERRUCA. Pl. 21.
VERRUCA.[136] _Schumacher._ Essai d'un Nouveau Syst. Class., 1817.
CLYSIA. _Leach._ Journal de Physique, tom. 85, July, 1817; _Clisia_,
Leach, Encyclop. Brit. Suppl., vol. 3, 1824; _Clitia_, G. B.
Sowerby, Genera of Recent and Fossil Shells.
CREUSIA. _Lamarck._ Animaux sans Vertèbres, 1818.
OCHTHOSIA. _Ranzani._ Memoire di Storia Nat., 1820.
LEPAS ET BALANUS AUCTORUM.
[136] According to Bock, in the 'Naturforscher' of 1778, this
term was used by Rumph for a Chelonobia, but as it was before the
adoption of the binomial nomenclature, according to the Rules, it
may be passed over, and does not interfere with the priority of
Schumacher.
_Distribution_, Northern Europe, Mediterranean, Red Sea, Madeira,
West Indies, Tierra del Fuego, Chile, Peru.
The shell in this genus is extremely unsymmetrical, not two of the six
pieces of which it is composed quite resembling each other. At first
it appeared hopelessly difficult to identify, in a homological sense,
these six valves, with those of ordinary cirripedes, but the difficulty
soon quite vanished. The operculum consists of two moveable valves on
one side, namely, a scutum and tergum, but without any moveable valves
on the opposed side: the scutum, though remarkable from being much
smaller than the tergum, can be easily recognised by giving attachment
to the animal's body and to the adductor scutorum muscle. The four
other pieces are articulated together, and form the shell surrounding
the sack, in which the animal's body is enclosed: of these, the two
against which the moveable scutum and tergum shut, are smaller, differ
greatly in shape, and are articulated together in a different manner
from the remaining two pieces; from these facts alone there would be
a strong presumption that they were of a different nature. The fixed
valve, against which the scutum shuts, is either furnished with a
remarkably prominent plate (_a_ in fig. 1 _c_, _s′_; compare this with
_s′_ in the reversed shell in fig. 1 _e_), or is hollowed out, as in
_V. nexa_, for the attachment of the adductor scutorum muscle. Thus
it is rendered probable that this fixed valve is a modified scutum;
but a surface of attachment for one end of the adductor muscle might,
perhaps, have been developed on any other valve, or a scutum might
have become fused with a lateral valve of the shell; the shell on
this latter view being rendered in idea more symmetrical. But when
a very young specimen is carefully examined, it is found that the
moveable and fixed scutum, the moveable tergum and its opposed valve
or fixed tergum, at the first period of calcification, resemble each
other quite closely; but that, as each zone of shell is added, the
differences become rapidly greater and greater: hence, it may be
considered as directly proved, that the two fixed valves (S′ and T′ in
all the figures in Pl. 21), which are opposed to the moveable valves
of the operculum (S and T), consist of an extraordinarily modified
scutum and tergum. It has been shown (p. 129), that at the period of
the metamorphosis, the two scuta, the two terga, and the carina of
the Lepadidæ, commence their growth, under the form of the so-called
"primordial valves," and so differ from all the other valves when such
occur: now, in two species of Verruca, I have found closely analogous
primordial valves on the apices of both the moveable and fixed scutum
and tergum (thus affording strong additional evidence that their nature
has been rightly interpreted), and on one of the two remaining valves,
namely, that at the posterior or carinal end of the shell. Hence, we
may safely infer, that this latter valve, which, though very much more
developed on one than on the other side, is so far medial as to curl
round and cover the line of opening between the moveable and fixed
tergum, is really a carina. The sixth valve differs only very slightly
in shape from the carina, and is directly opposed to it; therefore,
in accordance with all analogy, it must be the rostrum. Consequently,
the shell in Verruca consists of a moveable scutum and tergum, a fixed
scutum and tergum, a carina and rostrum, and, as we shall immediately
see, a membranous basis--the basis being, as in all sessile cirripedes,
the homologue of the peduncle in the Lepadidæ.
The moveable scutum and tergum stand at about right angles with the
fixed pair; and as these latter form a part of the wall of the shell,
which is always steep on this side, the moveable pair, which close the
orifice, are nearly horizontal or parallel to the basis and surface of
attachment. Hence, the animal's body, which is attached between the two
scuta, but nearest to the moveable scutum, also, lies nearly parallel
to the surface of attachment; and I was consequently at first led to
suspect that the basal membrane was one side of the shell in a modified
condition; but the presence of the prehensile antennæ of the pupa in
nearly the middle of this membrane, and the sheet of cement-tissue on
its under side, demonstrate that this membrane, though lying on one
side of the animal, is the true basis. To make all the parts in Verruca
hold the same position as in other cirripedes, relatively to the
surface of attachment, we must develope the carina and rostrum equally
on both sides of the true longitudinal axis of the shell, and insert
the newly-developed portion between the basis and the fixed scutum
and tergum, reducing the latter in size, and tilting a little up the
moveable scutum and tergum; and by this means the animal's body would
be turned, so that its dorso-ventral longitudinal plane would stand at
right angles to the basal membrane.
Extraordinarily great as is the difference between the right and
left sides of the whole shell, yet in all the species it seems to be
entirely a matter of chance whether it be the right scutum and tergum
with the right side of the rostrum and carina, or the left scutum and
tergum with the left side of the rostrum and carina, which become
abnormally developed. Nor does there seem to be any relation between
the side of the operculum to be attached, whether right or left, and
the nature of the surface of attachment; for I have seen many specimens
adhering to perfectly level surfaces, and to quite cylindrical branches
of Laminariæ; and in these cases, however the larva might attach
itself, there could be nothing to favour the development of one side
more than the other. Although the attached scuta and terga are larger
than the moveable pair, yet, owing to the small development of the
carina and rostrum on the attached side, the upper or unattached side
must be considered as the most developed. In this respect, and in
the circumstance of either right or left side being modified, we are
reminded of the structure of _Pœcilasma Kæmpferi_ (described in my
former volume on the Lepadidæ), in which the valves on the side of the
capitulum, nearest to the crab's body, to which the specimens were
attached, were somewhat less developed than those on the opposite side.
I may add, that in ordinary Crustaceans, as I am informed by Professor
Bell, the unequal development of the thoracic limbs seems quite
capriciously to affect either the left or right side of the body.
_General Appearance of the Shell._--The shell is in most cases much
depressed and irregularly circular; the side formed by the fixed
scutum and tergum is always steeper than the other side: the colour
is white or pale brownish, and in _V. nexa_ pale red. The surface is
naked. The size is small, rarely exceeding a quarter of an inch in
diameter, and the whole shell often appears like a mere scale on the
surface of attachment. The most remarkable feature in the external
aspect is due to the suture between the rostrum and carina, which is
formed by oblique, interlocking plates or folds; as all these plates
continue to be added to at their extremities during growth, the upper
plates become longer than the lower ones; and the plates on both
sides of the suture together form a triangular area, with the broad
end uppermost, somewhat like the radius of a sessile cirripede: they
act, also, like a radius, for their growth serves to separate these
two valves, and so adds to the diameter of the shell. The suture
between the rostrum and fixed scutum and that between the carina and
fixed tergum are nearly of the same nature, but the former is more
conspicuous than the latter; neither are so conspicuous as that between
the carina and rostrum: accordingly as the right or left scutum and
tergum are moveable, so the suture, second in plainness, (see Pl. 21,
fig. 1 _a_, and 1 _d_,) is placed to the left or right hand. The fourth
suture, between the fixed scutum and tergum, as viewed externally,
is straight, and so very obscure that it has been overlooked by some
authors, and the shell described as consisting of only three nearly
equal pieces, for the fixed scutum and tergum together are about equal
in size to the carina or rostrum. The orifice approaches more nearly
to an unequal-sided triangle, with the apex broadly truncated, than to
any other figure. The operculum fits with remarkable closeness, and
is surrounded by a slight rim, formed by the edges of the four other
valves.
_Moveable Scutum and Tergum._--The scutum (S in 1 _b_ and 5) is narrow
and very small, barely equalling half the size of the tergum, and
therefore proportionally much smaller than in any other cirripede; in a
very young shell, however, (of _V. Strömia_) less than a pin's head in
size, the scutum equalled the tergum in size. The valve is remarkably
thick; it is generally depressed down the middle; but in _V. nexa_
this part is longitudinally ribbed. The occludent margin is curved.
On the tergal margin there are two articular ridges (with a deepish
furrow between them), of which the upper one (′ in S, in fig. 1 _b_,
and 5) extends from the apex about half-way down the valve; and the
other, or lower articular ridge (′′ in S), generally runs down nearly
to the basal margin: an angle, running from the apex to the basi-tergal
corner of the valve, appears like a third articular ridge, but cannot
properly be considered such. The above two articular ridges interfold
with analogous ones on the scutal margin of the tergum, and so lock the
valves together. On the under side (fig. 1 _f_), the surface is bounded
along the occludent margin by a slight rim: there is generally a very
slight depression for the adductor muscle; but in _V. Spengleri_ there
is a straight, short, sharp (Pl. 21, fig. 2), prominent adductor ridge.
The _moveable tergum_ is broad and rhomboidal. Externally a prominent
axial ridge (′′′ in T, in fig. 1 _b_, &c.), which widens downwards,
runs from the apex of the valve to the basal point, and there
projecting slightly, causes the scutum to be indented; this indentation
on the scutum appears like a third articular ridge, lying beneath
(′′), S, in fig. 1 _b_, &c. Above the lower and axial ridge, on the
scutal margin of the tergum, there is a middle articular ridge, which
locks in, between the lower (′′) and upper articular ridges (′) of the
scutum (S). Again above the middle ridge there is an upper and third
articular ridge (′), which is either quite distinct, as in fig. 5, T,
or more commonly is formed by the occludent margin of the valve, as in
T, fig. 1 _b_. The broad extremity of this upper articular ridge is
often produced into a slight projection, or shoulder, and this always
underlies the scutum, of which the under and upper surface is indented
or furrowed (see fig. 1 _f_), in order to receive this shoulder. The
upper articular ridge of the scutum (′, S, 1 _b_) locks in between the
upper articular ridge or occludent margin (′, T), and the middle ridge
(′′, T) of the tergum.
Hence, altogether, there are three articular ridges on the scutal
margin of the tergum, the occludent margin being generally counted
as one; whereas, on the tergal margin of the scutum, there are only
two ridges, though, as before noticed, an outer indentation, which is
developed as a ridge in _V. nexa_ (fig. 5, S), might almost be counted
as a third articular ridge.
I may here just remark, that the furrow between the two ridges on the
tergal margin of the scutum, resembles the articular furrow in the
scutum of the Balanidæ; but it may be doubted whether the resemblance
be more than superficial, as this furrow, in the case of Balanidæ,
receives the edge itself of the tergum, whereas here it receives only
a ridge, proceeding from the apex of the tergum, to a nearly middle
point on its scutal margin. Finally, I may add, that the tergum in
this genus, in general shape, in growth (presently to be referred to),
in the manner in which the upper scutal shoulder is overlapped by
the scutum, and in the presence of the axial ridge, presents a very
striking resemblance to certain old fossil species of Pollicipes, and
to a limited extent to the living species of Lithotrya.
The scutum and tergum being interlocked, move together; they can be
firmly shut by the contraction of the long adductor scutorum muscle.
Their opening appears partly due to the elasticity of the membranous
hinge (representing the opercular membrane), by which they are attached
transversely, just beneath the summit of the carina and rostrum. No
doubt the protrusion of the cirri effectively aids the act of opening.
These valves are not capable (nor, of course, the other valves) of any
other movement; for there are no muscles for such movements.
_Fixed Scutum and Tergum._--The fixed scutum is larger than the fixed
tergum, and therefore has the same proportions as the homologous valves
in ordinary cirripedia, but reversed proportions compared with the
moveable scutum and tergum. The shape of neither valve can hardly be
described. The _fixed Scutum_ (S′ in all the figs.), externally, seems
at first to consist of two portions, namely, a curved occludent rim
(_a_), closely resembling the opposed occludent margin (_a_) of the
moveable scutum (S), having in fact undergone very little modification,
(as may be best seen in Pl. 21, fig. 1 _b_); and secondly, of a much
modified portion (marked ′′), which resembles in outline and state of
surface the rest of the walls of the shell, and may be called the
parietal portion. The rostrum (A) curls round the end of the occludent
portion, under an edge (_b_), evidently answering to the basal margin
(_b_) of the moveable scutum, and is simply united to this portion
by membrane, but beyond this part, it is articulated to the parietal
portion (′′) of the fixed scutum, by oblique interlocking ridges, like
those forming the suture between the rostrum and carina. In _V. nexa_,
however, (fig. 5) the rostrum (A) does not curl round any part of the
interlocking fixed scutum (S′), but articulates with it by a straight
suture. Internally, the fixed scutum (S′ in figs. 1 _e_, 1 _c_, of
reversed shells) has a surprisingly large, thin adductor plate (_m_ in
fig. 1 _c_, 1 _b_), with a rounded outline, projecting nearly parallel
to the basis or surface of attachment; the adductor scutorum muscle
is attached to its upper surface, and consequently the animal's body
lies between this plate and the moveable scutum. In the fixed scutum of
_V. nexa_, however, there is a deep pit, instead of a plate, for this
muscle.
The _fixed Tergum_, likewise, consists of two portions--a middle
and lower, or parietal portion, and a rim or upper portion; the rim
consists of two unequal arms, answering to the two upper margins (not
merely the edges) of the rhomboidal moveable tergum; the longer rim
(_x_ in T′, see fig. 5) answers to the carinal margin (_x_) of the
moveable tergum, and may be called the carinal rim; and the shorter
rim (_o_ and ′ in fig. 5) answers to the occludent margin (_o_ and
′) of the moveable valve, and may be called the occludent rim. The
carina curls round the end of the carinal rim, under an edge, _z_ (much
foreshortened in T′ in fig. 1 _b_, and best seen in fig. 5), answering
to at least a large part of the basal margin (_z_) of the moveable
tergum, and interlocks, by a serrated suture, with the edge of the
parietal portion of the valve. Internally (fig. 1 _e_, less plain in 1
_c_) there is a transverse ledge, notched in the middle, and sometimes
deeply hollow beneath, running across the valve in about the line of
the adductor plate of the fixed scutum: this ledge, in fact, marks
and is partly caused by, the line of separation between the central
or parietal, much modified, and the scarcely modified, upper or rim
portion of the valve. The use of this ledge is apparently to give
attachment, as does the under side of the adductor plate of the fixed
scutum, to ligamentous fibres, presently to be mentioned, by which the
shell is attached to the basal membrane: the carina and rostrum being
so much more gently inclined, do not stand in need of a ledge for their
attachment.
By comparing the moveable scutum and tergum with the corresponding
fixed valves, in all the species, the modification of the latter may
be clearly made out to have been effected as follows; and the case
appears to me a striking and interesting one. The moveable scutum
and tergum lie in the same plane, and are articulated by the means
of three ridges on the tergum (including the occludent margin), and
by two on the scutum. The fixed scutum and tergum have to be curved,
and to be greatly increased in size; and this is brought about, as we
shall see, by the large development of a certain small portion of each
valve. Comparing first the moveable tergum (T) with the fixed tergum
(T′), the umbo of growth matches the umbo of the four margins of the
moveable valve, the carinal (_x_), basal (_z_), and occludent (_o_ in
fig. 5), margins can be identified with certainty in the fixed valve,
from their close similarity in shape, their absolute apposition, or
correspondence in position. There remains only the scutal or articular
margin, with its three articular ridges; of these, the uppermost (′),
inasmuch as in most of the species it is hardly distinct from the
occludent margin, can, as we have just seen, be clearly identified, and
is overlapped, as it normally should be, by the upper tergal corner
of the fixed scutum: the second or middle articular ridge, though
not so distinct as in the moveable valves, can be plainly recognised
(′′), T′, in fig. 1 _b_, and 5; and it serves its normal function of
articulating the two valves together. But when we look in the fixed
valve for the third or axial ridge (′′′), we find in its exact place,
namely, extending from the umbo to the extreme opposite end of the
valve, between the second articular ridge (′′) and the basal margin
(_z_, see fig. 5), only that portion of the valve which I have called
the parietal portion; consequently, I do not doubt that this really is
the axial ridge largely expanded. So again in comparing the moveable
scutum (S) with the fixed scutum (S′); two of the three margins of the
former, namely, the occludent (_a_, see fig. 1 _b_) and basal (_b_),
can be identified without a doubt in the fixed valve: the third and
tergal margin remains; this should have two articular ridges; of these
the upper one, still serving its normal function, can be detected in
all the species (′ in fig. 1 _b_), and can be seen pretty plainly (′
fig. 5) in _V. nexa_: but of the lower and other articular ridge there
is no sign,--excepting indeed the whole parietal portion of the valve,
which, from holding an exactly homologous position with the lower
articular ridge of the moveable valve, I cannot doubt in this ridge
expanded and curiously metamorphosed. Hence, in both fixed scutum and
tergum, it is the outermost or lowest of the articular ridges which
has been modified and expanded, so as to rest on and be fixed to the
surface of attachment. It would appear as if it had resulted from one
ridge in each of these valves having been thus used up by expansion
(so to express myself), that the suture between the fixed scutum and
tergum is more simple than any other suture in the whole shell; and it
is owing probably to this straightness, and consequent _tendency_ to
weakness, that the valves do not grow along this line, and so do not
become separated from each other during growth, as on the three other
lines of suture. As it actually is, owing to this suture never being
separated, it is even stronger than the others; its edges on the inside
(fig. 1 _c_), I may add, are a little inflected or prominent.
_Rostrum_ and _Carina_: these valves differ from each other, only in
the former (A) being rather the largest, and in being more plainly
articulated with the fixed scutum, than is the carina (B) with the
fixed tergum. Their umbones stand in their normal places, at the
two ends of the orifice leading into the sack, that is, facing the
dorso-ventral longitudinal plane of the animal; but they are very
unequally developed on the two sides, and hence they rise very
obliquely from the surface of attachment. Their summits are nearly
square, which is caused by the continued growth on both sides of the
oblique plates or ridges, by which they are articulated with the
adjoining valves. These plates strikingly resemble, as already stated,
the radii in certain species of Chthamalus. Without these articulating
plates, the outline of the rostrum and carina would have been
triangular, with the apex upwards. In _V. nexa_, in which the walls
of the shell are almost perpendicular, the rostrum (A, fig. 5) is very
peculiar and patelliformed, with the umbo sub-central: this results
from the development of a border at the upper end of the valve. In this
same species, the basal edges of the rostrum, carina, fixed scutum and
tergum, are rectangularly inflected, so as to form a ledge round the
basis, as in the case of some few species of Chthamalus,--the ledge
appearing like part of the real basis. During the growth of the shell,
the upper internal ends of the carina and rostrum are either rendered
solid, or a ledge is formed on the inside across their summits, hollow
beneath, like the sheath of the Balanidæ, to which solid or hollow
ledge the basal margins of the moveable scutum and tergum are attached
by a rim of membrane, forming a hinge.
_Direction of Growth: Minute Structure of Valves._--The shell grows
downwards all round its basal margin. As far as the diametric growth
of its upper part is concerned, there may be said to be only three
valves, for the fixed scutum and tergum never become, as already
stated, separated; on the three other lines of suture, the valves are
added to on both sides; and thus the whole upper part of the shell,
and the orifice, increases in diameter. The moveable scutum and tergum
grow along their basal margins, and along the margins by which they
are articulated together; but the scutum in this latter respect, less
than the tergum. The summits of the moveable scutum and tergum, during
continued growth, become either worn away, or they project freely; in
this latter case, an internal ledge is added round the upper end of the
fixed scutum and tergum, so as to keep the orifice accurately closed.
In _V. nexa_ the rostrum, with its sub-central umbo, is anomalous, as
already stated, owing to a broad upper internal border growing in a
direction almost directly opposed to the basal growth of the moveable
opercular valves.
In young specimens, on the apices of both scuta and both terga, and
on the carina, but not on the rostrum, _primordial valves_ may be
distinguished, resembling the valves, so called, which first appear (p.
129, Introduction) after the metamorphosis in, the Lepadidæ. In the
Verrucidæ, however, they are calcareous; and the minute transverse
cylinders, of which they appear to be composed, stand further apart,
causing the surface of the primordial valve to be marked with little
separate circles, instead of by hexagons.
The shelly matter of which the valves are composed is translucent: it
is remarkably destitute of any investing membrane. The under surface
is marked with rows of minute approximate pores, parallel to the lines
of growth, into which the corium enters: after a portion of shell has
been dissolved in acid, these threads of corium are seen to change,
a short distance within, into cylinders of yellow chitine, running
obliquely through the substance of the valve. These cylinders are about
1/2000th of an inch in diameter, but in parts they are spindle-shaped
and twice as thick: they vary in length, about 1/100th of an inch
being the average length: these cylinders at their upper ends suddenly
contract into a point, more or less long, or are produced into a very
fine tortuous tubulus of chitine, imbedded in the shell: I have seen
in no other Cirripedes tubuli of this structure. There are other
ordinary tubuli, such as occur in the valves of most Cirripedes, about
1/6000th of an inch in diameter, and which sometimes alternate with
the above-described thicker cylinders. There are no external spines.
From the number and length of the tubuli of both kinds, the tissue left
after the action of acid is singularly complicated.
_Basis._--The basal membrane is thin, and is divided, but not very
plainly, into concentric slips, marking the successive increments of
growth. In the middle of it, in two young specimens, I found with great
difficulty the pupal prehensile antennæ: they were of small size,
measuring from the extreme edge of the main or second segment to the
end of the disc, only 27/6000ths of an inch: the disc appeared narrow
(as in Pollicipes and Scalpellum), with a single spine at the proximate
end: the ultimate segment, placed as usual at about right angles to
the disc, bore two groups of shorter and longer spines, but I could
not count how many. The antennæ were enveloped in a mass of cement of
a yellow colour, resembling in all its characters the cement of other
Cirripedes. In only one case, I believe I saw bifurcating cement-ducts,
of extreme tenuity, viz. 1/15,000th of an inch in diameter. The sheet
of cement on the whole under side of the basal membrane, not rarely
shows a very irregular reticulated structure. For convenience sake,
it will be best to defer the discussion on the very anomalous, though
slight, powers of excavation which this genus possesses, and which I
must attribute to the effects of some substance secreted probably by
the cement-organs. I will here only mention, that the specimens which
have excavated a depression, are less firmly attached than those, which
have not acted on their support; and that, in the former case, the
basal membrane, for a considerable space in the middle, becomes quite
detached.
_Animal's Body._--The body is much flattened and, owing to the
little development of one side of the shell, lies parallel to the
surface of attachment. The prosoma is but little protuberant. The
articulations of the thorax are unusually straight and transverse.
The _Mouth_ is also much flattened: it is placed rather distantly
from the adductor scutorum muscle, owing to the production of the
lower margin of the labrum. The _Labrum_ is not notched, or even
hollowed out in the middle, or (excepting in _V. nexa_) bullate;
its crest is surmounted by about eight (more numerous in _V. nexa_)
little teeth, or by some fine bristles. The _Palpi_ are of moderate
size, with their tips nearly meeting; they are slightly curved, and
have bristles only on their outer sides and extremities: they are
apparently capable of being lifted up and down by a muscle attached to
them, just outside the rounded swelling on each side of the labrum to
which they are articulated: in _V. nexa_, however, the palpi are very
small and narrow, and their tips do not nearly meet. In this genus,
therefore, we find the swollen state of the labrum and the size of
the palpi--characters generally invariable and of high classificatory
importance--variable. The mandibles have three upper main teeth,
with two or three minute lower teeth, or, in _V. nexa_, with the
lower part pectinated with small spines: in _V. Strömia_, I have seen
traces of the second tooth being laterally double--a character of
some importance. The _Maxillæ_ have a notch under the upper pair of
large spines, with the lower part bearing, as usual, a double row of
bristles, and forming a large step-formed projection: these organs are
furnished with the usual apodeme and muscles. The _Outer Maxillæ_ are
prominent, and deeply lobed on their inner surfaces, the two lobes
being clothed with bristles.
_Cirri._--The first pair are attached, as usual, on each side of the
mouth, and stand some way apart from the five posterior pairs. The
second and third pairs differ considerably in structure from the three
posterior pairs, which are much elongated. The _first pair_ (excepting
in _V. nexa_) is short, with the two rami slightly unequal in length,
and with the segments thickly clothed, as usual, with spines. The
_second pair_ is remarkable from the posterior ramus being more than
twice as long, and containing thrice as many segments, as the anterior
ramus, which is barely as long as the shorter ramus of the first pair:
the segments in the anterior ramus of the second pair (only five in
number in a full-sized specimen) are broader and more protuberant in
front, and more thickly clothed with spines (the terminal spines being
doubly pectinated), than are the segments on the posterior ramus; on
the latter, the uppermost segments have their bristles arranged in
front in simple pairs, with the dorsal spines long, the lower segments
being more thickly clothed with bristles, owing to the development of
lateral rows. The _third pair_ resembles in every respect the second
pair, except in being a little longer, and in the bristles on the
posterior ramus being less crowded, more resembling the arrangement of
those on the posterior cirri. In _V. nexa_, however, there is not so
great an inequality in length or dissimilarity in structure in the two
rami of the second cirrus, and only a very slight difference of any
kind in the two rami of the third pair. _Fourth_, _fifth_, and _sixth
pairs_ have numerous elongated segments, bearing four or three pairs of
long slender spines in front, with a single minute bristle between each
pair, and with two or three slender spines in the dorsal tuft.
There is a considerable amount of variation in the proportional
length, and in the number of the segments, of the several cirri in
_V. Strömia_; in some specimens the two rami of the fourth pair were
unequal in length; in some, nearly all the cirri on the lower or
attached side were shorter than those on the upper side.
_Caudal Appendages._--These are of most unusual length, sometimes
even exceeding those of _Ibla quadrivalvis_, which surpasses, in this
respect, all other cirripedes. They arise on each side and over the
anus. They consist of numerous (sometimes as many as twenty-four),
unequal, cylindrical, thin segments, bearing, at their upper ends,
a circle of long and very slender spines. They sometimes equal two
thirds or even four fifths of the length of the sixth cirrus; but
their length, and the number of their segments, (sometimes imperfectly
divided), varies much in different specimens of the same species,
and sometimes even on opposite sides of the same individual. In some
very young shells, as big as a pin's head, the caudal appendages were
proportionally extremely short, and consisted of only two or three
segments. No muscles enter these organs; and when the animal is taken
out of its sack, they project straight out behind, instead of being
curled in, like the cirri.
_Anatomical Structure._--The animal's body is attached to the two
scuta by the adductor scutorum, and by the other usual muscles running
towards the mouth, and surrounding the prosoma. The whole external
covering or shell has no other muscles; Verruca thus differing from the
Balanidæ and Lepadidæ; but the shell is attached all round, near its
circumference, to the basal membrane, by a band of very short fibres,
appearing like muscles, but really ligamentous, as determined for me
by Professor Quekett. _Branchiæ_ are entirely absent. The _alimentary
canal_ presents all the usual characters, but in the prosoma is rather
abruptly bent back on itself. The orifices of the two _olfactory
pouches_ are not at all prominent; they are placed directly under the
outer maxillæ, (homologically in their middle segment), just above
a small, medial, tongue-like apodeme. The orifices of the _acoustic
sacks_ appeared to be in their usual position beneath the basal
articulations of the first pair of cirri. The _vesiculæ seminales_
occupy their usual position in the prosoma; they are not much
convoluted; they unite before entering the penis. The probosciformed
penis is imperfectly ringed; it is thick and short, and tapers much
more abruptly than is usual; it supports a few very thin hairs. The
_ovarian cæca_ are spread over the basal membrane, at the bottom of
the sack; hence they in fact lie almost on one side of the animal: they
consist of two main trunks, proceeding out of the animal's body at the
rostral end of the sack, which then branch and inosculate. In specimens
of _V. Strömia_ collected by Mr. Peach for me, in Cornwall, during the
first week of April, there were included two ovigerous lamellæ, placed
transversely across the rostral and the carinal end of the sack: the
lamellæ were .11 of an inch in length; they appeared loose and not
attached, as in the Lepadidæ, to any ovigerous fræna. The ova, in their
earliest age, have one end much pointed, and are 8/1000ths of an inch
in length; they become blunter and increase a little in size before
the larvæ burst forth. The larvæ, both during their earliest stage and
after the first moult, have been excellently figured and described[137]
by Mr. C. Spence Bate: they present no particular characters distinct
from the larvæ of other Cirripedes. I will only further add, that
the structure of the prehensile antennæ still adherent to the basal
membrane, indicates that the larva in its last stage,--that is the
locomotive pupa,--has a normal character.
[137] 'Annals and Mag. of Nat. Hist.,' 1851, Pl. 7, fig. 8-10.
_Affinities._--These have been sufficiently discussed under the family;
I need here only remark that all the species, with the exception of _V.
nexa_, are intimately allied together.
_Range--Habits--Geological History._--The genus Verruca ranges, being
represented by four species, from Iceland to Cape Horn. The species
that is found in Tierra del Fuego extends up the west coast to Peru.
Our northern form, _V. Strömia_, (if I may trust a specimen in the
British Museum, apparently ticketed in an authentic manner), occurs
also in the Red Sea; and this is the only locality in the eastern
hemisphere whence I have seen this genus. The species seem generally
to live in rather deep water: I procured _V. lævigata_ from nineteen
fathoms, on the east coast of Patagonia: _V. Strömia_ is found,
according to information given me by Professor Forbes, on the British
shores, between five and fifty fathoms, and on the steep shore off
Mull, in ninety fathoms; but Mr. Thompson assures me that he once
saw it adhering to tidal rocks and likewise to some floating bark.
Generally the species are attached to living organic bodies, especially
shells of Mollusca and of Cirripedes, to Gorgoniæ, and Laminariæ;
less frequently to rocks. We shall immediately see that it has slight
powers of excavation. This genus is geologically older than any true
sessile cirripede or member of the Balanidæ: _V. Strömia_ is found in
the Glacial Deposits and in the Red and Coralline Crag of England:
another species (in a state not to be identified) occurs in the ancient
Tertiary formations of Patagonia; and another in the Chalk of England
and Belgium. The fact of this Family ascending to a Secondary epoch
accords, in an interesting manner, with its affinities; inasmuch as
though in appearance a sessile cirripede, it is almost equally related
to the Lepadidæ and Balanidæ, and is more nearly related to the
Lepadidæ than to the Balaninæ, or typical members of the Balanidæ: of
the latter, none have hitherto been found in any Secondary deposit,
whereas the Lepadidæ culminated during the Cretacean period.
_Powers of Excavation._
My attention was called to this subject by Mr. Hancock, whose excellent
researches on the boring of Mollusca are well known. _Verruca Strömia_,
when attached to shells destitute of an epidermis, excavates, as he
informed me, a slight depression, deepest in the middle; but when the
epidermis is present no effect whatever is produced. We shall presently
see that the central depression is in some degree distinct from that
of the circumference. I have since found Mr. Hancock's observations
strictly applicable to _V. lævigata_, _V. Spengleri_, and to an ancient
tertiary species from Patagonia. From having found that the following
cirripedes, viz., Lithotrya, Alcippe, and Cryptophialus, all form
their deep excavations by mechanical means, and from having read the
above-mentioned memoirs by Mr. Hancock on the boring of mollusca, I
was strongly impressed with the idea that the action in Verruca would
likewise prove mechanical: but from the following facts I have come to
the conclusion that the excavation must be due to a solvent, probably
poured out from the cement-ducts, which debouch on the under side of
the basal membrane.
In the first place, an epidermis, as just stated, perfectly preserves
the shells of the various species of mollusca and certain cirripedes,
to which I have seen Verruca attached: this is well shown by comparing
the effect produced on the same shell in parts covered by the epidermis
and in parts whence it has been abraded; or where the shell of the
Verruca had fixed itself, whilst very young, within a crack in the
epidermis, and had subsequently, by its growth, turned up the edges,
and had then acted on the underlying shell; whereas the specimens
attached to the sound epidermis had not produced the smallest effect.
Again, I have seen an epidermis-covered mussel-shell encrusted by a
hard nullipora, on which _V. lævigata_ was attached; and here the
calcareous nullipora, under the middle of the basal membrane, was
entirely corroded away, whilst the underlying epidermis and the shell
beneath it, were not in the least affected. The protection afforded
by the epidermis is still more strikingly shown by contrasting shells
with very sharp prominent ridges, when thus invested and when naked,
to which Verrucæ have been attached: I have given a figure (Pl. 21,
fig. 6) of a piece of an invested Venus, from the surface of which a
_V. Spengleri_ had been just removed; on the other hand, I have seen a
Peruvian Discina in which even sharper ridges, covered by epidermis,
were left absolutely untouched, although projecting deeply into the
shell of an attached _V. lævigata_. I have seen several specimens of
this latter Verruca (which has the power of corroding naked shell as
deeply as its congeners), attached to the membrane-covered variety of
_Balanus lævis_, the shell of which was thus perfectly preserved: now
this membrane is little more than the 1/2000th of an inch in thickness;
it is not hard, and so brittle that it generally separates with the
Verruca, leaving the underlying shell of the _B. lævis_ with its
lines of growth glossy and perfect: it appears to me impossible that
a membrane so thin and brittle could resist an action, if mechanical,
which has worn away from twenty to forty times as great a thickness of
hard shell; but the thinnest film of any matter on which acid does not
act, as of grease in certain forms of printing, will perfectly preserve
the underlying substance, and as I have ascertained by putting on a
drop of acid, is the case with this membrane. I have removed several
scores of shells of _V. Strömia_ from the stems of Laminariæ, and when
the latter were washed and slightly dried, generally not the least
effect could be seen, except that the spots where the shell had adhered
were glossy from the still adherent basal membrane: yet the stems of
Laminariæ are far from hard. In some cases, however, the attachment of
the Verruca seemed to have produced a very slight depression on the
Laminaria, but this, I think, may be safely attributed to the growth
of the surrounding surface; for I have seen exactly the same effect
produced by the attachment of the discs of the antennæ of a Lepas,
and these discs, with their long spines, could not possibly produce
any excavation; nor is Lepas or its pupa in any case a burrowing
animal. Again, I have seen a few specimens of Verruca attached to
Gorgoniæ, and they had not acted in the least on the horny axis. I
have examined numerous specimens of _V. Strömia_ attached to three
pieces of slate-rock, and to one piece of red sandstone, all from
different localities, and no effect whatever had been produced; yet
the slate-rock, especially in one instance, was soft. Mr. Bate, to
whom I am indebted for some of these specimens, also informs me that
he could discover no impressions on the slate-rocks, whence specimens
of the Verruca had been removed. On the other hand, I have had two
specimens of limestone, with attached Verrucæ, one coarse and very
impure, and the other hard and marble-like; and in both cases there was
a distinct central slight cavity, including loose gritty matter. The
loose particles evidently resulted from the unequal action either of a
solvent or of some mechanical power on the rock, for it is improbable
in the highest degree that the shells should have fixed themselves
exactly over small collections of loose particles, even if such could
possibly have remained on projecting surfaces of sea-washed rocks.
The above facts seem to indicate pretty plainly that the excavation
of the support does not depend on its hardness, but on its containing
calcareous matter, liable to be acted on by some solvent: but as this
view, considering what we know of Lithotrya and of the two other
burrowing genera of cirripedes to be hereafter described, appears
improbable, I will add a few additional observations. I most carefully
examined the shell and basal membrane of Verruca, and likewise the
tissues left after the dissolution of the shell in acid, and could
detect no structure at all fitted for boring; and what appears more
important, there was no apparent difference in the state of the
specimens which had and had not excavated a hollow; and this, I think,
would certainly have been the case (as in Lithotrya) if the action had
been mechanical. It is not easy to ascertain, owing to the small effect
at any time produced, at how early an age Verruca begins to act on
its support; but I found two sets of specimens only 1/20th of an inch
in basal diameter, which had certainly commenced. The ribbed shell,
(Pl. 21, fig. 6), especially the middle rib, shows, in a somewhat
exaggerated degree, the _typical_ form of the excavation; it may be
here seen that the excavation is of the same depth for some little
distance from the circumference towards the centre, but that in the
middle it suddenly becomes deeper. I have seen several specimens with a
central hollow, without any, or with scarcely any, marginal depression,
and likewise the reversed case. These several facts show that the
central excavation cannot be due to an equable action, prolonged
during the whole growth of the shell, having thus affected the middle
more than the circumferential parts, for in this case the excavations
would have sloped into each other. In specimens which have not at all
acted on their support, the whole basal membrane is firmly attached,
as in all ordinary cirripedes, to the supporting surface; but in those
which have acted, the middle portion of the basal membrane is quite
unattached, and the circumferential portion is, I think, less firmly
attached than is usual; but between these two portions, there is a
circular zone strongly cemented to the supporting surface, and which
alone keeps the shell in its place. Now, on the mechanical theory,
to account for the circumferential hollow, the basal edges of the
shell together with the circumference of the basal membrane must be
subjected to movement, but the shell is united to the basal membrane
by corium and by transparent structureless chitine (both of which may
be left out of question) and by a circle of short fibres, which adhere
at their lower ends to the firmly cemented circular zone, and by their
upper ends to the shell; and these fibres have been very carefully
examined by Professor Quekett, and pronounced to be not muscular,
but exclusively ligamentous, and therefore incapable of moving the
edge of the shell. The basal membrane over the central hollow is, as
stated, quite loose: its lower surface, formed by a reticulated layer
of horny cement-tissue, shows no signs of abrasion, and the membrane
is so brittle and tender, that in specimens which have been once dried
and then well soaked, it almost invariably cracks when the shell is
removed, owing to its mere adhesion to the delicate inner tunics of
the sack; yet on the mechanical theory, the wearing of the central
hollow must have been caused by the action of this middle portion of
the basal membrane,[138] which, it may be repeated, is destitute of
muscles. From the presence of the prehensile pupal antennæ, enveloped
in cement, nearly in the centre of the basal membrane, it is certain
that this spot was originally attached to the supporting surface, and
has since been detached from it; as, moreover, the central hollow goes
on increasing in diameter with the growth of the shell, it is certain
that the inner edge of the firmly attached circular zone of basal
membrane must likewise continually go on becoming detached: it may,
then, be asked by what force can the basal membrane, seeing that it
is united to its own shell above only by fibres of ligament near the
circumference, be continually torn away from the underlying support,
to which it is strongly cemented? On the other hand, on the theory
of a solvent slowly poured out from the cement-ducts, its separation
from its support is simply explained. It might be supposed that the
calcareous matter, when dissolved, would not be able to escape from the
central hollow, owing to the basal membrane being so firmly cemented
all round it; but the attachment is by a reticulated layer of cement;
and I infer that it must be permeated by open passages, from the fact
of the hollow being often filled, in dried specimens, by a bubble of
air, instead of the basal membrane being pressed closely down into the
hollow, as would have been the case had the hollow been hermetically
sealed up. I have seen a few instances in which the bottom of the
central hollow was occupied, (as was remarked to me by Mr. Hancock),
by a little chalky and gritty matter; and in the case of one of the
specimens of calcareous rock, before alluded to, by coarse grains and
oxide of iron; this seems quite compatible with a solvent acting more
readily on certain parts of the rock or shell than on other and less
soluble parts or particles.
[138] Mr. Hancock suggests to me that the basal membrane, on the
mechanical theory, need not itself move; the motion of epithelial
scales, were they transferred into cutting agents, might be
supposed to be sufficient. But of such scales, though I used very
high powers, I could see no trace; and their presence on the under
side of the layer of cement seems hardly possible. Moreover,
according to Von Siebold ('Anatomie Comparée,' tom. 1, p. 412),
ciliary action has not been observed in any Crustacean, or indeed
any Articulate animal. This same statement is likewise made in
Annals and Mag. of Nat. Hist. 1854, p. 136, by Dr. T. Williams.
The greatest depth of the central hollow, in any specimen seen by me,
even measuring from the top of a rib in the case of a ribbed shell,
to the deepest point, was only 1/50th of an inch; but considering how
much depressed the shell of Verruca is, I have no doubt that this small
gain of space is of service to the animal: we must suppose the loose
middle portion of the basal membrane is stretched slightly, or splits
and is repaired, so as to fit the hollow. With respect to the even
much slighter circumferential excavation, it barely equals in depth
the thickness of the extreme edges of the walls; it must, I presume,
give strength to the shell when _laterally_ pushed; but it certainly
appeared to me that the individuals which had excavated a depression
for themselves, could be pried vertically up much more easily than
those which had not acted on their support. Finally, we must suppose
that the hypothetical solvent is poured out of the cement-ducts at
the extreme circumference of the basal membrane, which is almost
loose and destitute of cement, so as to slightly corrode outwards and
downwards the calcareous support; the action here then stops, and this
rim of basal membrane becomes, after a new rim has been formed and
as the shell grows outwards, firmly cemented down to the now slightly
excavated surface of attachment; but during all the time the solvent
goes on acting in the middle, and continues, during the whole growth
of the shell, to encroach on and dissolve the supporting surface from
under the inner edges of the previously cemented down, circular zone of
basal membrane. I have discussed this subject at considerable length,
as it appears to me an interesting one. In this case we have the action
of ciliæ[139] and of respiratory currents, to which in the case of
Mollusca so much has been attributed, entirely eliminated. It is, also,
an interesting fact, that within the same Order we should have some
Cirripedes boring by simply mechanical means, and others by a chemical
solvent.[140]
[139] See the previous note to p. 516.
[140] The solvent may be carbonic acid gas, as suggested by Mr. C.
S. Bate in the case of Mollusca ('Report of British Association,'
1849, p. 73), but here, under the basal membrane, we cannot have
the respiratory currents, or the ciliary action (see note, supra),
as likewise suggested by Mr. Bate.
1. VERRUCA STRÖMIA. Pl. 21, fig. 1 _a_-1 _f_.
LEPAS STRÖMIA. _O. Müller._ Zoolog. Dan. Prod., No. 3025, 1776.
---- ---- _Ib._ Zoolog. Dan., vol. 3, Tab. 94, 1789.
---- STRIATA. _Pennant._ British Zoology, vol. 4, Tab. 38, fig. 7,
1777.
DIE WARZENFORMIGE MEEREICHEL. _Spengler._ Schriften der Berlin.
Gesell., 1 B., Tab. 5, fig. 1-3, 1780.
LEPAS VERRUCA. _Spengler._ Skrifter af Naturhist. Selskabet, 1 B.,
1790.
---- ---- ET STRÖMIA. _Gmelin._ Syst. Nat., 1789.
BALANUS VERRUCA. _Bruguière._ Encyclop. Meth., 1789; _Clisia
verrucosa_, Deshayes, in Tab.
---- INTERTEXTUS. _Pulteney._ Catalogue of Shells of Dorsetshire,
1799.
LEPAS STRIATUS. _Montagu._ Test. Brit., 1803.
---- VERRUCA. _Wood's_ General Conchology, Pl. 9, fig. 5, 1815.
VERRUCA STRÖMII. _Schumacher._ Essai d'un Nouveau Syst. Class., 1817.
CREUSIA STRÖMIA ET VERRUCA. _Lamarck._ Animaux sans Vertèbres, 1818.
OCHTHOSIA STROEMIA. _Ranzani._ Memoire di Storia Nat., 1820.
CLISIA STRIATA. _Leach._ Encyclop. Brit. Suppl., vol. 3 (sine
descript.), 1824.
CLITIA VERRUCA. _G. B. Sowerby._ Genera of Recent and Fossil Shells,
Plate.
VERRUCA STRÖMII. _J. E. Gray._ Annals of Philosophy (new series),
vol. 10, Aug., 1825.
_Moveable scutum, with the lower articular ridge not half as broad as
the short upper articular ridge: shell generally ribbed longitudinally._
_Var., with the shell not longitudinally ribbed._
_Hab._--Shores of Great Britain and Ireland, Shetland Islands;
and, according to various authors, Denmark, Iceland, and shores of
northern Europe. Red Sea, Brit. Mus. Attached to shells, laminariæ,
rocks, crabs, and floating bark, from low tidal mark to fifty or
ninety fathoms.
_Fossil_ in Glacial deposits of Scotland, Mus. Lyell; Red Crag
(Walton, Essex), Coralline Crag (Sutton), Mus. S. V. Wood.
I have given so full a description of the genus that little remains
to be said under the species. Generally the whole shell is covered
(independently of the interfolding, oblique, articulating plates) by
narrow, longitudinal ridges or folds; and by this character alone the
ordinary variety of _V. Strömia_ can be distinguished (as far as I
have seen) from all the other species. The shell is white or dirty
yellowish-brown. The scutum has the lower articular ridge on its tergal
margin very narrow (but somewhat variable in width), appearing like a
mere slight shoulder, against which the longitudinal axial ridge of
the tergum abuts: it is not half as wide as the short, upper articular
ridge. On the under side there is a very slight depression for the
adductor scutorum muscle. There is considerable variation in the degree
to which the transverse ledge on the under side of the fixed tergum
projects, and therefore in the depth of the hollow thus formed. The
specimens with the right-side, and those with the left-side opercular
valves moveable, are apparently about equally numerous.
The specimen in the British Museum, from the Red Sea, was attached
to a Gorgonia, and was in the same box with a Pyrgoma--circumstances
favouring the correctness of the locality--but I am much surprised
from the general distribution of the species, that _V. Strömia_ should
occur in so distant and isolated an area. After careful examination,
I can discover no constant difference between the Red Sea and British
specimens.
The specimens from the Crag have not their moveable opercular valves,
which offer much more important diagnostic characters than the shell;
but as far as the latter is concerned, no difference whatever can be
perceived from _V. Strömia_.
2. VERRUCA LÆVIGATA. Pl. 21, fig. 3 _a_, 3 _b_.
VERRUCA LÆVIGATA. _G. B. Sowerby._ Genera of Recent and Fossil
Shells, Plate.
_Moveable scutum, with the lower articular ridge broader than the short
upper articular ridge; moveable tergum broader than high, with the
upper articular ridge produced into a point._
_Hab._--Tierra del Fuego; Eastern Patagonia, nineteen fathoms; Chile;
Peru; Mus. Brit., Cuming, Stutchbury, Darwin: attached to shells, and
often to _Balanus lævis_ and _psittacus_.
I can point out no difference in the shell between this species and
_V. Strömia_, excepting that its walls seem invariably to be smooth,
which is rarely the case with _V. Strömia_; perhaps also the oblique
interfolding articular plates between the several compartments are here
more prominent. It appears that specimens with the left side uppermost,
and therefore with the left opercular valves moveable, are considerably
more common than those with the right valves moveable. The moveable
scutum and tergum are articulated together by much more prominent
articular ridges than in _V. Strömia_, and the two valves together are
broader in proportion to their height,--the height being measured from
the apex to the basal margin. In the scutum the lower articular ridge
is considerably broader than the short upper ridge. In the tergum, the
basi-carinal corner is more rectangular, and the whole valve is nearly
square: owing to the deep furrow receiving the lower articular ridge of
the scutum, the axial ridge of the tergum is proportionally narrower
but more prominent than in _V. Strömia_; the uppermost ridge (formed by
the occludent margin of the valve) projects, especially when viewed on
the under side (fig. 3 _b_), as a moderately sharp point.
In the _mouth_, the lower teeth of the mandibles are more distinct than
in _V. Strömia_; the lower part of the edge of the maxilla is very
prominent. In the second and third pairs of cirri the terminal spines
on the shorter rami are coarsely pectinated; on the sixth pair there
are only three pairs of main spines on each segment; but these several
points, according to the analogy of other species, I should expect to
be variable.
This species is alluded to by Bruguière, in the 'Encyclopédie
Méthodique,' but was confounded by him with the _V. Strömia_ of Europe.
3. VERRUCA SPENGLERI. Pl. 21, fig. 2.
_Moveable scutum, with a sharp, straight, medial adductor ridge: fixed
scutum not larger than the fixed tergum._
_Hab._--Madeira, Mus. Lowe; attached to shells.
It would appear that the present species does not attain quite so
large a size as the more northern _V. Strömia_; the walls are not
longitudinally ribbed as is usual with this latter species. The
proportional sizes of the compartments seem to be somewhat different;
the fixed scutum is either equal to or even smaller than the fixed
tergum, instead of being larger, as in _V. Strömia_; but in young
individuals the proportions are reversed. In several specimens the
fixed scutum and tergum together were larger than the carina. The
rounded adductor plate of the fixed scutum is extremely large. The
lines of growth, especially on the moveable opercular valves, are
rather more plainly crenated than in _V. Strömia_. In the moveable
scutum the lower articular ridge on the tergal margin varies a little
in size, and is sometimes larger than in _V. Strömia_ (but never so
large as in _V. lævigata_), and is placed more in the middle of the
tergal margin: but by far the most important character by which this
species can be distinguished from all the others, is the presence,
on the under side of the moveable scutum, of a straight, prominent
adductor ridge, which runs up to and even under the apex of the valve,
for it is there slightly hollowed out. In the moveable tergum, owing
to the medial position of the lower articular ridge of the scutum, the
middle of the scutal margin is more hollowed out, and the axial ridge
narrower, than in _V. Strömia_.
In the animal's body the only difference which I could perceive was
that the shorter rami of the second and third pairs of cirri were not
so short, compared either to the other cirri or to the longer rami of
these same cirri. In the second cirrus, in a moderately-sized specimen,
the segments were six and thirteen in number in the two rami, and in
the third cirrus, seven and fifteen.
Had it not been for the specimen in the British Museum of _V.
Strömia_, from the Red Sea, I should have concluded, from geographical
considerations, that _V. Spengleri_ probably was the species found
in the Mediterranean, and noticed by Spengler ('Schriften der Berl.
Gesell.,' 1 B., 1780), as a small variety of the northern _V. Strömia_;
and likewise that it was the _Creusia echinoides_ of Risso ('Hist. Nat.
Product. de l'Europe,' tom. 4, p. 382, 1826), which is certainly a
Verruca, but not described with sufficient minuteness to be recognised.
4. VERRUCA NEXA. Pl. 21, fig. 5.
_Shell reddish: moveable scutum, with three strongly prominent
longitudinal ridges, besides the articular ridges: fixed scutum larger
than the carina, with no distinct adductor plate._
_Hab._--West Indies, Mus. Brit.; attached to a Gorgonia.
This species differs considerably from all the others in the genus.
The shell is brownish-red, tinted yellow: it is not at all depressed
like the former species, but the walls are almost perpendicular or
even overhang their bases, and the summit of the shell consequently
is broad. This form may be in part, but only in part, due to the
attachment on the thin branches of the Gorgonia. The umbones of the
compartments are remarkably prominent and sharp. Although the parietes
are nearly smooth, yet from being so steep, they are little seen, and
owing to the very prominent but rounded ribs by which the compartments
and opercular valves are articulated together, the whole shell has a
strongly ribbed appearance. The diameter of the largest specimen was .2
of an inch.
The _rostrum_ (A, fig. 5) is patelliformed, with the umbo of growth
sub-central, but rather above the middle point; hence this valve,
differently from the carina, and differently from the rostrum of the
other species, grows not only at its basal margin, and on both sides
where opposed to the carina and fixed scutum, but also along its upper
margin where opposed to the basal edges of the moveable scutum and
tergum: owing to the perpendicularity of this valve, the upper part
forms a ledge almost parallel to the orifice of the shell. The carina
(B) is of unusually small size, being about only half the size of the
rostrum, and scarcely exceeding in size the fixed tergum. The _fixed
scutum_ (S′) is large, larger even than the carina; it is oblong, and
its shape is more simple than in the other species; this is chiefly
owing to the rostrum articulating with the whole of that margin (_b_)
which answers to the basal margin of the moveable valve; whereas
in the other species (fig. 1 _b_) it curls beyond this margin, and
articulates with the very protuberant, so-called, parietal portion of
the valve. Three or four rounded prominent longitudinal ribs, exactly
like the homologous ribs on the moveable scutum, run from the apex
of the fixed scutum to the basal margin, and their extremities form
the teeth by which it articulates, as just stated, with the rostrum.
Its upper articular ridge (′) is more prominent, and placed much
lower down in the suture between it and the fixed tergum, than in
the foregoing species. The ledge (_o_) by which the orifice is kept
neatly closed, is here more distinct than in _V. Strömia_: this ledge
is necessary, as well as in the case of the fixed tergum, owing to
the altered shape of the summits of the moveable scutum and tergum,
due to their corrosion and to their coming to project freely. But the
most remarkable character of the fixed scutum is, that on the under
side there is no great adductor plate, but a rounded hollow with its
lower edge only slightly prominent; the absence of the adductor plate,
which is present in all the other species of the genus, is no doubt due
to the under side of this valve being inclined even outwards, and so
standing in some degree opposed to the moveable valve; thus affording
on its under surface a place for the attachment of the lower end of
the adductor scutorum muscle; whereas in the other species this muscle
could not possibly have been attached, without the aid of an adductor
plate, to the under side of the much depressed and sloping fixed
valve. The _fixed tergum_ (T′) is a little more simple in form than
the corresponding valve in the other species; the two arms, answering
to the occludent and carinal margins of the moveable tergum, are more
nearly equal in length: the internal transverse ledge, separating these
rims or margins from the parietal portion of the valve, is but little
developed.
All four valves forming the shell are remarkable from having, when
full-grown, but not whilst young, their basal edges abruptly inflected
inwards, thus forming a ledge all round the basal membrane, as in
_Chthamalus intertextus_ and _Hembeli_.
_Moveable Scutum._--This is slightly larger in proportion to the
tergum than in the foregoing species: it is chiefly remarkable from
the presence of three prominent longitudinal ridges on the main part
of the valve, like the two articular ridges on the tergal margin; of
these latter, the lower one extends down to about the middle of the
tergal margin. The _moveable tergum_ is rhomboidal, with the whole
carinal portion marked only by lines of growth: it is only remarkable
by the upper of the three articular ridges on the scutal margin being
unusually distinct from the occludent margin.
With respect to the animal's body, its several peculiarities have
already been pointed out under the genus. The _labrum_ is decidedly
bullate, triangular in section, with a row of minute bead-like teeth on
the crest; the palpi are very narrow and short, and do not nearly touch
each other: this variation in the structure of the labrum and in the
size of the palpi, is very remarkable, considering how important, in a
classificatory point of view, these parts are in all other Cirripedes.
In the _mandibles_ there are either two or three main teeth, with the
whole lower part of the organ pectinated with sharp spines. _Cirri_:
the first pair is not short; in the individual examined, the two rami
had eleven and twelve segments. In the second pair, the shorter ramus
was two thirds of the length of the longer ramus, the segments being in
number ten and fifteen; in the arrangement of the spines this second
pair resembles its homologue in the three other species. In the third
pair, the two rami are very nearly equal in length, having sixteen and
eighteen segments; and the segments of the anterior ramus are only a
little thicker and more thickly clothed with spines than those of the
posterior ramus. The remaining cirri and the caudal appendages are as
in the other species.
5. VERRUCA PRISCA. Pl. 21, fig. 4.
VERRUCA PRISCA. _Bosquet._ Monographie des Crustacés fossiles du
Terrain Crét. de Limbourg, Tab. 1, fig. 1-6; 1853.
_Shell smooth: moveable scutum, with the lower articular ridge somewhat
broader than the upper articular ridge._
Fossil--'Système Senonien et Maestrichtien,' Belgium, Mus. Bosquet;
in Chalk, Norwich, Mus. J. de C. Sowerby.
M. Bosquet has admirably figured and described the several separated
valves belonging to this species, and I owe to his great kindness an
examination of some of them. In Mr. J. de C. Sowerby's collection,
also, there is a single specimen, attached to a Mollusc, with the four
valves of the shell united together, but without the two moveable
opercular valves; it cannot be positively asserted that this is the
same species with that of M. Bosquet, but such probably is the case.
This is the species to which I alluded in the Introduction to my
'Monograph on Fossil Lepadidæ.' It is an interesting species, from
being the only known Secondary one, but in itself it is a very poorly
characterised form, and I can point out no important character in the
shell by which it can be recognised. The rostrum and carina, which are
of nearly equal sizes, are locked together by the usual interfolding
plates, and likewise to the fixed scutum and tergum; but these latter
plates seem to have been less developed in M. Bosquet's specimen than
in the English. The fixed scutum has a large adductor plate, which
seems to have been chipped in M. Bosquet's specimen; this valve and the
fixed tergum in all essential respects resemble the same valves in _V.
Strömia_. The surface of the shell is very smooth.
The _moveable scutum_ has its occludent margin considerably arched:
the lower articular ridge is broader than the upper ridge, in which
respects it resembles the same valve in _V. lævigata_, but the whole
valve is not so broad as in that species. There is no adductor ridge on
the under surface. The _moveable tergum_ has its upper articular ridge
narrow, and slightly produced into a point on the scutal margin: in
this latter respect this species also resembles _V. lævigata_, but the
whole valve is not so broad in proportion to its height.
3. Family LEPADIDÆ.
_Cirripedia having a flexible peduncle, provided with muscles: scuta
and terga, when present, not furnished with depressor muscles: other
valves, when present, not united into an immoveable ring._
This Family has been fully treated of in my former volume, published
by the Ray Society,[141] and I should here only have alluded to its
existence, had it not been for the genus Alcippe, which differs in
so many important characters from the other members of the Lepadidæ,
that formerly I did not even suspect that it could belong to this
Family, and therefore deferred its examination. The genus Alcippe was
discovered, well described and illustrated, in 1849, by Mr. Hancock;
to whose very great kindness I am indebted for permission to dissect
and examine his entire stock of this truly remarkable Cirripede. In the
classification of the whole class I have not felt so much doubt, as
whether I ought to institute a family for the reception of this genus.
Alcippe differs from all other Cirripedes (putting on one side for the
instant, the males and complemental males of Ibla and Scalpellum) in
the very singular fact of being destitute of a rectum and anus;--in
the three segments of the thorax, which usually support the second,
third, and fourth pairs of cirri, being without any appendages;--in the
fifth and sixth pairs of cirri having their inner or posterior rami
metamorphosed into very singular roughened cushions or buttons, which
apparently serve to triturate the food;--in the caudal appendages being
muscular, and being used conjointly with the cirri;--and lastly, in
the pupa having a lesser number of segments in its abdomen and caudal
appendages than in (as far as I have seen) any other Cirripede. It will
be thought that these characters are amply sufficient to justify the
placing Alcippe in a separate family, more especially when the close
general resemblance in the animal's body in most of the other members
of the Balanidæ, Verrucidæ, and Lepadidæ, is borne in mind. On the
other hand, the males and complemental males of Scalpellum and Ibla
must indisputably be considered as members of the Lepadidæ; yet the
male of _Scalpellum vulgare_ and _ornatum_ has no stomach, anus, or
mouth, which is a far more abnormal structure than the absence only of
the anus in Alcippe: the cirri, also, in these same males, differ from
the ordinary cirripedial type decidedly more than in Alcippe. Again,
in the male of Ibla, all the cirri, excepting the fifth and sixth
pairs, are aborted, and these two pairs are usually only uniramous;
here, then, we have a decided resemblance to Alcippe. Hence, if we
might assume that the female Alcippe had partially assumed characters
confined to the males of the other genera, it would assuredly stand
amongst the Lepadidæ. Independently of this comparison with the
foregoing males, the affinities of Alcippe are so special to several
genera amongst the Lepadidæ, that it seems unnatural to force it out
of the position which it well occupies between Ibla and Anelasma, and
place it in another family by itself: thus, in being bisexual, and
in the general character of its very curious males, Alcippe shows an
affinity to Ibla and Scalpellum; and to the former of these genera it
is related in several particulars, such as in the body being lodged
within the peduncle, and in the structure of the larval antennæ, &c.:
to Anelasma and Alepas it is allied in the general character, and to a
certain extent in the muscles, of the capitulum; Anelasma, also, has
all its cirri to a certain degree rudimentary, and _Alepas cornuta_ has
the inner rami of the fifth and sixth pairs of cirri,--namely, the very
same rami which are so curiously modified in Alcippe,--small, destitute
of muscles, and functionless for their proper purpose: to Anelasma
and Lithotrya it is allied in the peculiarity of the lower end of the
peduncle becoming elongated by growth, and in being imbedded; and to
Lithotrya by its powers of excavation and manner of attachment. Now,
I believe it generally holds good that when a form is really distinct
from another group, its affinities are general, or only in a slight
degree special to the members of that group. Nor, indeed, can it be
asserted that Alcippe differs much more, somewhat more it certainly
does, from the other genera, than does Anelasma, with its more singular
mouth, spineless rudimentary cirri, and fimbriated peduncle; and I have
never regretted having included this genus amongst the Lepadidæ. Hence,
after much consideration, I have resolved to consider Alcippe as one of
the Lepadidæ, though so curiously modified,[142] and having characters
confined to the males of the other genera. Perhaps I have been in some
degree influenced by the difficulty of finding external characters by
which to separate Alcippe as a family from the other Lepadidæ.
[141] The fossil species have been described in a separate
Monograph published by the Palæontographical Society. Since
its publication, M. Bosquet has produced an excellent memoir,
containing descriptions, with the most beautiful illustrations, of
several new Cretacean species of Pollicipes and Scalpellum. The
memoir is entitled a 'Monographie des Crustacés Fossiles du Terrain
Crétacé, du D. de Limbourg.'
[142] Adrien de Jussieu, in his 'Memoir on the Malpighiaceæ,'
'Archives du Museum,' tom. 3, p. 86, when speaking of the
characters afforded by the degraded flowers, which in certain
genera are borne together with ordinary flowers, makes the
following observations bearing on the question here discussed,
viz., whether or not to include Alcippe amongst the Lepadidæ.
"Ces exemples peut-être aideront à comprendre comment à des
genres d'une organisation assez compliquée, viennent quelquefois
s'en rattacher d'autres d'une organisation beaucoup trop simple
en apparence, membres appauvris et dégradés d'une même famille,
qui lui appartiennent sans la représenter; comment le type,
s'y présente comme effacé, ne conservant plus pour se laisser
reconnaître que quelque trait isolé, mais caractéristique, dont la
valeur, essentiellement ordinale, peut être ainsi constatée." Under
the point of view, so strongly and admirably insisted on lately by
Milne Edwards ('Annales des Sciences Nat.,' 3d series, tom. 17), of
describing types without regarding whether the different members
blend together on their confines, perhaps Alcippe should be raised
to the rank of a Family: I feel quite unable to decide how properly
to act.
But we shall presently find, when we come to Cryptophialus, that all
the above difficulties, great as they are, are greatly enhanced,
for Cryptophialus is certainly allied in a very direct and curious
manner (in decided opposition to the remarks just made on special
affinities) to Alcippe, and yet in all the more important parts of its
organisation, and in its metamorphosis, it differs so fundamentally,
that I have felt myself obliged to form not merely a Family, but a
distinct Order for its reception.
_Genus_--ALCIPPE. Pl. 22, 23.
ALCIPPE. _Hancock._ Annals and Mag. of Nat. Hist., vol. 4, 1849, Pl.
8, 9.
_Fem.--Capitulum without valves, with the orifice spinose: peduncle
with the basal end added to during growth; its rostral surface
depressed and covered by a horny disc: capitulum and peduncle imbedded
in a cavity excavated in the shells of molluscs._
_Labrum very large, with a row of long hairs on each side: palpi
rudimentary: mandible one-toothed: second, third, and fourth cirri
absent: fifth and sixth cirri with the posterior ramus represented by a
button-like body: caudal appendages four jointed, muscular: anus none._
_Males,--several, adhering to the upper end of the horny disc of
the female: capitulum naked, transparent, elongated, with a small
orifice at the end: peduncle lobed, with the lower end extending far
beyond the pupal antennæ: eye, testis, and vesicula seminalis single;
probosciformed penis very long: mouth, stomach, thorax, abdomen, and
cirri none._
ALCIPPE LAMPAS, _Hancock_ ut suprà.
_Hab._--North-eastern shores of England, fifteen to twenty fathoms,
imbedded in dead shells of _Fusus antiquus_ and _Buccinum undatum_
(A. Hancock); south-eastern shores, off the Eddystone, Lighthouse (C.
S. Bate).
FEMALE. Pl. 22.
I may premise that after the sketch of the leading peculiarities of
Alcippe, and after the discussion on its affinities, just given under
the Family, I think it would be superfluous to institute a full generic
description, separately from the following detailed account of this
most anomalous cirripede.
_General Appearance._--The whole animal is from .2 to .3 of an inch in
length, of a soft texture, colourless or yellowish, and lives concealed
in a cavity of its own formation in the shells of certain Gasteropods.
This cavity communicates with the water by a narrow fissure-like
orifice (Pl. 22, fig. 4), broadest at the posterior end, where the
cirri are exserted; narrow, closed, and generally curved at the other
(_a_) end: the two sides of the fissure (_b_) are commonly bordered
by a calcareous inorganic deposit: the walls of the cavity are worn
so thin over the peduncle, at the narrow end of the fissure, that the
orange-coloured ovaria can generally be seen through the shell of the
mollusc, and hence there is here a distinct fan-shaped stain (fig. 3)
on the surface. The animal consists of a compressed capitulum, without
valves, and of a sort of peduncle depressed on its rostral face, and
covered with a broad, oval, thin, horny disc. We must remember that
in the Lepadidæ the peduncle does not essentially differ from the
capitulum, being only the flexible lower or anterior end of the animal,
and is separated from the capitulum only by shape, and generally by the
direction of the lines of growth. The disc, when most regular (fig.
1, H), lies in a plane at right angles to the sides of the capitulum,
and almost in a line with the orifice leading into the sack; but the
peduncle is often very irregular (fig. 2), and the disc comes even to
occupy a position nearly parallel to one or the other side of the
capitulum. On the carinal side, the capitulum is generally separated
from the peduncle by a rather deep fold (_f_, in the section fig. 5),
but this depends in some degree upon the state of distension of the
mass of ovarian cæca. I have given a drawing, fig. 1 (partly taken
from Mr. Hancock), of a very regular individual, and of an extremely
distorted specimen (fig. 2). The distortion, I believe, is generally
caused by the animal, during its excavation, breaking into some old
cavity.
_External Structure._--The orifice leading into the sack is about
one third of the total length of the animal: its edges or lips are
thickened, horny, and brownish: at the lower end, exactly where the
orifice ends, the lips are formed, from being deeply notched, into
two sharp projections (_a_, figs. 1, 5, 6), unlike anything occurring
in any other Cirripede. The external membrane (_c_, fig. 6) of the
lip supports an irregular but nearly straight band of sharp, thick
spines of chitine, about 1/1000th of an inch in length, together with
a few hairs: at the carinal or upper end of the orifice the spines are
largest and most numerous; at the other and lower end, they decrease in
size; and on the two projections (_a_, fig. 6), and on the adjoining
parts of the external membrane, they graduate into the small dentated
points which cover the whole surface of the animal. The inner tunic of
the sack (_b_), on each side along the upper half of the orifice, is
remarkable from having a moderately broad, curved band of short, sharp
spines, not quite so thick as those on the external surface, closely
adpressed together and pointing upwards, like the javelins of an
ancient phalanx, thus probably preventing the ingress of any intruding
animal. This band of spines curves at the upper end, conformably with
the shape of the orifice. The inner tunic of the sack in this upper
part is yellowish, and, what is very unusual, is thicker than the
external membrane. A little way down, within the orifice, and more
especially in front of an elegant row of hairs on the two sides of the
great labrum, there is a band of very fine but stiff hairs (5/1000ths
of an inch in length), pointing upwards, and making together with those
on the labrum a hedge, barring ingress into the sack.
The external membrane over the whole animal, excepting the horny
disc which covers the rostral face of the peduncle, is very thin and
transparent; it is periodically and often moulted, as may be inferred
from the many old lines of junction round the edges of the horny disc:
it is irregularly and pretty thickly (but not so thickly as in fig.
7) studded with star-headed, minute points, from 2 to 5/10,000ths
of an inch in diameter, composed of hard chitine, seated on a short
footstall, and this on a circular, yellowish, slightly thickened disc
of the general investing membrane, appearing like a halo surrounding
each little point. These points are directed obliquely upwards. There
are none on the horny disc, though particularly numerous close to its
margin. Their state varies much: just after a moult, when newly formed,
the spines are regularly star-headed, with quite sharp rays, from two
to six in number, with some of them occasionally bifid; but these
points or rays soon become blunted, and ultimately half the star is
worn away, so that the appearance then presented is that of a crescent
with a few blunt points on its convex side. At each exuviation, the
thickened membrane of the orifice with its strong external spines (the
condition of which also varies according to the period elapsed since
the last moult), and of course the whole internal tunic of the sack,
with its spines and hairs, are all moulted, together with the external
membrane and the little star-shaped points. In most specimens a barely
distinguishable band or bar of yellowish, slightly thickened membrane,
runs from the points (_a_), at the lower end of the orifice, for some
way obliquely downwards; and at the lower end of this bar the weak
adductor scutorum muscle (having transverse striæ) is attached. This
bar is often strengthened by a prominent external fold of membrane, but
yet it is so flexible, and as it is united only to the lower end of the
orifice, I can hardly believe that it can, by means of the adductor
muscle attached to its opposite extremity, have much power in closing
the orifice. I believe that this muscle acts simply in narrowing the
whole animal, so as to favour its movement within the cavity in which
it is imbedded. Owing to this position of the adductor muscle, and its
consequent little power in closing the orifice, we can understand the
necessity for the defence afforded by the bands of spines and hairs on
the inner tunic of the sack and on the labrum, which do not occur in
other Cirripedes.
_Horny disc._--The general shape of the disc, its irregularity and
position, have been already described. It never extends, as remarked
by Mr. Hancock, to the extreme lower point of the peduncle; upwards it
reaches to a little below the lower end of the orifice. It consists of
successive layers of membrane, either moderately thick and opaque, or
only a little thicker than the general membrane of the body, but never
furnished with the little sharp points; it increases in size, in like
manner as the calcareous valves of other Cirripedes, the undermost and
last formed layer extending beyond the others, with its edge united,
till the next exuviation, to the general membrane of the body. The disc
is attached, at its upper end, apparently in the usual way, by cement,
to the roof of the cavity of the shell in which it is imbedded; but
the lower parts of the disc are also slightly and partially attached,
chiefly along the lines of growth or exuviation; and this, I suspect,
is effected by an inorganic calcareous deposit; anyhow I could not
perceive here any cement or cement-ducts. Beyond the circumference of
the disc the whole animal lies free in its cavity. The lines of growth
in the middle part of the disc are generally obliterated by the decay
of the older and outer layers. These lines, though of course ordinarily
conformable with the general outline of the disc, are not always so,
for the disc sometimes becomes during growth slightly changed in form,
and the animal, consequently, slightly changed in position; sometimes
either one or the other side or the upper end of the disc is left
deserted by the new layers of the growing disc; these being formed on
the deserted side of less size or extension, instead of larger size, as
they normally should be all round the disc.
The upper end of the disc is always produced into a projection of not
regular shape, but generally hollowed out or embayed in front (fig.
1), and almost always hollowed out on the two sides. This projection
stands directly over the adductor muscle (_b_ in fig. 5), and on the
exterior surface is generally convex, being concave on the under
side for the attachment of several muscles presently to be described.
The horny layers are in this part usually thicker than elsewhere. The
disc is thus upwardly produced, owing apparently to the fissure which
leads into the cavity of the shell of the mollusc becoming, during the
process of excavation, considerably longer than is necessary,--that
is longer than the orifice leading into the sack; and consequently,
for the protection of the imbedded animal, the lower and narrow end
of the fissure is closed on its under side by this upward production
of the horny disc, formed of layers of membrane of unusual thickness.
A deposition, also, of lime, hereafter to be mentioned, gives further
protection.
In the bays on each side of the upward production of the horny disc,
and likewise a little lower down on its edges, and therefore somewhat
protected by lying within the narrow, pointed, lower end of the fissure
in the shell of the mollusc, the short-lived Males (Pl. 22, fig. 1,
_m_) are attached often in groups of two, three, or more together.
It may be asked to what part, in other Cirripedes, does the horny
disc answer? Not considering the upward prolongation, which has been
developed for a special purpose, the disc is irregularly circular,--is
added to all round,--serves for the attachment of the whole animal
to the supporting surface,--is covered on the under surface by a
conformable and parallel mass of ovarian cæca, and the latter by
the inner tunic of the sack; therefore in every character, and in
its relation to the other parts of the animal, the disc answers to
the end of the peduncle, or to the basal cup in Lithotrya, or still
more closely to the basis in sessile cirripedes, with the important
exception that it lies in a line with the longitudinal axis of the
whole animal instead of at right angles to this axis. We know that all
ordinary cirripedes become first permanently attached in their pupal
state by their antennæ, which are seated on the ventral or rostral
surface, near to the anterior end of the body; and that from the young
cirripede, after the act of metamorphosis, being turned vertically
upwards, and from the extreme anterior, now lower, end of the body not
being rapidly developed, the surface cemented down, or the basis,
encroaches almost equally on the dorsal, lateral, and ventral surfaces.
But if we were to suppose the extreme anterior point of the body to be
rapidly developed, the surface of attachment or basis, without it grew
still more rapidly, could not possibly reach the dorsal surface, and
would, consequently, be confined to the ventral or rostral surface. I
have not seen the young of the ordinary or female Alcippe soon after
its metamorphosis, but in the male the development of the extreme
anterior end of the body is extraordinarily rapid, and from analogy we
may fairly conclude that this is likewise the case with the female.
Hence I believe that the horny disc answers to the cemented down, lower
end of the peduncle, in other members of the Lepadidæ, and to the basis
in the Balanidæ, and that it is confined to the ventral or rostral
surface, owing to the anterior or lower end of the body having been
rapidly developed. To make all the parts, internal and external, of
Alcippe, correspond with those of other cirripedes, the main circular
part of the horny disc must be turned up at nearly right angles to its
present position (the dorsal or carinal integuments, to the right-hand
in fig. 5, being shortened), and then we should have a peduncle,
certainly very short and broad, but holding its proper relative
position.[143]
[143] In the genus Lithotrya, as long as the animal continues to
bore into the rock, the calcareous discs by which it is attached in
its cavity, stand, as in Alcippe (Pl. 8, fig. 2, 2_a′_, Darwin's
'Monograph on the Lepadidæ'), parallel to the longitudinal axis,
but as soon as the animal ceases to bore, and the discs become
converted into a cup, they occupy a normal position at right angles
to the peduncle. According to Reinhardt, these discs, in Lithotrya,
are situated on the carinal or dorsal surface of the peduncle, at
which statement I now feel considerable surprise, as undoubtedly
the pupa must first permanently attach itself by its prehensile
antennæ on its ventral or rostral surface. In Anelasma I failed to
discover any cement or cement-ducts; but I am now strongly inclined
to believe, considering that the extreme lower or anterior end goes
on growing, that the surface of attachment will be found to occur,
as in Alcippe, on the rostral surface, a little way below the
orifice.
_Sack and its Muscles._--I have already described the curious phalanx
of spines, the long fine hairs, and thickened condition of the inner
tunic of the sack along the sides of the orifice. This inner tunic
is a reflexion from that enveloping the body of the animal, in the
usual manner, as may be seen in the section (Pl. 22, fig. 5). Between
the external membrane and the inner tunic of the sack (_e_), there
is of course the usual double fold of corium, these two folds being
united by minute, transverse, ligamentous fibres, branched at the two
ends, as in other Lepadidæ. Imbedded in the corium there are numerous,
longitudinal, striæ-less muscles, which do not run quite up to the
orifice, but to an oblique line beneath it. Externally to these muscles
there are, as in the other Lepadidæ, fine transverse muscles, confined
to the middle part of the animal, and running from the carinal margin
more than half way round both sides. Attached to the upper notched or
folded end of the orifice (_g_, fig. 5, above the upper ends of the
longitudinal muscles), there is a fan of rather strong, striæ-less
muscles, expanding downwards, with their lower extremities attached to
the outer membrane of the capitulum; these muscles apparently serve to
open the orifice: there is a somewhat analogous muscle in Lithotrya,
but in no other member of the Family: in Cryptophialus, however, there
is a closely similar muscle. Owing to the action of these several
muscles, the tissues forming the capitulum and peduncle are, according
to Mr. Hancock, highly contractile.
I have stated that the under surface of the upper produced end of
the horny disc is concave, and serves for the attachment of several
muscles. Of these some run to the basal margin of the great labrum, and
no doubt, as usual, move the whole mouth; others, as usual, run to the
skin between the labrum and the lower end of the orifice,--_i. e._ in
fig. 5, between the lower end of the row of fine hairs (see fig. 11),
which shows where the basal margin of the labrum is situated, and the
lower side of the point (_a_), where the orifice terminates: others
run obliquely on both sides towards the point of attachment of the
small adductor scutorum muscle (_b_, fig. 5): others, of considerable
strength, and these are more peculiar, run and are attached to the
lower end of the orifice (_a_), and serve apparently to draw up the
orifice from within the fissure-like cavity, in which it lies lodged:
others, again, extend transversely on both sides, close beneath the
inner tunic of the sack, a little beyond the line whence the ovigerous
fræna or branchiæ arise. These transverse muscles lie within the
longitudinal muscles, and therefore are quite different from the
exterior transverse muscles, which are situated more towards the
carinal portion of the peduncle and capitulum, and which are common to
most Lepadidæ. The internal transverse muscles, and those running to
the lower end of the orifice, are peculiar, but we shall hereafter meet
with them even more developed in Cryptophialus.
Along the medial carinal line there is, between the two layers of
corium, the usual circulatory channel. On each side of this line, on
the inside of the sack, there are generally some slight irregular
swellings, and sometimes a large extent of the inner surface is
irregularly carunculated with little knobs. The sack (_e_ in fig. 5)
extends down almost to the basal point of the peduncle (_d_), more
especially when the ovarian cæca are not gorged with ova.
_Branchiæ, or ovigerous Fræna._--Within the sack, on each side of the
body, rising not far from the ends of the adductor muscle (_b_), there
is a large fillet or fold; the two occupy so exactly the position
of the ovigerous fræna that I cannot doubt such is their nature,
though, as happens in the case of some species of Pollicipes, they
are destitute of their proper glands, and so do not serve for the
attachment of the ovigerous lamellæ; this attachment probably is not
required, owing to the protected situation which the lamellæ hold in
the sack, under the animal's body, and over the ovarian cæca. From
the unusually large size of these so-called fræna, I cannot doubt
that they serve as branchiæ, equally well with the plicated folds of
membrane, believed to be homologous with the fræna, in the Balanidæ,
which have by every one been considered as branchiæ. The fræna are
broad and truncated at their upper ends; their margins are sinuous, and
their outer surfaces papillose; they run longitudinally down the sack,
narrowing as they extend, almost to the basal point of the peduncle,
and hence are of considerable length; they are hidden in the section
(fig. 5) by the medial, somewhat protuberant mass (_c_) of ovarian
cæca, and partly by the (_i_) prosoma.
_Body._--The body is constructed on the usual type, and indeed does
not differ greatly from that of Ibla. The labrum is very large, its
lower or basal margin is separated by an unusual space (capable of
being contracted or folded) from the lower end of the orifice of the
sack; hence the labrum and whole mouth is placed quite remarkably
near the upper (or carinal) end of the orifice. This upper end of the
orifice, I may remind the reader, is homologically the posterior end
of the general covering or carapace, and all that portion of the whole
animal (as the sectional figure, 5, stands) below the lower margin of
the labrum, on the rostral or ventral surface, is formed by the three
anterior segments of the head. The main part of the body, carrying the
mouth, is formed by the great development of that segment of the thorax
which bears the first pair of cirri (_h_), here closely adpressed, as
usual, to the sides of the mouth. The lower portion of this segment
forms the prosoma (_i_), and has the characteristic outline, but is not
much developed. On each side of the prosoma an oval space of membrane
is yellowish and is thickened, and so gives support to this part of
the body. The five succeeding thoracic segments, which ought to carry
the five succeeding and posterior pairs of cirri, are together of very
small size (as in Ibla), in comparison with either the prosoma, or
the whole anterior part of the animal. The segment (_k_) which should
have borne the second pair of cirri, is considerably longer than the
following segments, and is at the same time less distinct, owing to an
oval convex shield of thickened membrane on the sides, not extending
the whole length of the segment, thus causing two transverse creases,
which, when the thorax is contracted, appear like two additional
segments. Had this segment borne cirri, they would have stood, as in
Ibla, at a considerable distance from the first pair. The segments
(_l_, _m_) which should have borne the third and fourth pairs of cirri
are like each other, except that the former is rather the longest.
The membrane covering all the thorax is surprisingly thin; and at the
articulations, which are straight and transverse, is deeply folded, so
that the thorax must be highly extensible, to a degree which I have
not seen equalled in any Cirripede except in the males of _Scalpellum
vulgare_ and _ornatum_. The thorax is represented as somewhat extended
in fig. 5. This part of the thorax is amply furnished with striated and
striæ-less muscles for its retraction and protrusion, and for lateral
movements. The segment (_m_) which should have borne the fourth pair
of cirri, at first sight falsely appears like the terminal segment
of the thorax: in one monstrous specimen it bore a single cirrus,
showing (if there had been any doubt) that it was a true segment.
The three terminal pairs of articulated appendages, form together a
brush; they consist of the fifth and sixth pairs of cirri and of the
caudal appendages: my reasons for considering the last-named organs
as of caudal origin will be given hereafter. A moderately careful
inspection, especially of the ventral surface, will show that the fifth
pair of cirri are borne on a small segment (_n_, fig. 5, but plainer
in fig. 13), which is quite distinct from, but partially concealed
by, that which ought to bear (and did bear in the monstrous case) the
fourth pair of cirri: this segment is oblique, and cannot be traced
distinctly all round the dorsal surface. The segment (_o_, fig. 13)
bearing the sixth pair is much less distinct, and can only be seen by
a longitudinal section, or when the cirri are a little separated, but
it certainly exists,[144] as is likewise shown by the presence of small
apodemes dipping in amongst the muscles, between this and the last
segment. The posterior or caudal appendages are closely approximated;
they are not separated by any fold from the sixth thoracic segment;
but _appear_ as if they were articulated on the dorsal surface of the
sixth pair of cirri, in exactly the manner usual in the other Lepadidæ.
The segments bearing the fifth and sixth pairs of cirri are highly
oblique to the preceding segments, and consequently the cirri, which
they support, instead of projecting inwards, lie like a brush in a
line with the longitudinal axis of the main part of the thorax. The
membrane forming the two small oblique terminal segments of the thorax
is strengthened by irregularly shaped plates of thicker and yellowish
membrane.
[144] In the middle, the fold is slightly prominent and pointed,
and being most finely villose, I for some time looked at this
projection as a rudiment of the probosciformed penis.
_Mouth._--The mouth is constructed on the strictly normal type of the
Family, but is peculiar in every part; it is remarkable from being
situated so near the upper (or posterior) end of the capitulum, this
being caused by the great length of the labrum, and of the space of
body between the latter and the lower end of the orifice. The _labrum_
is a very singular part of the mouth from its vast size and outline:
in fig. 8, we have a front view of the mouth, of which the whole upper
pointed part consists of the labrum, and _h h_ is the first pair of
cirri; in fig. 11, we have a lateral view of the labrum, with the
surrounding thin membrane of the body, _a_--_a_, still adhering to its
edges; _h_ is the first cirrus on the near side; _m_ the mandible, a
little distorted in order to show its tooth, marking the position of
the transverse crest of the labrum and of the orifice of the œsophagus;
_b b_ is the medial longitudinal ridge of the labrum. In the Balaninæ
the labrum forms a mere rim to the back of the mouth, consisting of
an inner fold running down the œsophagus, and of an outer fold, both
close together: in the Lepadidæ the folds are separated, the outer one
being swollen or bullate; and here this structure is carried to even a
greater extreme than in Ibla and its allies. The distance between the
transverse crest over the œsophagus and the blunt projecting point on
the summit of the medial ridge, _b b_, equals twice the longitudinal
diameter of the rest of the mouth. The lower margin on each side of
the labrum is produced into two projections (fig. 11), the longer
one curling round to a point beneath the jaws, with its extremity
imbedded as an apodeme. Another very peculiar character in the labrum,
prominently noticed by Mr. Hancock, is caused by a longitudinal row,
on each side, of closely approximate, long, very finely pointed hairs,
which, as already stated, are fronted on the opposed internal surface
of the sack by an irregular band of still finer hairs. The surface
of the labrum is partially covered by minute toothed scales, and
these, seen on the longitudinal medial ridge, _b b_, give it a finely
denticulated structure. At each end of the transverse crest which
overhangs the œsophagus, there is a knob, such as occurs on the labrum
of every Cirripede. United to these two knobs, which are formed of
thick and yellowish membrane, and springing from the adjoining sides of
the mandibles, there are two swellings formed of thin membrane (fig.
8), which occupy the exact position of the palpi, and may be considered
as these organs in a rudimentary condition and destitute of bristles.
The _mandibles_ are simpler than in any other Cirripede; they are
minute; they consist of an oblong plate, with only one very strong
tooth at the upper end: the face towards the labrum is swollen: beneath
the upper free part there is a small, sub-triangular piece of thickened
membrane, let in and forming part of the general outer surface of
the mouth, and representing the large square plate found in other
Cirripedes. The _maxillæ_ (fig. 15) are smaller but broader than the
mandibles; they have an upper tooth and a smaller lower one, lying not
quite in the same plane with the upper one, but nearer the mandibles.
The apodeme (fig. 8, 15) is of remarkable length, extending beneath
the basal fold of the mouth: it does not arise from the ridge or outer
edge of the maxilla, but a little on one side, from the face directed
towards the mandible. Between maxillæ and mandibles there is a very
singular prominent fold of membrane (fig. 8, 15), which resembles,
but probably falsely, the supposed rudimentary palpus attached to the
mandible. Altogether the maxillæ differ considerably from the same
part in other Cirripedes. In structure they seem adapted to assume
the function of mandibles; but they do not stand directly over the
œsophagus. The _outer maxillæ_ (fig. 8) appear like a minute, deeply
notched lower lip: each consists of a simple, oblong, rounded plate,
with a few small bristles at its upper end. The basal fold of the
mouth in front, beneath the outer maxillæ is distinct, and runs in a
line with the basal articulation of the first pair of cirri. In the
rudimentary palpi, minute and little developed outer maxillæ; and in
the inner maxillæ, taking the function of the mandibles, the mouth of
Alcippe presents some resemblance with that of Anelasma.
_Cirri._--These consist of the first, fifth, and sixth pairs: the other
pairs are absent, except in one monstrous specimen, in which there was
a fourth cirrus quite like the fifth. _First pair_, fig. 14, these are
seated on each side of the mouth in the usual position. They are formed
of very thin and flexible membrane. The pedicel, as usual, consists of
two segments, the upper one is short and not very distinct; but when
viewed on the inner side can be seen to have the ordinary structure:
both segments are destitute of bristles. There are two short rami,
being about one third of the length of the pedicel: they are directed
either in the line of the pedicel, or more commonly posteriorly, that
is towards the other cirri, and therefore in an unusual direction. The
anterior ramus is generally rather longer and thinner (as is commonly
the case with other Cirripedes) than the posterior ramus; but there is
some variation in this respect. On neither ramus is there any trace
of the ordinary articulations: both are thickly clothed with fine
bristles, which are singular from being thickened in their lower parts,
and plumose, like a feather. These cirri have some resemblance, as
remarked by Mr. Hancock, to a pair of pincers; but they cannot act as
such; they serve, I believe, as brushes. Delicate muscles, transversely
striated, enter and are attached within both rami and within both
segments of the pedicel, on the usual type, showing that these organs
(if there had been any doubt) are truly cirri.
The _fifth_ and _sixth pairs_ of cirri (fig. 13, _n′_, _o′_) are almost
exactly alike: they are of very small size: each cirrus consists of
four segments: the lower or basal segment is broad, with a few minute
bristles scattered on its inner surface: the second segment is also
broad, but shorter, with a few, generally hooked bristles, in two short
irregular rows, in the upper part: these two segments answer to the
two segments of the pedicel of ordinary cirri. The third segment is
thinner and longer than the second; it bears two or three longitudinal
rows of bristles, most of which are neatly hooked at the point; its
upper end is surrounded with a circle of bristles. The fourth and
terminal segment is short, thin, and simple, with only a few bristles
at the apex. These two upper segments are bent a little inwards; they
answer to one of the two normal rami of ordinary cirri. The third
segment does not stand exactly on the middle of the summit of the
second segment,--the posterior corner of the latter being occupied by
a very curious, convex, oblong, rather hard (especially in the lower
part), protuberant cushion (as called by Mr. Hancock) or button (fig.
9, _c′_), transversely wrinkled by fine, distinctly crenated ridges.
This button presents a considerably different appearance according to
the point of view, fig. 9, 10: on one of its sides it projects beyond
the outline of the second segment, whence it arises; on the other
side it is prolonged, as a smooth ridge, on the top of the second
segment, embracing to a certain extent the base of the third segment.
On the face opposite to that which has been drawn (fig. 9) as most
characteristic, it is seen to be somewhat constricted round its base;
this constriction, representing, I believe, an articulation. When
viewed directly in front (fig. 10) its outline is oval, passing into
shield-shaped. Its longitudinal axis is 3/1000ths of an inch in length;
but it varies a little in shape and size. I shall presently assign my
reasons for believing that these buttons are the posterior or inner
rami of the fifth and sixth pairs of cirri in a rudimentary and much
modified condition.
_Caudal Appendages._--These (fig. 13, _p_) are placed close together,
being articulated between the bases of the sixth pair of cirri, the
lines of junction being internally marked by minute apodemes. They
consist of four segments, resembling in every respect those forming the
cirri, with the important exception that there is not a vestige of the
button on the summit of the second segment; the segments are not so
thick as those of the cirri, and the terminal segment is smaller.
_Muscles and Functions of the Cirri._--For their size, the cirri and
caudal appendages have voluntary muscles of remarkable strength,
attached within their basal segments, and springing from the dorsal and
ventral surfaces of the so-called third and fourth (_l_, _m_) thoracic
segments. Other muscles, rising from within the basal segment of each
limb, run to the second segment, and from that to the third segment.
I could not distinctly make out whether any entered the terminal
segment. I have seen no other instance of muscles entering the caudal
appendages, but as in the pupa they are so furnished, we here have only
an embryonic character preserved. I may remark that the fifth and sixth
cirri, consisting of two large basal and two thinner terminal segments,
is likewise an embryonic character. From the position of the cirri, the
four hard protuberant buttons or cushions, tend to oppose each other
at a common point; and the caudal appendages fill up a gap behind,
between the cirri of the sixth pair. I at first thought, with Mr.
Hancock, that these buttons served to catch the prey; but, reflecting
on their convexity and hardness, they appear very badly adapted for
this purpose; it would, in fact, be a marvellous feat to secure, in
the dark, any moving object between four balls. On the other hand,
this very convexity, the hardness, and especially the crenated ridges,
and the powerful muscles (which from the first surprised me), are all
well explained, if we suppose the prey, being secured by the terminal
segments, to be triturated between these four balls: any part which
escaped upwards would, moreover, be retained in a sort of cage, formed
by the inwardly inflected terminal segments with their hooked spines.
This view of the very curious and unparalleled use made of a modified
portion, not of the haunch, but of an upper part of the two posterior
pairs of thoracic limbs, is in some degree confirmed by finding that
Cryptophialus, which has apparently analogous habits, requires its food
to be triturated, though in this case it is effected by very different
means, namely, by four beautifully toothed discs, with brushes of
hairs, developed within the lower end of the œsophagus.
The prey, when caught, would probably at once be carried by the
movement of the articulated thorax to the mouth (itself moveable),
and being there secured by the mouth in front, the caudal appendages
behind, the tips of the cirri above, and the broad pedicels of the
first pair on the two sides, it would be triturated by the four
crenated buttons, and would then be forced down the œsophagus by the
action of the simple jaws. I looked in vain in several specimens for
any object within the stomach. I believe, that when the specimens are
first taken, all half digested food is ejected by the mouth. Whether we
may thus account for the extremely foul condition of the rami of the
first cirri in all the many specimens examined by me, I know not; but
that these rami, which are thickly clothed with fine plumose hairs, and
are furnished with delicate muscles, act as brushes, so as to clean the
orifice of the sack, I can hardly doubt.
_Homologies._--I have as yet, to a certain extent, assumed that I have
correctly named the different parts; and a few remarks on this head may
be desirable, considering the absence of certain cirri, the singular
condition of the others, the close general resemblance of the cirri
and caudal appendages, and the fact of the latter being furnished with
muscles. The only cause for any doubt regarding the thoracic segments
is the shield of thick membrane on that segment (_k_, fig. 5), which
ought to have borne the second pair of cirri, causing two transverse
wrinkles (not distinguishable, however, on the ventral surface),
and sometimes making the segment appear as if it consisted of three
segments: if it did consist of three, as there can be no doubt about
the nature of the first pair of cirri, (not in a more rudimentary
condition than in Anelasma) or about the segment whence this first pair
arises, the two terminal oblique segments, with their appendages, would
be abdominal instead of thoracic: but this is improbable, inasmuch as
the abdomen is unusually little developed in the pupa (as presently to
be shown), and more especially from the circumstance of a monstrous
cirrus, identical in structure with the two succeeding pairs, having
been borne on a segment (_m_), which, in any case must be considered as
thoracic, for it is well known how very rarely thoracic and abdominal
limbs resemble each other. I cannot myself feel hardly any doubt on the
nature of these three pairs of appendages; for, in the first place,
the posterior appendages are articulated on and between the bases of
the adjoining pair, _exactly_ as the undoubted caudal appendages are
articulated in all other members of the family on the sixth pair of
cirri or terminal thoracic appendages. Secondly, we see in the male
of the allied genus Ibla, the very same appendages preserved as in
Alcippe, namely, the caudal, and the fifth and sixth pairs of cirri,
which latter, moreover, are generally uniramous. Thirdly and lastly,
in the likewise allied _Alepas cornuta_, we have the posterior rami of
these same fifth and sixth pairs of cirri in a rudimentary condition,
and resembling in every respect the caudal appendages. Assuming,
then, that the several appendages in Alcippe have been rightly
denominated, we have to consider the nature of their segments: in all
cirripedes, the pedicels of the cirri consist of two segments, of
which the lower one (as here) is longer than the upper one, and both
(as here) considerably thicker than the segments of the rami: in all
cirripedial pupæ, likewise, the thick pedicels of the limbs consist of
two segments, and each ramus, also (as here), of two segments: now,
with these coincidences, and bearing in mind that in Alcippe the two
upper segments do not arise from the exact middle of the summit of the
second segment, but from rather its anterior side,--bearing, also,
in mind the case just cited of _Alepas cornuta_ with the posterior
rami of these very same cirri rudimentary,--we may, I think, safely
conclude that here in Alcippe the two lower segments form the pedicel;
the two upper segments, the anterior ramus; and that the button-like
protuberance is the posterior ramus in a modified condition. As the
caudal appendages in none of the Lepadidæ, either in the mature state
or in the pupa, have two rami, we can satisfactorily understand the
absence of any trace of the button-like protuberance on the top of the
second segment.[145]
[145] I almost wish I could persuade myself that I had taken
an erroneous view of the thoracic segments, and therefore that
the three pairs of terminal appendages were all abdominal, for
then Alcippe would come into much closer relationship with
Cryptophialus; though even in that case it would form a distinct
family from it: but I cannot alter my opinion.
_Alimentary Canal._--The œsophagus runs down from the mouth, beneath
and nearly parallel to the straight row of hairs on the two sides of
the labrum: it is surrounded by the usual muscles: at the lower end it
bends down, and expanding a little, like a bell, enters the stomach.
The stomach is of considerable size and fills the main part of the
body, bulging out under the mouth, and prolonged as far as about the
middle of that segment (_l_), which ought to have borne the third pair
of cirri; here the stomach terminates in a blunt rounded point. The
tissue surrounding the stomach, and keeping it in its proper place, can
be traced to the posterior end of the thorax, but there is no rectum or
anus. I am prepared to assert positively that this is the case,[146]
for I made repeated longitudinal sections of the whole thorax in two
planes, and I subsequently cleaned the outer tissues with boiling
potash, and then, when as transparent as a sheet of glass, I examined
every part, and certainly there is no rectum (which in every case
is formed of chitine, and so is not acted on by potash) nor an anal
orifice. Singular as this fact is, it is not so improbable as it at
first appears; inasmuch as I have shown, in my former volume, that the
Lepadidæ can reject half digested food by their mouth, and secondly,
that the final stage of digestion appears to take place in the upper
part of the stomach. In the male of this very species, as we shall
immediately see, there certainly is no mouth or stomach, and apparently
no rectum or anus; so is it likewise with the males of _Scalpellum
vulgare_ and _ornatum_: in Proteolepas, there is a mouth and an
œsophagus, but no stomach, rectum, or anus. There are, I believe, no
other known instances, in the whole great class of Crustacea, of the
absence of an anus.
[146] I may venture to remark that I succeeded in every attempt,
which I made, in seeing plainly the œsophagus, and the acoustic
and olfactory orifices and sacks, which, according to all analogy,
would be of much smaller size, and far more difficult to discover,
than the rectum and anus. I may mention that, according to Mr.
Newport ('Annals of Nat. Hist.,' 1849, p. 277), the larvæ of
certain parasitic Hymenoptera have a stomach without any anus.
No crustacean, according to Milne Edwards, is destitute of this
orifice.
The stomach, in Alcippe, is much corrugated, so as to be deeply
pitted; but there are no regular cæca. The enveloping hepatic layer
is thick, brownish, pulpy, and formed of pellets of cellular matter,
not distinctly arranged in lines as is general; there is the usual
delicate muscular layer. The stomach was in every case empty, and I did
not notice the separated epithelial coat, so generally found in other
cirripedes.
_Organs of Sense._--I failed in discovering the eye, which I have no
doubt exits, as it is conspicuous in the pupa and in the male. The
olfactory pouches are seated rather laterally under the maxillæ, as in
Ibla. As in this same genus, the acoustic sack is seated remarkably
low down (fig. 5), at a very considerable distance beneath the basal
articulation of the first cirrus: the orifice is seated on a slight
prominence: the acoustic vesicle, I believe, is sub-cylindrical, with
irregular projections. I did not make out anything distinctly on the
nervous system.
_Female Generative System._--The animal we have thus far described
is exclusively female: when a longitudinal section of the thorax is
made, and the stomach removed, it can be most plainly seen that there
are no vesiculæ seminales or testes. Mr. Hancock has remarked on the
absence of the usual probosciformed penis. The male of Alcippe will
be subsequently described in detail. The female organs differ in no
respect from those of other members of the family, excepting in so
far that the layer or mass formed by the ovarian cæca (_c_) does not
lie transversely to the longitudinal axis of the whole animal, but
longitudinally under the horny disc. The ovigerous fræna are largely
developed, but serve, as previously stated, as branchiæ, and not for
their proper function of giving attachment to the ovigerous lamellæ.
The ovigerous lamella is single, and nearly corresponds, in size and
shape (as would ensue from the manner of its formation) to the under
side of the horny disc. The ova are broadly oval, and rather above
1/100th of an inch in length.
_Metamorphoses._--The larva in the first stage has been fully described
and figured by Mr. Hancock: it differs in no essential respect from
other larvæ of the family. Mr. Hancock overlooked the inferior minute
antennæ. With respect to the larvæ in the last stage, or pupa, I
obtained several specimens attached to the disc of the female, and
which were on the point of being developed into males; and another
specimen identical in all respects, but attached independently to the
shell of the mollusc, and which, therefore, I have every reason to
suppose, would have been developed into a female. In any case these
pupæ may be conveniently here described (Pl. 23, fig. 16.) They are
.025 of an inch in length; they are of the usual shape, with the
anterior end not very blunt and the postero-ventral surface somewhat
produced. The whole carapace or shell is very thin and smooth. There
are six pairs of thoracic natatory legs, situated far back towards the
posterior end of the body; each leg has the usual articulations, and
the two rami their usual long but not plumose spines; the presence of
the legs deserves notice, considering the rudimentary and modified
state of their homologues in the mature animal. The abdomen differs
considerably from the same part, as far as I have seen, in other
pupæ; it consists of only a single almost globular (fig. 17, _q_)
segment, instead of three segments; and the two caudal appendages
(_r_) are very long, and are composed each of only a single segment
(instead of two), carrying at its tip two short spines. There are two
purple eyes, 4/3000ths of an inch in diameter, which, after having
been dried and then soaked, could be seen to be compound; they are
fixed in the usual manner to two rather short apodemes, which latter
have their usual origin. But the pupa has a very unusual appearance
owing to the presence of a single dark purple eye, half the diameter
of the two larger eyes, situated behind and above the latter, and
quite disconnected with the apodemes; this is the eye of the mature
animal, which, for some reason, is here developed earlier than usual.
The prehensile antennæ are remarkable from being seated very close to
the anterior extremity: owing to this, the articulation of the second
or main segment with the basal segment, is hardly at all oblique.
The whole pupa is of exactly the same length as the pupa of _Ibla
quadrivalvis_, and so are the antennæ, (see p. 286 of my volume on
the Lepadidæ), viz., 32/6000th of an inch, but the second segment is
narrower, (being only 8/6000ths in breadth in the broadest part), and
is longer in proportion, for the disc which forms part of the total
length is only 4/6000ths in length, whereas in Ibla it was 8/6000ths;
the disc is here hoof-shaped, as in Ibla. The ultimate segment is
remarkably short and narrow, (being only 3/20000ths in width, and less
than half the size of that in Ibla); it carries (I believe) three
terminal spines, and is not notched. Altogether the antennæ more nearly
resemble those of Ibla than of any other genus in the family. From the
position of the antennæ, and from the length of the second segment, the
pupa, when cemented by the disc or third segment, to the supporting
surface, adheres, with its posterior end almost vertically upwards.
With respect to the young cirripede within the pupa, I could only
observe that its anterior end was formed into a blunt point.
_Powers of Excavation; Inorganic Deposit of Calcareous Matter;
Attachment._--Alcippe, according to Mr. Hancock, attacks only dead
shells of the Fusus and Buccinum, and always on their inner sides,
especially on the columella. The excavations, in the specimen which I
examined, were so numerous as almost to touch, and sometimes to run
into each other, the included animal being thus rendered distorted. The
orifices are directed with respect to the shell indifferently upwards
or downwards. From the shape and size of the cavity corresponding
to that of the included animal, there can be no doubt, as stated by
Mr. Hancock, that Alcippe forms its own cavity. That the action is
mechanical I think may safely be inferred from the whole outer membrane
being studded with minute, star-headed points of hard chitine, which
rise from halo-like little discs of thickened membrane, which latter
are well adapted to allow the underlying adherent muscular layer to
act on the points, and thus on the surrounding shell. Consequently
the points generally show signs of severe attrition, but they are
periodically and often replaced, at each exuviation, by new and much
sharper points. There are no points on the permanently attached layers
of the horny disc, but it particularly deserves attention, that the
renewable membrane always extends beyond the circumference of the
disc, and is there most thickly studded with the points. We have
met, in Lithotrya, with a precisely analogous fact in the extension
of the periodically moulted membrane of the peduncle, furnished with
star-headed points of chitine, and in addition with minute calcareous
beads (which, however, seem soon worn away), beyond the calcareous
discs, by which this cirripede is attached in its cavity. We need not
feel much surprise at points of chitine being hard enough to wear away
shell, when we consider what work the jaws of insects, likewise formed
of chitine, will effect.
With respect to the first commencement of the excavation, the pupa,
owing to the position of its prehensile antennæ, fixes itself with its
posterior end almost vertically upwards; and the young cirripede, after
its metamorphosis, from the greater length of the ventral integuments
formed round the eye-apodemes, must be thrown backwards into nearly
the position represented in Pl. 22, fig. 12, _b_. I have not seen a
young female at this early age, but I have traced the development of
several males, and have found that the lower end of the peduncle,
(_i. e._ what was the anterior end of the pupa), grows at quite a
remarkable rate, so as very soon to form a great bag extending beyond
the attached prehensile antennæ. Now if we suppose an analogous
structure in the female or ordinary Alcippe, and the supposition is
quite allowable, we shall almost immediately have the anterior or lower
end of the young cirripede, just in advance of its antennæ, pressing
against the surface of the shell of the mollusc; and if armed with
triturating points, as we have every reason to believe it is, it would
wear for itself a cavity. The horny disc on the ventral surface of
this protuberant anterior end of the young animal will, we may assume,
soon become cemented to the near side of the cavity just supposed to
have been excavated. And the whole animal, by further slight changes
in direction, namely, by working down more and more obliquely, will
take, as shown at (_c_), its final position. As the whole surface of
the animal, with the exception of the horny disc, is provided with
triturating points, the animal, when once imbedded, can and does
increase its cavity at both ends in length, in depth, and all round
the edges of the horny disc,--in short, in every direction excepting
directly over the horny disc. I believe, as already explained, that the
young Alcippe, (_b_, diagram), first bores obliquely into the shell;
and whatever amount of downward extension the horny disc attains before
the young cirripede assumes its proper position, with its ventral
surface upwards and parallel to the inner surface of the shell of the
mollusc, that amount determines the thickness of the plate of shell
hereafter to be left unabraded over the horny disc, as the latter
continues to extend in circumference. This plate of shell over the
horny disc is so thin, that, as mentioned at the commencement, the
colour of the ovaria is seen through; and until I reflected on the
following considerations, I was much surprised how the instinct of
the animal could so neatly guide it not to grind too deeply, and yet
to grind till only a very thin plate of shell was left over its horny
disc: these considerations are, that whatever thickness was first
given to this plate of shell, when the animal was very young and first
assumed its ultimate position, that thickness would in most cases be
always retained, owing to the flatness of the disc, and to the membrane
armed with triturating points protruding very slightly beyond and
above the horny disc, only just enough to wear away the surrounding
shell to the thickness necessary to allow of the formation of each new
zone of disc; as the disc itself is not armed, it subsequently has
no power of wearing away the plate of shell above it. Thus the horny
disc, besides giving support and attachment to the peduncle, is of this
peculiar service that it seems to guide, (somewhat like the wood-part
in a plane), the rasping powers of the lower extreme margin of the
peduncle.
I may here observe that certain radiating and often punctured lines,
mentioned and figured by Mr. Hancock, which help to render the thin
plate of shell over the peduncle conspicuous (fig. 3), are formed by
the burrows of an excessively minute annelid, the punctures being
apparently the exit orifices: I imagine that these annelids find it
difficult to commence their burrows on the smooth surface of the shell,
and that they congregate at these particular spots and thence burrow in
radiating lines, owing to their having taken advantage of the little
cliff-like edges, at the narrow and disused ends of the fissures
leading into the cavities occupied by the Alcippe, where alone they
would not be disturbed by the action of the cirri, when first they
commenced making their little burrows in the shell.
The fissure leading into the cavity is required to be broad at the
posterior end, in order that the cirri may be there freely exserted
out of the sack; and narrow in other parts, to prevent, as it would
appear, anything injurious getting in between the animal's body and
the cavity in the shell of the mollusc. As the fissure is increased in
length by attrition at the broad posterior end, which end during growth
becomes broader and broader, the lower part of the fissure has to be
narrowed, and this is effected in a very singular manner, namely, by
advantage being taken of the strong tendency, which triturated shell
with animal matter, has to set into a solid shelly mass, although
constantly agitated.[147] Mr. Hancock noticed this edging of hard
shelly matter, and naturally thought it was a secretion. Lines of
deposition (Pl. 22, fig. 4, _b_), parallel to the edges of the furrow
can often be perceived in it: its thickness and extension vary much: I
have seen it on one side alone of the orifice: it is, of course, never
found at the broad end where the process of enlargement goes on. The
peculiar worn surface with which it irregularly thins away downwards,
on the sides of the cavity, made me (together with the apparent
impossibility of such a secretion proceeding from an animal wholly
invested by a chitine membrane) suspect it to be inorganic; and this
view is certainly correct, for when a fragment is dissolved in acid,
a considerable residuum is left of bits of membrane, rubbish, and, in
one instance, even of the remnants of a foreign animal, apparently an
annelid. We have here all the circumstances favorable for inorganic
deposits of this nature, namely, finely triturated shell and chitine or
animal matter, produced by the excavation of the chamber, sea-water,
and movement.
[147] I have given some remarkable cases in my volume on 'Volcanic
Islands,' (p. 49), in which limestone, having almost the hardness
and specific gravity of marble, has been thus deposited. Almost
every coral-reef offers similar examples. The curious substance
described by Mr. Horner and Sir David Brewster, ('Philosoph.
Transact.,' 1836, p. 65), which is formed during the manufactory of
cloth, offers another example of the strong tendency which lime and
animal matter have to unite. Lately, Dr. Horsford, in 'Silliman's
North American Journal,' Jan. 1853, has discussed the chemical
theory in an analogous case on the coast of Florida; he attributes
the aggregation to the formation of a hydrate of lime through
the action of the animal matter. Mr. G. B. Sowerby, Junr., has
described a case very analogous to that of Alcippe, ('Proceedings
of Zoolog. Soc., Mollusca,' Pl. 5, fig. 4, p. 162, 1850), namely,
that of _Pholas calva_, in which a tube is formed of _inorganic_
calcareous matter, serving to narrow the entrance.
From the manner of growth of the animal, the fissure leading into the
cavity in the shell becomes much longer than the orifice leading into
the sack, and to prevent the body being unnecessarily exposed, the
upward projection of the disc, already described, is formed under the
narrow and disused end of the fissure; moreover, the two rims of the
inorganic calcareous deposit sometimes here approach so closely, as
almost or actually to touch each other; and between them, as remarked
by Mr. Hancock, there is usually a little accumulation of grains of
sand. This narrow end of the fissure is generally curled either to the
right or left hand; and I can only account for this fact by supposing
that, whilst the cirripede is young, and has not a large horny disc
attached to the cavity, it cannot keep its body straight during the
long-continued boring process.
The animal is attached by its horny disc to the thin shelly roof over
the peduncle, and likewise to the under side of the narrow end of the
fissure, but is elsewhere quite free. I carefully examined the disc
in many specimens, but could not see any cement-ducts: I believe I
saw layers of cement at the upper end of the disc, but it is not easy
to discriminate between this substance and the yellowish, somewhat
disintegrated, layers of the horny disc. The pupa certainly becomes
attached by ordinary cement, so that the attachment in early life,
at least, is normal. In some full-grown specimens, I found the lower
parts of the horny disc attached, along the edges of the layers, to the
roof of shell; and as I looked here in vain with the highest powers
for cement-ducts, or for cement, it appears to me probable that the
rough edges of these layers were united to the roof by a thin layer of
the inorganic calcareous deposit. The animal, from its very protected
situation, certainly requires to be less firmly cemented than other
cirripedes; and even in Lithotrya, which is less deeply imbedded
than Alcippe, the cementing apparatus was feebly developed. From the
length of the pupal antennæ, cemented by their terminal segments, the
position of the young cirripede (Pl. 22, fig. 12) can be changed to
a considerable extent, like a ship swinging at her moorings, but in
order to assume its final position, the animal must, I think, travel
like Lithotrya, but to a much less extent, by a short succession of
overlapping horny discs,--the old discs being partially deserted, each
new one extending beyond the last-formed one: even in the case of the
mature animal, we have seen that, under certain circumstances, it
changes, to a certain extent, its position; portions of the old disc
being deserted and attached to the roof of a deserted portion of the
cavity.
_Affinities._--In the preliminary remarks under the Family, I have
discussed this subject almost sufficiently: I will here only remark,
that the genus, though so abnormal, yet stands naturally between
Ibla and Anelasma, having clear affinities, on the one side, through
and beyond Anelasma to Alepas; and on the other side, beyond Ibla to
Scalpellum, and so to Lithotrya. Moreover, it is very distinctly
related to Cryptophialus in the succeeding Order.
MALE. Pl. 23.
On every specimen of the female Alcippe, which I carefully examined,
I found some minute parasites (or epizoons) attached to the lateral
edges of the upper part of the horny disc, and therefore lying within
the narrow end of the fissure leading into the chamber excavated in the
shell of the Buccinum. Although having had some experience in the very
anomalous forms which male cirripedes assume, yet when I first casually
inspected these parasites under a weak lens, from their transparency,
their elongated and lobed body, including an internal folded up organ,
I actually threw them away, thinking that they were probably Bryozoa.
Subsequently, a more careful inspection immediately showed the cemented
prehensile antennæ, and their cirripedial nature was demonstrated. I
soon found specimens with the perfect still adherent exuviæ of the
locomotive pupa, undistinguishable from the pupa already described as
probably belonging to the female Alcippe. But as this latter fact,
may perhaps be doubted, I must show that there is other evidence
sufficient to prove that these cirripedial parasites are the males of
the female Alcippe. Of the females, I inspected many specimens, and all
certainly were without external male organs; and in the four or five
specimens which I rigidly examined, there were no testes or vesiculæ
seminales, the latter being in all hermaphrodite and male cirripedes so
conspicuous. On the other hand, I examined at least thirty specimens
of the parasite, and they were all exclusively males, for all had
a probosciformed penis, and the greater number had their vesiculæ
seminales filled with spermatozoa, and hence were ready to perform the
act of impregnation, but undoubtedly they contained no ova. It would,
then, be very strange, if these two cirripedes of opposite sexes, thus
attached together, were not sexually related. Wonderfully different as
the parasite is from the female Alcippe, yet, in one very important
character it is related to Alcippe, and to no other member of the
Family, namely, in the sack extending down to the extreme lower point
of the peduncle; the male organs, I may add, occupying an analogous
position with the peculiar position of the female organs in Alcippe.
The lateral lobes of the peduncle in the parasite seem to represent
the sides of the broad depressed peduncle in Alcippe; and in both the
peduncle grows at its lower end--a very rare circumstance--observed
only in two genera in this Family, namely, in Anelasma, and in a
slight degree in Lithotrya. Besides these points of resemblance
between Alcippe and its parasite, which are striking, considering
their external utter dissemblance, the affinities of both point,
judging from certain small characters, in the same direction, namely,
towards Ibla and Alepas. Finally, then, I think, we may confidently
admit that this parasite or epizoon is the male of the female Alcippe:
indeed, considering the facts given in my former volume, on Ibla and
Scalpellum, I have, perhaps, here discussed the question at unnecessary
length.
The males are generally attached, as already stated, to the two
hollowed out sides of the upward prolongation of the horny disc; they
adhere by means of little patches of cement, proceeding from the
terminal segments of their antennæ, to the overlapping edges of the few
later-formed zones of the disc; hence, they lie protected, within the
narrow end and a little under the edges of the fissure leading into the
cavity in which the female is lodged. In some specimens, however, the
males are attached rather lower down on the disc, and are not confined
exclusively to its upper margin, so that they live fairly under the
roof of shell which covers the main part of the disc: but they are
never attached very low down, so as to lie far from the lower end of
the orifice leading into the sack of the female. I have two or three
times seen as many as three males on each side, but sometimes there is
only one on each side, or none on one side. A large distorted specimen
actually had twelve males, and two pupæ on the point of undergoing
their final metamorphosis, all fourteen attached on one side, and all
evidently must have been alive together! Another specimen had nearly
the same number, a few on one side, and the rest on the other side.
The male immediately, after the exuviation of the pupal carapace,
25/1000th of an inch in length, is only 23/1000th of an inch long,
but ultimately it becomes, chiefly from the growth of the lower end
of the peduncle, nearly twice this length; for the largest specimen
which I have seen, that figured, was 45/1000th of an inch long (_i.
e._ under 1/20th of an inch), and 1/100th of an inch in breadth across
the peduncle, beneath the lateral lobes. The whole external membrane
of the animal (as well as the internal membrane of the sack), is very
thin, quite structureless, and as transparent as glass; hence, even
the spermatozoa, within the vesicula seminalis, can be seen from the
outside. The whole structure of the animal is very simple. The ventral
surface can be at once recognised by the attachment of the antennæ
(fig. 19, _a_), and these organs mark the point which was the anterior
end of the male, just at the period of its metamorphosis, and before
the lower end of the peduncle had grown. These antennæ have already
been fully described; they are conspicuous from being composed of
membrane, rather thicker than that investing the body of the male,
and which external membrane can be traced entering these organs, and
appearing like cement-ducts; but within these tubular prolongations of
the outer membrane, I could obscurely see the real cement-ducts.
The part answering to the capitulum is much flattened and elongated;
it widens but little from the upper to the lower end, where it blends
with the carinal or dorsal surface (the under surface in fig. 19)
of the lobed peduncle. At the upper end there is a small orifice,
and close to this, on the ventral or rostral side, there is a thin,
apparently double projection (_i_, fig. 19) or flap of membrane, one
flap lying exactly over the other. The whole length of this capitulum
probably corresponds with that small portion of the capitulum in the
female, between the upward prolongation of the horny disc and the
lower end of the orifice; and the two broad flattened projections in
the male, probably answer to the two sharp narrow points (_a_, fig. 1,
Pl. 22) in the female. The peduncle has two lateral lobes (_h_, _g_,
fig. 19), and, whilst young, what may be called a third and medial
lobe, but this soon increases largely by growth, and forms the main
part of the peduncle. The lateral lobes are intimately connected with
the ventral surface; they tend to lie in a plane, at right angles
to the compressed capitulum, but owing to the excessive thinness
and flexibility of the whole external membrane, it is difficult to
ascertain the relative position of the different parts. Moreover, owing
to the pupa being so much flattened, these lobes are necessarily formed
folded up; and, I believe, it depends on the position, with respect
to surrounding objects, which the male ultimately holds, whether the
lobes ever assume, their apparently normal position, in a plane at
right angles to the sides of the pupa; owing, also, to the form of the
pupa, the two lobes seem generally to be actually formed of unequal
sizes, that formed in the dorsal region of the pupa being the largest.
I believe that these lobes correspond with the lateral margins of
the upper end of the peduncle of the female, which margins project
laterally beyond the sides of the capitulum. The lower lobe, or end
of the peduncle, is depressed in the same plane with the lobes; it is
of variable length; when first formed it hardly extends beyond the
basal articulation of the prehensile antennæ. Commonly it does not lie
quite in a straight line of the capitulum; and I have seen specimens
in which it stood at nearly right angles to the capitulum and to what
was the ventral surface of the pupa; this irregularity in the relative
position and sizes of the different parts of the peduncle, no doubt,
to a considerable extent, depends on the form of surface to which the
male becomes attached, just in the same way as we have seen that the
peduncle of the female becomes altered in shape during the excavation
of the chamber in which it is lodged.
I feel some difficulty on one point: in the pupa the single eye of the
future male can be clearly distinguished, and it lies some way from
the anterior end of the body; but in two males, which certainly had
just moulted, and in which none of the internal organs were as yet
developed, the eye lay close to the anterior end, directly over the
basal articulation of the antennæ. I suspect this is somehow caused by
the great change of form which supervenes, during the metamorphosis,
at this anterior end of the body; the extremely compressed body of the
pupa having to become depressed and lobed in the young male. I have
given a figure of a young male, just as it appeared (Pl. 23, fig. 18),
somewhat distorted from lying on a flat surface; _c_, being the eye.
The sack extends, in a very remarkable manner, down to the lower end
of the peduncle, the whole inside of the animal being thus freely open
to the water. In the upper part, the sack forms a mere narrow tube; it
does not appear to have been formed in the same manner as in all other
cirripedes, namely, surrounding the thorax and natatory legs of the
pupa, but in an abnormal position, along the dorsal surface, above the
sack and thorax of the pupa: a transparent line, where the new narrow
sack is in process of formation, is the first indication of the coming
metamorphosis. The sack in the capitulum of the male is not central,
but lies near the dorsal surface; the ventral interspace, between the
outside and the sack, is occupied by oblique fibres (_l_, fig. 19),
which may be striæ-less muscles, but I suspect are ligamentous fibres,
giving support to the whole projecting capitulum. These fibres enter a
little way within the lobed peduncle; they are probably homologous with
the strong muscles, which run from beneath the upper end of the horny
disc of the female to the lower end of the orifice leading into the
sack. Round the lobed peduncle,[148] there are two bands (_e_, _f_) of
thin muscular fasciæ, slightly oblique to each other, and attached at
the ends to the outer membrane; they are evidently homologous with the
external transverse muscles, which are best developed round the same
part in the female. Some of these muscles present a singular chain-like
appearance, from being strangled at intervals: they act probably in
aiding the long probosciformed penis to protrude itself out of the
sack. I could not detect any longitudinal muscles, and the lower part
of the peduncle seems destitute of muscles of any kind.
[148] I believe I saw in one specimen, most delicate transverse
muscular fibres round the lower part of the elongated capitulum.
The internal structure of the animal is very simple. Within the lower
end of the peduncle there is a dark purple eye (_c_), under the
1/1000th of an inch in diameter, a testis (_d_) and a (_b_) vesicula
seminalis. These organs falsely appear as if suspended in the middle
of the peduncle, but they are really attached, I believe within a
separate partition, to the ventral surface, occupying the same position
as the mass of ovarian cæca in the female. The eye lies on the line of
junction between the testis and the vesicula seminalis, and on their
ventral side. The testis is rounded, and consists of a mass of cells,
on an average 1/5000th of an inch in diameter. The vesicula seminalis
varies extremely in condition, being either a mere rather broad vessel,
enlarged where it joins the testis, or a bag fully as large as the
testis itself, and distended with spermatozoa, all arranged parallel to
its longer axis. There was an evident relation between the size of the
vesicula seminalis and that of the testis, the number of the cells in
the latter decreasing as the mass of the spermatozoa increased: there
was also an evident relation between the age of the male and the state
of these organs; younger and more opaque individuals, having their
testes of large size; and older specimens, with the lower end of the
peduncle arrived at its full dimensions, having the vesicula distended.
Some few old specimens had evidently discharged their spermatozoa. By
dissection I more than once distinctly traced the vesicula seminalis
entering the broad lower end of the penis. The membrane, forming
the vesicula, is ringed, and I presume is, as in other cirripedes,
contractile, so as to expel the spermatozoa. The probosciformed penis
(_m_) is of extraordinary length: it is plainly ringed, or rather
articulated, in this respect resembling that organ in Ibla and Alepas;
it tapers gradually, and terminates (as usual) with a brush of fine
bristles; it is furnished with delicate voluntary muscles, arising from
the body round its basis, and extending no doubt up to the apex, but
too fine to be traced all the way. Its broad lower end is attached in
a slight depression, on the ventral side of the sack, a little above
the point of attachment of the pupal antennæ. According to all analogy,
the spot whence the penis springs must be considered as representing
the thorax and abdomen; and the outer membrane of the penis is here, as
on this view it should be, reflexed and is continuous with that lining
the sack. Ordinarily the penis lies coiled up in complicated folds,
appearing like a large intestinal worm, and fills the lobed part of
the peduncle, which apparently serves for no other purpose than its
reception. In one case in which I dissected out the penis, I found
it in its contracted state; 41/1000th of an inch in length, equal to
that of the entire capitulum and peduncle; in a specimen, in which the
penis had been naturally exserted, the part which protruded (_m_) was
by itself rather longer than the whole animal; and as this specimen
had been placed in spirits of wine, the organ no doubt was contracted;
hence I think it probable that the probosciformed penis, when fully
stretched out, would equal twice the length of the entire animal.
There must be a nervous system; and there must likewise be a gland
(homologous with the ovaria) for secreting the cement; but I could not
distinguish parts so small. Certainly there is no mouth, or stomach, or
thorax, or limbs of any kind, or abdomen.
It is obvious that these males must be very short-lived: they perform
their masculine functions and then perish. We have seen, however,
that after the act of metamorphosis they do grow a little, and I have
reason to suspect that this is effected, as with other Cirripedes,
by moulting. The growth must be absolutely dependent on the store of
nutriment laid up within the pupa. The young male, immediately after
the exuviation of the integuments, thorax, natatory legs, abdomen,
and eyes of the pupa, consists of a pulpy cellular mass, without any
internal organs as yet formed.
Judging from the different sizes of the females which included
perfectly developed ova, I infer that they must breed more than once
during their lives; and therefore, that successive sets of males, as
in the genus Scalpellum, must become attached to them. I was not,
however, able to discover the prehensile antennæ or other remains of
the old males adherent to the females; a circumstance which I presume
is accounted for by their attachment being weak. Considering the
very small size of the male, it is not surprising that so many,--in
one case fourteen,--are required to impregnate the numerous ova of a
single female. How the males know the proper period when the ova,
lying in a sheet at the very base of the sack of the female, are ready
for impregnation, I cannot say, without it be that they perceive
the moulting of the external membrane, close to the edge of which
they are attached; for this moulting would indicate the period when
the ovigerous lamella came to the surface of the sack, and the ova
would then be soon ready for impregnation. From the position in which
the males are attached, and from the extraordinary length of the
probosciformed penis, capable of voluntary movements, I have no doubt
the males can insert the tip of this organ within the lower edge of
the orifice of the sack, and there discharge the spermatozoa, which,
by their own movements, must pass down the sides of the sack of the
female till they reach their proper destination. The position of the
males, with respect to the female's body, is almost exactly the same
as that occupied by the complemental males of _Scalpellum Peronii_
and _villosum_; the lower and narrow end of the fissure, worn in the
gasteropod shell, here affording that protection to the males, which
the edges of the opposed scuta afford to the _complemental_ males
of the above two species of Scalpellum. We cannot doubt that these
latter males aid in the impregnation of the ova of the hermaphrodites,
but they are not furnished with a very long penis, probably for the
very reason that they are _complemental_ males, and therefore not so
absolutely necessary for the impregnation of the ova as are the males
of Alcippe.
I have, in my former volume, expressed my astonishment at the extent
to which abortion had been carried in the male Ibla; but it has been
carried much further in the male Alcippe. In Ibla, the thorax is
reduced to a mere flap, and only two pairs of cirri exist in a most
useless and rudimentary state, but there is a well organised mouth,
stomach, and anus. In the males of _Scalpellum vulgare_, _ornatum_,
and _rutilum_ there is no mouth or stomach, but there is a thorax with
four pairs of minute, modified cirri, and a large abdominal lobe. Here,
in the male Alcippe, all these negatives are united, we have no mouth,
no stomach, no thorax, no cirri, no abdomen! The archetype crustacean
consists of twenty-one segments; of these the seventeen anterior
segments can be clearly made out in the archetype Cirripede: now, in
the male Alcippe, the first three segments are largely developed,
forming all that is externally visible, but the remaining fourteen
segments are absolutely aborted, but in idea may be considered as
forming the membranous depression whence the probosciformed penis
springs; for this organ normally arises at the extremity of the
seventeenth segment. To show the wonderful diversity of nature, even
in the same sub-class, I may be permitted to remark, that whilst in
Alcippe only the three anterior segments are developed, the fourteen
succeeding segments being rudimentary, in Proteolepas (hereafter to be
described) these fourteen segments are all largely developed, whilst
the three anterior segments are quite aborted, being represented only
by a thin envelope to the two threads by which this Cirripede is
attached to the supporting object.[149]
[149] It may be worth stating, that in order to procure perfect
specimens of the female and male Alcippe, pieces of the shell
inhabited by them should be dissolved in weak acids.
ORDER II.--ABDOMINALIA.
_Cirripedia, having a flask-shaped carapace; body consisting of one
cephalic, seven thoracic, and three abdominal segments; the latter
bearing three pairs of cirri; the thoracic segments without limbs;
mouth with the labrum greatly produced, and capable of independent
movements; œsophagus armed with teeth at its lower end: larva, firstly
egg-like, without external limbs or an eye; lastly binocular, without
thoracic legs, but with abdominal appendages._
I feel compelled to form an Order for the one genus and species,
namely, _Cryptophialus minutus_, to be here described. We must, I
conceive, attribute much greater value, in classification, to internal
parts and organs, at least where such are not known to vary, than to
external structure. Now in Cryptophialus, the body consists of eight
segments, of which the first two are not developed in any cirripede
hitherto described. Of the eight, the seven posterior or thoracic
segments are quite free, or detached from the carapace, and do not
bear any appendages; whereas in all the foregoing cirripedes of the
order Thoracica there are (at least in the normal sex) six pairs of
cirri; Alcippe alone must be excepted, in which there are only two
pairs. Again, in the Thoracica there are no abdominal appendages,
excepting the terminal or caudal, whereas in Cryptophialus the abdomen
bears three pairs of biramous cirri. In the crest of the labrum,
being produced into a special, lancet-formed organ, articulated at its
base and capable of movement, and in the palpi projecting straight
upwards, we have a great difference from all other cirripedes;
and these organs, we have every reason to believe, possess a high
classificatory importance. The œsophagus in Cryptophialus, where it
enters the stomach, is armed with teeth and hairs, moved by muscles,
forming a beautiful structure, of which we have not a trace in any
other cirripede. Lastly, and perhaps most importantly of all, the
metamorphosis is different; for the early larval stages are passed
under an egg-like condition within the sack of the parent; and the pupa
differs from the pupæ of all other cirripedes, in not having natatory
thoracic limbs, and is therefore only able to crawl about by the aid of
its great prehensile antennæ.
Thus far the evidence is decisive in favour of Cryptophialus being
placed in a separate Order; but if we were to trust to the characters
derived from the external covering or carapace,--and such characters
are of high importance, as we may safely infer from the natural
arrangement of the foregoing families which depends on the structure
of the carapace,--we should place Cryptophialus close to Alcippe
amongst the Lepadidæ. These genera agree in their burrowing habits,--in
their attachment by a horny rostral disc,--in the external membrane
being covered with triturating points,--in the spinose and notched
orifice, with an external lateral bar on each side, and in the inner
tunic of the sack being protected by hairs and spines. They agree to
a considerable extent in shape, and in the peculiar arrangement of
the muscles of the whole external covering of the animal: they agree,
also, in their manner of growth, and in the sack extending down to
their basal extremity. Some of these resemblances may possibly be
analogical, and due to similarity of habits, and not to affinity; and
we must attribute to mere similarity in function, a certain amount of
resemblance in their labrums, for this part is essentially different
in the two genera; and to the same cause, the resemblance between the
brush formed by the two pairs of thoracic cirri and caudal appendages
at the end of part of the thorax in Alcippe, with the three pairs of
abdominal cirri at the end of the whole thorax in Cryptophialus. I
allude to this latter resemblance, for it was owing to it, and to the
similarity in the habits of Cryptophialus and Alcippe, that I stated,
in the introduction to my former volume on the Lepadidæ, that the two
genera would probably fall into the same order. In the structure of all
the parts of the mouth and of the cirri, in the digestive organs and in
the metamorphoses, Cryptophialus is not more closely related to Alcippe
than to any other genus whatever amongst the Lepadidæ.
Nevertheless I am confirmed in the view that the external resemblances
between these two genera are due to some real affinity, and are not
merely analogical, by a very remarkable fact,--namely, that Alcippe and
Cryptophialus are both bisexual, and have males, several in number,
attached exactly in the same position, and which males are so closely
similar that, considered by themselves, they might absolutely be almost
classed as species of the same genus! For they agree in the absence of
all internal organs and parts, excepting the single testis, vesicula
seminalis, and immensely long probosciformed penis; and they agree,
also in manner of growth, in the arrangement of the muscles, and even
in shape. The whole case seems to me very singular, and, as far as my
knowledge extends, unique: we have two animals, of which the females,
if classed by their external parts (homologically consisting of the
three anterior segments of the head), would be placed alongside each
other in the same family; but when classed by the whole rest of their
organisation, certainly must be ranked in distinct orders;[150] yet
the males of these very same animals might almost stand in the same
genus. If it be asked why the position of Cryptophialus in the system
should not be determined by the male, instead of by the female, the
answer would be that the male is here abnormal and rudimentary in
its whole structure; and I believe systematists are agreed that less
perfect parts (and therefore a less perfect whole) offer less valuable
characters than the more perfect parts or whole. We see this conclusion
plainly verified in the case of the hermaphrodite _Scalpellum vulgare_
and _Ibla quadrivalvis_, for there can be no doubt where these species
should be arranged, yet if we attempted to place them by their
complemental males, we should utterly fail: exactly in the same manner,
if _Ibla Cumingii_ and _Scalpellum ornatum_ were ranked by their males,
they would be quite misplaced. So again, if we were to attempt to class
the six species of Scalpellum by their males and complemental males,
undoubtedly the first three and last three species of the genus would
have to stand in distinct orders! Hence we may reject the males as a
foundation for classification, but no doubt they serve to show that the
resemblances in the carapaces of Alcippe and Cryptophialus, are not
merely analogical or functional, but evince a true affinity, though
these genera differ so greatly, in mouth, body, œsophagus, cirri, and
especially in their metamorphoses.
[150] M. Milne Edwards would, perhaps, in accordance with the
profound views lately propounded by him on classification, consider
Cryptophialus as an extremely modified, and, to a certain extent,
degraded member or satellite of the _type_ of the Lepadidæ; but
I do not myself feel able to draw a line of distinction between
the being a very abnormal member of one group, and belonging to
a distinct group. I may add that I have several times tried to
persuade myself, with no success, into the belief that I have
somehow misunderstood the homologies of the thoracic segments and
cirri of Alcippe and Cryptophialus; for if this were so, the two
genera could be brought into much closer relationship; but with
any conceivable amount of error on my part, there remains the
great difference in the metamorphosis, not to mention the palpable
differences in the cirri, the parts of the mouth, and in the whole
course of the alimentary canal.
CRYPTOPHIALUS MINUTUS. Pl. 23, 24, fig. 1 to 19.
_Hab._--Chonos Archipelago, Southern Chile; imbedded in the
_Concholepas Peruviana_.
FEMALE. Fig. 1-18.
_General Appearance._[151]--This, the smallest known cirripede, is
flask-shaped and compressed, with a small orifice, on a more or less
produced neck, placed at one corner: one of the narrow sides of this
carapace is somewhat flattened or depressed, with its superior edge
prolonged a little upwards; by this side, the animal is attached to
the cavity in the shell, within which it is lodged. There is here no
distinction between a peduncle and capitulum, that is between the
lower or anterior, and the upper or posterior end of the animal,
as seen externally. The small orifice is toothed and hairy: it is
coloured purple, as is likewise the projecting labrum; the rest of
the animal being tinted only by the muscles and internal parts seen
through the outer integuments. The largest specimen (fig. 2) which I
have measured did not quite attain the length of one tenth of an inch.
This cirripede inhabits, in vast numbers, the shells of the living
Concholepas Peruviana, amongst the Chonos islands; the whole outside
of the shell being sometimes completely drilled by its cavities,
almost touching each other, as happens in the case of Alcippe with the
shells of Buccinum. The oval aperture leading into the shell-cavity,
in full-sized specimens, is between (2-3)/100 of an inch in length: it
is generally surrounded by a narrow, internal, calcareous rim, which
apparently has the same inorganic origin, as in Alcippe. The toothed
orifice of the carapace leading into the sack, fills up the orifice of
the shell-cavity; but it can be voluntarily withdrawn a little: when
opened, and the animal is in action, the lancet-formed, moveable crest
of the labrum, and the abdominal cirri, are exserted.
[151] I am greatly indebted to Dr. Hooker, for having several years
ago, when I examined this my first cirripede, aided me in many
ways, and shown me how to dissect the more difficult parts, and for
having made for me several very correct drawings, which, with some
subsequent alterations, are now engraved.
_Integuments._--The external membrane is colourless, thin, but strong;
it is studded with minute bifid, trifid, and quadrifid points of hard
chitine, which are the agents of excavation: they are directed upwards,
except towards the lower end, where they are directed from the disc
or surface of attachment. These points beneath the orifice, and on
each side close along the lateral bar, are larger than elsewhere.
There are no points on the disc or surface of attachment, which is
formed of somewhat thickened, yellowish membrane, and is not moulted,
like the rest of the external membrane, but is formed of successive
layers extending beyond each other; the lines of growth, however,
being obscure, and only occasionally distinguishable. The disc is
oval, not extending to the lower end of the animal, and with the upper
edge thinning out and produced upwards (fig. 1). The animal, during
its growth, moves a little downwards in its cavity, by means of the
new layers of the attached disc being formed, not symmetrically with
respect to the old layers, but beyond them or at a greater depth in
the shell; hence when the animal is removed quite perfect, by the
Concholepas being dissolved in acid, the upper and deserted margin
of the disc or surface of attachment generally projects as a free
edge, but in a tattered and worn condition. In full grown specimens,
which have ceased burrowing downwards, nearly the whole disc, though
occupying the same position relatively to the animal, becomes in fact
deserted, and is lined by membrane continuous with, and like that,
investing the rest of the body, but furnished only with simple blunt
points, instead of with the sharp bifid and trifid triturating points.
The orifice is formed on each side by a toothed rim of hard chitine
(fig. 3, 4), which can be opened and shut, owing to its being flexible
at the rostral end, and folded inwards at the carinal or posterior
end. The teeth vary in arrangement and sharpness: generally they form,
taking a rim on one side (fig. 3), two prominences or groups of points
at the rostral or anterior end, always separated by a broad notch,
from the bottom of which the lateral bar extends downwards, from the
posterior, larger and less regularly toothed half of the rim. Of these
teeth the larger ones project nearly straight up, and the smaller and
lower teeth outwards, graduating into the smaller teeth, just below
the rim, which again graduate into the minute points, studded over the
whole surface. These outer teeth probably serve to prevent any creature
crawling into the cavity, between the shell and animal. Scattered
bristles rise from all over the rim. The lateral bar, above alluded
to, consists of the general membrane of the body, thickened, hardened,
rendered elastic, coloured yellow, and apparently formed into a fold:
where attached to the under side of the rim, at the above-mentioned
notch, the bar is thinner and more flexible than elsewhere: it runs
half way down the animal, first straight and then curved towards
and closely approaching the disc or surface of attachment. At the
lower end, the bar, or more strictly the thickened membranous margin
of the bar (_b′_, fig. 3), expands into an oblong, slightly rigid
plate, studded with from three or four to ten or twelve points, which
have their ends expanded and truncated, or even slightly bifid. The
extremity of this plate projects freely from the general surface of the
body. We shall afterwards refer to the use of this extremely peculiar
bar and plate.
The two rims forming the orifice cannot be quite closed; but ingress
by any foreign object into the sack is beautifully prevented by an
internal membranous lip on each side (_d′_, fig. 3, 4), and by a
third inwardly folded lip (_d′_) at the posterior and broad end of
the orifice. These three lips can be brought together, and form a
valve. The lateral lips are very narrow at the mouth or rostral end
of the orifice, where the hairy lancet-shaped crest of the labrum
closes the orifice, and largely expand towards the posterior end: they
are produced from the inner tunic of the sack: they appear formed of
the finest hairs, placed parallel and approximate, but when examined
under the highest powers, these hairs (for they still appear such) are
found to be united by delicate membrane, which has its extreme edge
fimbriated. The third, or posterior and inwardly folded lip, differs
in being composed of much coarser, flattened hairs, which are united
towards their bases, and are free at their extremities, where they are
serrated or coarsely plumose on both sides.
The sack extends down to the lower end of the animal. It is lined by
delicate membrane. At the orifice on each side, a little posteriorly
to the lateral external bars, this inner membrane is strengthened by a
pair of thin yellowish bars (_c′_, fig. 3), which run parallel to the
straight portions of the external bars. These inner bars at their lower
ends become pointed and die out: at their upper ends, and close to the
rim, they are broader, but more flexible, and so transparent as hardly
to be distinguished from the rest of the membrane. At the rostral end
of the orifice, in a medial line, this same inner tunic of the sack
is thickened for a short space downwards, so as to form a fifth bar
(_c′′_); which separating from the inner tunic, runs inwards between
the outer and inner membranes of the carapace (_i. e._ between _b_ and
_c_, fig. 3, 5), behind the mouth, as far down as opposite to the lower
end of the œsophagus, and there becoming thinner and ligamentous,
gives attachment to some powerful muscles.
At each exuviation, the external membrane with the dentated hardened
orifice, the lateral bars, the inner tunic of the sack with its bars,
are all moulted, together with the usual integuments of the animal's
body. New and sharp triturating points are thus periodically formed for
the work of excavation. The whole animal increases during growth in
every direction, and therefore, at its lower or basal end, as was the
case with Alcippe. The disc or surface of attachment, is added to by
new underlying layers, extending beyond the old layers at the lower end
and on the sides, but not at the upper end, where, as in the case of
the calcareous discs of Lithotrya, the old layers are deserted and worn
away. I saw what I believed to be little globules or patches of cement;
but I was not able to discover any cement-ducts.
_Muscles of Sack and Orifice._--The animal is surrounded by rather
strong longitudinal muscles, not running up close to the orifice: these
muscles exhibited, to my surprise, distinct traces of transverse striæ:
there are no external transverse muscles, as in all the Lepadidæ.
Attached to both sides of the inward fold or hinge, at the posterior
end of the orifice, some striated or voluntary muscles run for a short
distance downwards, diverging like a fan: their contraction would cause
the dentated rim to open: a strictly homologous muscle occurs only in
Alcippe. At the opposite end of the orifice, a remarkably powerful
voluntary muscle is attached to the ligamentous bar above described as
proceeding from the rostro-medial end of the toothed rim (_c′′_); and
at its lower expanded end, it is attached under rather above the middle
of the disc: this muscle corresponds with a similar one in Alcippe,
and with some much weaker muscles in other Lepadidæ. Its action would
be to draw down within the shell-cavity the whole dentated rim, and
likewise to close the orifice; and here, I believe, come into use
the lateral elastic horny bars with their curious basal projecting
plates, furnished with expanded points, for much friction would thus
be caused by, yet some play be allowed for, the several movements;
the elasticity of the bar bringing up the dentated orifice, when the
powerful muscles attached to the rostral end of the latter became
relaxed. Round the space where the just-mentioned muscles are attached
to the horny disc, a sheet of other muscles radiate, a few on both
sides obliquely upwards, but the greater number transversely and within
the first-described longitudinal muscles; they extend on both sides
about half round the animal. There are similar muscles in Alcippe, but
not extending so far round the animal. Their action must be to draw the
whole carapace towards the surface of attachment; the action of the
longitudinal muscles being to shorten it; the orifice supported by the
lateral horny bars, serving as the fulcrum for the contraction of the
longitudinal muscles. I could not see any adductor scutorum muscle,
although I looked particularly under the expanded plates at the ends of
the lateral external horny bars.
_Body._--This is laterally compressed: it is widest and thickest at the
upper end, and thence tapers to the lower or posterior end. The last
three or four thoracic segments are bent under the anterior segments,
giving the whole something of the appearance of certain crustaceans,
divested of their legs. The somewhat conical mouth, with its singular
labrum, is very large. The body consists of eight segments. The first
segment (fig. 5, 1), or that succeeding the mouth, is the seventh or
last cephalic segment of the archetype crustacean; it is the largest of
all eight segments; it is joined by its dorsal surface to the carapace
or external covering of the animal, and the membrane with which it is
invested is prolonged upwards and downwards (_c_, _c_, fig. 5), and so
forms the inner tunic of the sack. The succeeding seven segments are
thoracic; they are free, and are destitute of limbs; the articulations
separating them are transverse. The first and second thoracic segments
give rise, on their medial dorsal surfaces, each to a remarkable
tapering curved appendage, presently to be described. At the end of the
last thoracic segment, there is a minute abdomen, bearing three pairs
of biramous cirri.
The _Mouth_ consists of three pairs of organs, namely, the outer
maxillæ, maxillæ, mandibles with their palpi, and of a great and very
curious labrum. These organs, by the fusion (as in other cirripedes)
of their lower segments, form a large, somewhat conical, projecting
mouth, which is separated on the ventral surface from the rest of the
body by a distinct fold or articulation, where the muscles proceeding
to the above gnathites are attached. The labrum (_e_, fig. 5, 9) is of
large size; and the crest close over the opening of the œsophagus is
produced into a great, lancet-shaped, moveable organ, wholly unlike
anything occurring in any other cirripede: it is coloured purple, and
is thickly fringed in the upper part by very fine hairs: it is bowed a
little backwards from the mouth: the base, which rather overhangs the
œsophagus, is a little contracted, and is transversely marked by an
articulation: two small, parallel, voluntary muscles (with transverse
striæ) are attached at their lower ends close beneath the articulation,
and extend about one third up the organ: their contraction would serve
to erect it; and their relaxation would, apparently, allow it to fall
backwards on a little knob (_e′_, fig. 5) behind. This little knob
resembles a similar projection in many of the Lepadidæ. As the labrum
is formed of similar membrane with that of the succeeding segment of
the body, its limit downward, beneath the knob, can be told only by a
small apodeme which projects inwards, at a little distance within the
line where the membrane of the body is reflexed upwards (_c_), so as to
form the inner tunic of the sack.
The mandibles, palpi, and maxillæ, all project more than is usual. The
_Palpi_ (_f_, fig. 9, 5) are narrow, flattened, and taper a little;
they support a few long bristles on their tips, and on one of their
sides. In every other cirripede (in which the palpi are developed) they
are directed transversely across the mouth, one towards the other, and
are for a considerable space united to the labrum: here they project
straight up, and seem to rise exteriorly to the bases of the mandibles;
they are, however, united to the basal lateral edges of the labrum, and
when the latter is torn from the rest of the mouth, the palpi separate
with it. I could not distinguish the knob on which, in every other
cirripede, the palpi are articulated. The _Mandibles_ (fig. 8) have an
upper, lower, and middle tooth, with some finer intermediate points
and hairs. The _Maxillæ_ (fig. 7) are narrowed in at their spinose
edge, where there are three large spines and several finer bristles,
together forming a flattened brush: this organ is remarkable from the
apodeme (_a_) being bent into the shape of a scythe, with the terminal
or blade-portion a little expanded, and directed backwards and inwards.
The _Outer maxillæ_ are sub-triangular in outline, with several
bristles on their summits and along their outer surfaces.
_Segments of Body._--I have stated that the mouth is succeeded by
eight segments. As in all the cirripedes hitherto described, the
body consists of only six segments, the number eight at first seems
very improbable, and therefore I may be permitted to state that both
Dr. Hooker and myself, when first examining this animal, and having
no notion whatever regarding its homologies or the structure of
other cirripedes, came to the conclusion, judging only from external
appearances, that is, from the transverse folds, and from the lines of
movement when the body was bent by a needle, that there were really
eight segments. I have since carefully looked to this point: when the
outer membrane is cleaned and examined, the four posterior segments are
very plain, owing to a dorsal medial line, being alternately either
thickened and coloured yellowish, or thin and colourless: the four
anterior segments are less plain, but yet the membrane on the dorsal
surface, on the line of each fold or articulation, does present some
difference, from being destitute of the fine, transverse, toothed
scales which occur on other parts. But I lay most stress on the fact,
that all these eight articulations were used for the attachment of
muscles. Hence I conclude that the eight segments are real; and we
shall see, in the next order, that the very same eight segments are as
plain in Proteolepas, as in the larva of an insect or as in an annelid.
There is good reason to believe that the general covering or carapace
consists in all cirripedes of the three anterior segments, and that the
mouth (judging from its appendages) also consists of three segments,
consequently the first segment of the body in Cryptophialus must be the
seventh or last cephalic segment, and the seven next free segments must
be the normal seven thoracic segments.
The first segment of the body (_i. e._ last cephalic, fig. 5, 1) is,
as stated, the largest, and is attached dorsally to the carapace:
its ventral surface is flattened, and is formed of somewhat thickened
membrane: on each side, a little below the articulation separating this
segment from the mouth, there is a small blunt projection, with the
free part only 1/500th of an inch in length. Each of these appendages
bears four or five bristles on one side near the summit, and a few
on the other side, lower down: from their position I believe them
to be rudiments of a first pair of maxillipeds (tetartognathites of
Milne Edwards), of which no trace occurs in any other cirripede. The
differences between this segment and the seven succeeding segments,
is of interest, as offering some confirmation of the belief, lately
disputed by some naturalists, that the cephalic and thoracic segments
in the class Crustacea, do differ in their nature,--a conclusion which
we shall see further confirmed under Proteolepas. The second and third
(_i. e._ first and second thoracic) segments (2, 3, fig. 5) are the
next largest, and are remarkable from supporting singular appendages,
already alluded to. The sides of the second segment are formed of
thickened yellowish membrane. The fifth and sixth segments are the
smallest, and mark the point of chief flexure of the body. The eighth
segment is a little elongated, formed of thicker membrane than the
other segments, and dorsally is indented by the anus.
The singular tapering appendages (fig. 5, _i_, _k_) arising dorsally
from the second and third segments, differ from each other only by the
upper one being smaller, less curled, and perhaps rather smoother. When
first examining this animal, not knowing that it was female, and not
finding a probosciformed penis, I concluded that these organs were of
this nature,--an excusable mistake, considering their almost ringed
structure, their somewhat constricted bases, the direction of their
curvature, and their position in the midst of the ova within the sack.
On careful examination, however, these appendages are seen not to be
truly ringed or articulated, but are covered with transverse lines of
scales, hirsute on their edges; these scales being less distinct, or
even quite absent on the smooth upper portion; they do not include
any muscles; they are imperforate at the apex, which is not furnished
with bristles (as seems always to be the case with the probosciformed
penis); and, lastly, they are lined by corium, but are not occupied
by any vessel, gland, or organ of any kind. The only function which
I can assign to these appendages, is that of aiding the retention
of the ova within the sack: for the ova, when first produced, are
aggregated round them; at this period I several times observed long,
somewhat curled, very thin fibres, not tapering like hairs, adhering
to both appendages, the nature of which fibres I cannot explain. In
very many cirripedes there seems a strong tendency to the production of
tapering, filamentary appendages, somewhat like the two (_i_, _k_) here
described,--namely, at the bottom of the sack in some Balaninæ, at the
bases of the anterior cirri in Lepas and in some other genera, and on
the dorsal surface of the prosoma in certain species of Pollicipes; in
this latter case some of these appendages were covered by scales; and
the prosoma whence they arose answers to the third segment of the body
in Cryptophialus, or that supporting the lower and larger appendage.
Appendages of this nature, in several cirripedes, serve for the
lodgment of the testes, but in some cases they are of no apparent use,
excepting, perhaps, in aiding respiration by the expansion of corium
thus exposed, and this partially may be their function here, for there
are no proper branchiæ.
_Cirri._--There are three pairs, together forming a nearly straight
brush, of considerable length, projecting in a line with the last
thoracic segment. Each cirrus is biramous. In a moderately large
specimen there were twenty segments in one of the longest rami. Each
segment (fig. 14, _a_) is strengthened by a dorsal or posterior shield
of thickened yellowish membrane, from the upper edge of which a single
smooth spine projects; the anterior surface is likewise strengthened
on the two edges by thickened membrane, and supports two pairs of long
spines, which are plumose, or rather hirsute, on both sides. In the
lower segments of both rami of the several cirri, the inner spine of
each pair is considerably shorter than the outer spine,--evidently in
relation to the little power of divergence of the two rami. All the
cirri resemble each other, excepting that the rami of the anterior
pair, are rather shorter than those of the other cirri, but the dorsal
spines on their segments are longer. In all ordinary cirripedes the two
rami are equally supported by the upper segment of the pedicel, which
latter is very short compared with the lower segment, and is separated
from it by a transverse articulation. Here (fig. 13) the exterior or
anterior ramus is articulated on the pedicel, almost like a branch, in
front of the other ramus, which seems more directly continuous with
the pedicel. The upper segment of the latter is separated from the
lower segment, both being of about the same size, by a very oblique
articulation. On the front surfaces of the pedicels there are two or
three pairs of spines.
_Abdomen._--The three cirri on one side are separated from their
opposite pairs by a prominent, longitudinal fold, formed of thickened
yellowish membrane, which, when foreshortened by being viewed dorsally,
looks like a style projecting immediately beneath the anus. The
transverse folds separating the three pairs, are exceedingly slight.
The inner basal edges of the pedicels of the cirri project slightly
inwards as apodemes, giving, I believe, attachment to some muscles. The
anus lies between the posterior pair of cirri, and deeply indents the
last thoracic segment. The transverse folds separating the three pairs
of cirri, little developed as they are, must, according to all analogy,
be considered as representing three segments of the body, and as we
have accounted for seven cephalic and seven thoracic segments, we must
conclude that these are three abdominal segments. We know, moreover,
that the abdomen in the pupæ of the Thoracica, with a single exception,
does actually consist of three segments.
_Movements of the Mouth, Thorax, and Cirri, &c._--Judging from the
fact of the lancet-shaped appendage of the labrum being more or less
exserted in dead specimens out of the shell-cavity, and from the
analogy of other cirripedes, I do not doubt that the whole mouth can be
considerably raised and depressed; we have seen, also, that the rostral
end of the toothed orifice of the sack can be pulled down, which would
aid in exposing the mouth and labrum. The well-articulated thorax,
by the aid of the muscles attached to every segment, can certainly
be doubled up and contracted, so that the cirri with their pedicels
(coloured purple like the exposed labrum) can be wholly protruded out
of the sack and shell-cavity. The three cirri no doubt can be separated
a little from each other, both transversely and longitudinally; and
according to analogy, the two rami of each cirrus can likewise be
separated: there are, also, muscles for moving the two segments of the
pedicel of each cirrus; and other muscles run up the many segments of
the rami. We have seen that the great lancet-formed appendage of the
labrum, laterally fringed with fine hairs, can be erected; and I do
not doubt that the prey when entangled by the expanded cirri, is borne
against this appendage, and is then, by the retraction of the thorax,
dragged down its smooth surface to the mouth, where it is seized by
the mandibles and maxillæ, which lie like a trap at the bottom of an
inclined and moveable plane.
_Alimentary Canal._--The œsophagus is long; it runs backwards from
the mouth and then downwards; at its lower end, where it enters
the stomach, the part, which in other cirripedes is expanded and
bell-shaped, is modified in a most singular and quite peculiar manner;
for the lower part of the œsophagus, after widening a little, becomes
converted into what appears at first like a square box, 8/1000ths
of an inch across. This box is really deeply folded (see diagram,
Pl. 24, fig. 11) into six longitudinal ridges and hollows: two of
these hollows, facing each other, are wider than the others, and when
the organ is dissected out of the body, it generally lies (fig. 10)
with one of these faces exactly over the other, the narrower lateral
folds being thus hidden, and the whole consequently appearing like
a simple square box with concave sides. But when a section is made,
or the lower open end is turned upwards, we see that the organ is
six-rayed and elongated, with the longer axis standing parallel to
the flattened sides of the animal's body. The edges of the folds are
formed of yellowish, thickened membrane, with a sinuous or beaded
outline, which serve to strengthen the organ. Internally, the two
broad concave sides are armed, in their upper inwardly prominent (fig.
12) part, each with a disc, 2/1000ths of an inch in diameter, crowded
with short, thick, brownish, inwardly projecting teeth. The two discs
stand exactly opposed to each other. The bases of the teeth, seen from
the outside (Pl. 24, fig. 10), seem like brown little circles, with
a smaller circle within. The disc obscurely appears to be formed by
the confluence of two smaller discs which lie, I believe, at a very
small angle to each other: beneath each of these half discs there is
a longitudinal band of very fine hairs, the two bands uniting into
one, lower down within the organ. The internal longitudinal edges,
also, of the four lateral smaller folds are likewise clothed with fine
hairs; hence we have six parallel longitudinal rows of very fine but
stiff hairs, or eight, if the united bands on the two broader faces
under the discs be counted each as two. These bands of hairs, and the
opposed discs, armed with very strong teeth, can be separated and
brought together with force, by the action of strong constrictor and
diverging muscles. Hence any prey carried down the œsophagus, before
entering the stomach, would have to pass, as it were through a mill,
and be subjected to a severe trituration by the discs of teeth, and
immediately beneath to a brushing by the six longitudinal bands of
hairs. This curious and unique structure answers, I believe, the same
purpose as the four convex, hardish, crenated buttons on the posterior
thoracic cirri in Alcippe, which are likewise unique in that genus. I
observed that in some specimens the teeth had been worn quite blunt,
but the teeth and hairs are periodically moulted and renewed, together
with the whole œsophagus.
The stomach is broadest at the upper end, and extends from a little
beneath the mouth down to the fifth segment of the body, where it
becomes narrow. It presented an irregularly contracted appearance, and
was covered by a pulpy hepatic layer. The rectum is of rather large
diameter; it extends from the middle of the fifth segment to the end
of the eighth segment of the body, or seventh of the thorax, where
the large anus is situated, lying between the posterior abdominal
cirri, and partly hollowed out in this seventh segment. The rectum,
as in all other cirripedes, is periodically moulted. The food is of
a bright green colour, as if of a confervoid nature; the triturating
and brushing action of the œsophagus seems to roll this matter into
pellets, which apparently retain this form until finally expelled as
excrement: certainly the excrement is in pellets, and I have several
times seen pellets within the stomach.
_Organs of Generation._--The specimens as yet described are exclusively
female, there being certainly no testes or vesiculæ seminales. As in
every specimen collected (early in January) there were within the sack
either nearly mature eggs, or young larvæ, it was the worst period for
seeing the ovarian cæca, and I have failed to discover them in the
specimens now long kept in spirits of wine; but I cannot doubt they
would be found, between the inner and outer tunics of the carapace or
general covering, near the disc. I have stated in my original notes,
made when the specimens were alive, that the ova are at first perfectly
detached; but some appearances make me believe that I overlooked (as
might easily happen) the often _excessively_ fine membrane which in
other cirripedes unites the ova together, and so forms the ovigerous
lamellæ. The ova are much less numerous than in other genera, varying
from only nineteen to about sixty. In the same individual all the eggs
were always in the same state of development.
_Metamorphoses._--The true ova, in their earliest condition, when in
the sack, are ovate (Pl. 24, fig. 15), orange-coloured, quite smooth,
and barely 10/1000th of an inch in their longer axes. They soon become
broader at one end than at the other; and by degrees the narrow
posterior pointed end becomes developed into a slightly club-shaped,
almost transparent (fig. 16) horn, and the broader anterior end, into
two rather longer horns. The length of the oval part, not measuring the
horns, is nearly the same as in the primary true egg condition. There
is as yet no trace of internal organs, the whole contents consisting
of pulpy granular matter. How far the above changes are effected by
moulting, either of the whole or of part of the integuments of the
egg-like body, I cannot say; but the pulpy matter within the ovum, even
in its earliest stage, was included within an inner envelope or case.
In the next distinct stage (there being, however, slighter intermediate
changes) the posterior horn has shrunk, and become converted into a
bluntly-pointed conical termination for the whole body (fig. 17),
whereas the two anterior horns have approached each other on the
future ventral surface, and have increased considerably in length and
thickness, and contain within them the prehensile antennæ, which can be
externally seen, and which I dissected out of these horn-like cases.
The oval part of the egg-like larva (for I hardly know what to call it)
is now very slightly shrunk, being hardly more than 9/1000ths of an
inch in length. At this stage, these bodies adhere by the tips of their
anterior horns, containing the antennæ in process of formation, to the
inner tunic of the sack, and likewise in little groups one to another:
as the included prehensile antennæ ultimately become attached by cement
(proceeding, no doubt, as usual, from a modified portion of the ovarian
tubes), it seems probable that some cement may at this early period be
excreted, but I could not make out the exact means of attachment. The
egg-like larvæ are, also, aggregated round the tapering curled dorsal
appendages of the second and third segments of the body, and it is
possible that at this, or at an earlier period, these appendages may
act like the ovigerous fræna in the Lepadidæ, and serve to retain the
egg-like larvæ within the sack.
We come now to the last larval, or pupal condition, before the final
metamorphosis into the mature animal; the changes above described have
been, at least to a great degree, if not absolutely gradual; but the
pupa suddenly appears perfectly developed, from the moulting of the
last-described horned, egg-like larva. It is now a free animal crawling
about the sack of its female parent. It has increased a little in
length, as compared with the oval part of the egg-like larva in its
second stage, namely, from a little above 9/1000ths to 16/1000ths of an
inch: from the position of the prehensile antennæ in the two states,
I have no doubt that this increase of size is entirely due to the
anterior part of the pupa being doubled up whilst within the egg-like
larval envelope. The pupa in shape (fig. 18) somewhat resembles a
coffin, and is far less laterally compressed than other pupæ, and
hence can easily be placed either on its dorsal or ventral surface.
The prehensile antennæ are of large size: when the animal was alive,
they were concealed under and partially included within, the front
part of the carapace or shell, which in this condition was not so much
truncated as in the drawing given (fig. 18) of a specimen lying on its
back, with its antennæ protruded. Some specimens formerly examined for
me by Dr. Hooker, had their antennæ and whole ventral surface forced
outwards, apparently from the endosmose of the spirits of wine. The
whole dorsal surface, and the overlapping sides of the carapace are
elegantly punctured, and are formed of a rather brittle substance, here
and there supporting, especially at the front end, some fine and rather
long bristles,--which latter I have not seen on the pupæ of other
cirripedes. The ventral surface is very narrow towards the posterior
end of the animal; it is formed of thin, structureless membrane. On
this surface, close to the posterior end, there is a minute orifice,
through which three pairs of bristles are protruded, attached to (as
I believe) the rudimentary abdomen; the bristles apparently cannot be
withdrawn.
The antennæ, (fig. 18) as stated, are of large size compared to the
whole animal: they resemble, in all essential respects, the same organ
in other cirripedes. The ultimate segment is unusually thick;[152] it
is terminated by five bristles, one of which is longer than the others,
and stands rather separated from them. The disc-segment is large,
nearly circular, with the broad edges transparent and membranous;
on its posterior edge there is a single small spine. The second or
main segment, counting from the base, has a single spine on its upper
margin, close beneath the spine on the disc; it is articulated to
the disc-segment, a little way from the disc itself,--which is a
peculiarity I have not elsewhere noticed. The basal segment is thick
and not so short as usual. These organs are furnished with powerful
muscles. They are generally protruded alternately; and by the adhesion
of the sucker-like disc, the animal drags itself along. The sucker-disc
has great play, and when observing specimens alive, I compared its
action to that of a wrist-joint. The antennæ, when retracted within
the carapace, lie parallel to each other.
[152] As I have given, in my former volume on the Lepadidæ, p.
286, so many measurements of the antennæ, I may here add those of
Cryptophialus,--the length from the end of the disc to the end
of the second segment, (formerly called by me, erroneously, the
basal,) is 26/6000ths of an inch; the greatest width of the second
segment, 9/6000ths; the length of the little ultimate segment,
3/6000ths, and its width under 3/20000ths of an inch.
Posteriorly to the antenna, I distinctly saw the apodemes to which the
eyes are attached: I was not able to distinguish any middle fork to the
apodeme, which consequently does not resemble a UU, but U. The eyes are
dark purple, and, as usual, compound: in one specimen I counted twelve
ocelli within the common spherical envelope.
I could not distinguish any thorax, and certainly there is no mouth;
nor, from analogy, could the latter be expected, excepting as forming
part of the young cirripede: there are no natatory legs, which the
pupæ in all other cirripedes possess. Of the three postero-ventral
pairs of bristles, the most posterior or dorsal pair, differs from the
other two pairs in being considerably smaller, and in being mounted
on elongated pedicels: the two anterior pairs of bristles are strong:
the three pairs are articulated, one behind the other, on a small
body, apparently enclosed in a minute sack, and certainly attached all
round by membrane to the internal edges of the orifice, through which
the bristles are protruded. These bristles, when the pupa was alive,
were often moved, and served apparently to steady the body during the
alternate protraction and retraction of the prehensile antennæ. From
the fact of the pupa of other cirripedes having an abdomen, formed of
three segments, placed exactly in the same position as the minute body
here supporting the three pairs of spines, I believe this body to be
the abdomen. In other cirripedes only the posterior segment of the
abdomen bears spines, which are supported on little limbs or pedicels,
namely, the caudal appendages, the other segments being naked. But as
the mature Cryptophialus, unlike other cirripedes, has abdominal cirri,
the presence of spines on the corresponding abdominal segments in the
pupa, is explained and rendered probable: there can, I think, be little
doubt that the small terminal pair of spines, supported on elongated
pedicels or limbs, answers to the caudal appendages found in many
cirripedes.
The whole course of the metamorphosis is very peculiar. The gradual
changes in the egg-like larvæ (for I suppose they must be called larvæ)
from a simple oval egg, to pointed oval, to oval with three horns, and
lastly to oval with the two anterior horns larger, and the posterior
horn reduced to a mere point, seems to me very curious; and offers, as
far as I know, a unique case. It is interesting to reflect how perfect
a series, in the development of an animal, we have, in different
members of the Articulata,--from an ordinary egg, in which all the
changes go on unperceived, and whence a perfect animal is matured,--to
an egg-like larva which undergoes the changes just described, and which
turns into a pupa that does not eat or increase in size,--to a larva
which eats and increases in size, but undergoes only one great change,
as in most insects,--to a larva undergoing several great changes, as in
the case of ordinary cirripedes, before its final metamorphosis into
the mature animal. The first larval condition of other cirripedes, in
which there is a single eye, three pairs of thoracic limbs, and a much
elongated pointed body, covered by a prolongation of the carapace,
is here not fully developed or matured; but this stage is, I think,
clearly and very curiously indicated by the posterior horn of the
egg-like larva, which we may suppose represents the posterior pointed
end of the body, for it disappears in the succeeding stage, just as it
does in the second larval condition of other cirripedes. In the first
stage of ordinary cirripedial larvæ, the anterior horns are always
present, serving, as in the case of these egg-like bodies, to enclose
and protect the antennæ during their formation. The second egg-like
stage answers to the second larval condition of ordinary cirripedes,
as described (and figured, Pl. 30, fig. 1) in the introduction to the
Balanidæ. The third or pupal state is fully developed in all cases.
Finally, the pupa of Cryptophialus is peculiar in its punctured, hairy
surface, and in its shape, which, in being so much more depressed than
usual, retains an earlier larval condition; but its chief and highly
remarkable character consists in the entire absence of natatory legs;
and, in consequence, instead of there being a large sack within the
carapace, with an elongated orifice on the ventral surface, there is
only a quite minute orifice at the extreme posterior end of the animal,
through which the bristles, borne apparently on all three segments of
the minute abdomen, are protruded.
The pupæ of the male and female are exactly alike in all their general
characters, and probably in every point of detail; but my later and
more minute observations were made only on pupæ, which, from their
place of attachment, would certainly have turned into males. As these
pupæ, without any further metamorphosis, were developed into males, we
may, I think, safely infer that such is the case with the females: and,
consequently, that the whole course of the metamorphosis has been, in
this cirripede, seen and described. During this whole course, no food
could possibly have been obtained, for the pupa is destitute of a mouth
or organs of prehension, and the stock of cellular matter, enclosed
within the ovum, has been sufficient for all the above changes, and for
the final metamorphosis. We shall, moreover, immediately see, in the
case of the male, that the stock of cellular matter has also sufficed
for the development of testes, spermatozoa, and a wonderfully elongated
probosciformed penis.
MALE. Pl. 24, fig. 19.
By throwing pieces of the perforated shell of a Concholepas into
acid, I examined several scores of specimens of the Cryptophialus,
and on all, with the exception of a few young individuals, males were
attached. They were attached by cement, proceeding in the usual manner
from the prehensile antennæ, outside, to the edges of the upper half
of the disc formed of the thicker not-moulted membrane, by which the
female adheres in her chamber: hence the males are included in the
upper part of the same cavity with the female, into which they must
have crawled as pupæ. I found from one or two up, in one case, to seven
males, attached to the same female; four or five being the most usual
number. In the early part of January, when all my specimens were taken,
many of the males had not shed their pupal integuments, and of those
that had, the majority were immature, a few only having spermatozoa:
all the females had within their sacks, either ova including almost
perfect pupæ, or fully developed pupæ: we may, consequently, conclude
that these young males were maturing in order to impregnate the next
set of eggs.
The male, immediately after its metamorphosis from its pupal condition,
which has been fully described, is almost globular, but slightly
bilobed, and is formed of strong, structureless, transparent membrane,
including a mass of cellular matter, apparently without any included
organs: it is attached by about the middle, between the anterior and
posterior lobes, by the not-moulted prehensile antennæ. When the male
is mature, its greatest length, measured from the posterior end, where
the orifice is seated, to the anterior and blunter end, is about
13/1000 of an inch, and therefore rather less than the pupa, which
was 16/1000ths in length. Relatively to a full grown female, the male
slightly exceeds half the diameter of the toothed orifice leading into
her sack, see (_z_) fig. 1, Pl. 23. In the mature condition, (fig. 19),
one lobe, namely, the upper or posterior, has become more pointed, and
is terminated by a minute orifice, 8/6000ths of an inch in diameter.
This orifice is formed by a rim of thickened brownish membrane, which,
on what was the ventral surface, has a few very minute, but strong,
sometimes bifid spines;--in this one character, the male resembling
the female. The other and lower (homologically anterior) end or lobe
is broader, and contains a mass of cellular matter, which, from its
close resemblance in appearance and position to similar matter within
the male Alcippe, I have no doubt forms the contents of the testis.
In one single specimen, I succeeded in isolating a vesicula seminalis
of small size, containing perfectly distinct spermatozoa. Across the
middle, between the two lobes, close under the outer integument,
there is a broad layer of rather strong transverse muscular fasciæ.
I did not observe any eye, the presence of which I should have
expected from analogy. Internally there is no mouth, thorax, cirri,
or other organs, excepting the testis and vesicula seminalis just
mentioned, and an immensely elongated probosciformed penis, coiled up
and filling the rest of the inside of the sack down to the testis,
which latter occupies the whole anterior, and generally lower end
of the animal. This penis is plainly articulated, and includes fine
transversely-striated muscles: no doubt it can be protruded through
the minute orifice, and voluntarily moved about. Out of a male,
12/1000ths of an inch in length, I dissected a penis, which, when not
stretched, measured 50/1000ths of an inch in length; when a portion
was pulled between two needles, it could be stretched to apparently
three times its former length, and I should think that this organ
could be extended by the animal to, perhaps, even the 100/1000ths of
an inch,--that is, to between eight and nine times its own entire
length! The use of this enormously elongated penis obviously is,
that the spermatozoa of these males, which are so extremely small in
size, compared to the female, should all be conveyed within the sack,
and none be lost. It should be borne in mind, that the whole male,
including every part, is scarcely larger than a single ovum, of which
sometimes sixty have to be impregnated by only two or three males. In
a full-grown female, the distance from one of the attached males to
the middle of the orifice leading into the sack, is about the 5/100ths
of an inch, equal to the length of the coiled up, not-extended penis:
the further distance from the orifice of the sack to an ovum lying at
the bottom of the sack, would be almost 10/100ths of an inch, so that
the spermatozoa have to pass a distance of 15/100ths of an inch from
the testis of the male to the lower ova. I believe two thirds of this
distance would be passed safely along the probosciformed penis.
The resemblance between the male of Cryptophialus and of Alcippe is
truly surprising; and is the more wonderful, considering the great
dissimilarity of their pupæ. Hardly any characters can be pointed
out in which these males differ, excepting such as might have been
thought of only specific value, namely the relative proportions of the
different parts, and mere external shape. The peduncle growing a little
after the metamorphosis, in the male of Alcippe, and the prolongation
of its capitulum with the included oblique ligamentous fibres, are the
greatest differences. Having fully remarked, under Alcippe, on the
wonderfully rudimentary condition of these males, destitute as they are
of so many parts and organs, I will here say nothing further on these
singular creatures, destined to discharge their spermatozoa, die, and
be succeeded by a fresh set of short-lived male successors.
Order III.--APODA.
_Cirripedia, with the carapace reduced to two separate threads, serving
for attachment: body consisting of one cephalic, seven thoracic, and
three abdominal segments, all destitute of cirri. Mouth suctorial, with
the mandibles and maxillæ placed back to back, enclosed in a hood,
formed by the union of the labrum and palpi. Metamorphoses unknown._
The characters above given fully justify, I think, the formation of
this order; though it contains only one species, the _Proteolepas
bivincta_. The mere external appearance (Pl. 25, fig. 7), so
wonderfully different from that of every other cirripede, would by
itself prompt to this same conclusion. At first sight the Proteolepas,
if of fresh-water origin, might even have been mistaken for the larva
of some insect, fastened by two threads to its prey. The entire
absence of the three anterior segments of the head and therefore of
the carapace, or, speaking strictly, the mere rudiment of these parts,
forming an envelope to the two cement-ducts,--the absence of a stomach,
rectum, and anus,--the entire absence of thoracic and abdominal
appendages or cirri,--the absence of a probosciformed penis,--are all
negative characters, which might ensue from degradation, so common
with parasites; and which might, therefore, have been esteemed of not
high classificatory value. But the suctorial mouth, with the palpi and
labrum united into a hood, and with the mandibles and maxillæ reversed
or turned back to back, so as to be utterly incapable of prehension,
is a type of structure not hitherto met with, I believe, in any other
animal, and cannot be explained away by degradation. The formation of
the ova within the segments of the body, a peculiarity confined to
this one cirripede, evidently results from the non-development of the
anterior part of the head, within which the ova are usually formed;
but the compound structure of the vesicula seminalis is a peculiarity
which cannot be thus explained. Proteolepas has no particular affinity
to any other cirripede; it resembles, indeed, Cryptophialus in one
important point, but only in one point, namely, in the number of the
segments of its body. It is really beautiful to see how the homologies
of the archetype cirripede, as deduced from the metamorphoses of
other cirripedes, are plainly illustrated during the maturity of this
degraded creature, and are demonstrated to be identical with those of
the archetype Crustacean. I was at first inclined to rank Proteolepas
in one division, and all other cirripedes in another division of
equal value; but as it may be inferred from the characters of the
prehensile antennæ, that the pupa did not differ much, if at all in any
important character, from the pupæ of other cirripedes, I have thought
the three orders, which I have instituted, would be the most natural
arrangement. As any one looking at the drawing given of Proteolepas,
might very naturally feel inclined to protest against its being ranked
as a cirripede, I must reurge the importance of the pupal antennæ
being constituted on the common type, for from their structure, by the
law of correlation, that of the whole pupa may be inferred; and even
still more I must insist on the importance of the one great character
of the antennæ being cemented to the surface of attachment by matter
proceeding, as we shall see, in a modified state, from the great
ovarian sack. The structure, also, of the mouth (to a certain extent),
the segmentation of the body, though in appearance so peculiar, the
hermaphrodite condition, the single penis, the absence of oviducts, all
accord with, and taken together demonstrate, its cirripedial nature.
PROTEOLEPAS BIVINCTA. Pl. 24, 25, figs. 1-7.
_Hab._--Parasitic within the sack of the _Alepas cornuta_, from St.
Vincent's, West Indies, Brit. Mus.
_General Appearance._[153]--The entire animal, as already remarked,
curiously resembles, at the first glance, the larva of some insect.
It is rounded, but somewhat compressed, and tapers gently towards the
posterior end. It lies curved in an arc, the ventral surface being
concave, and the dorsal convex, but a little flattened dorsally at the
anterior and blunter end. Its length, if straightened, would rather
exceed one fifth of an inch. The body consists of eleven segments,
which, excepting the three terminal, are conspicuously plain. The
first segment is surmounted by a rather small mouth, which any one
would, assuredly, at first consider as the entire head, though he
would in vain search here for eyes, antennæ, or other parts of the
three anterior cephalic segments. On the dorsal surface, low down on
the second segment of the body, two, quite flexible, thin, but strong,
flattened threads arise, which terminate in a pair of prehensile
antennæ, having the usual cirripedial structure. From the penultimate
or disc segment of these antennæ, cement has been excreted, by which
the antennæ are firmly cemented low down to the rostral end of the sack
of the cirripede, the _Alepas cornuta_, on which it is parasitic: hence
Proteolepas lies with its back downwards, and with its ventral concave
surface fitting the convex body of the Alepas: its mouth lies under
the middle of the soft prosoma of the latter cirripede, which I cannot
doubt that it lacerates and sucks.
[153] I may be permitted to premise, that though I procured only a
single specimen, yet perceiving its very singular nature, I took
such care and length of time in the dissection, and repeated every
observation so many times that I think reliance may be placed
on the description here given. Fortunately I had acquired, from
dissecting many much smaller specimens of various cirripedes, all
the advantage which full experience could give me, when I commenced
on Proteolepas.
_Mouth._--The mouth is suctorial, and is constructed on a different
plan from that in any other cirripede, or, indeed, in any other,
as far as I know, articulated animal. It is narrower, in both a
longitudinal and transverse plane, than the first segment of the body,
and is distinctly separated from it. The lower part on the ventral
side, is protuberant and rounded. The summit is square, and is formed
by the crest of the labrum, with two large palpi (_d_, fig. 3), having
nearly the usual form amongst the Balaninæ, and pointing towards each
other, but differently from in any other Cirripede, they are united
for their whole length to the labrum, and by their extremities to
each other. These parts together thus form an arch or hood, within
which stand the other gnathites. The palpi are roughened by groups
of very minute spines. At their bases they can be obscurely seen to
be separated from the rest of the mouth by an oblique joining or
articulation. The back of the mouth is formed entirely of the labrum,
which becomes narrow towards its base: it is, from top to bottom,
20/1000ths of an inch in height. Within the hood formed by the palpi
and labrum, a pair (_c_, fig. 3), of very singular, compounded,
mandibular organs project freely, straight up, with their convex outer
edges placed parallel and close together, and their teeth pointing
directly from each other, so that they stand in a reversed position
compared with the jaws of all other cirripedes, and are absolutely
incapable of prehension.
This compounded organ is singularly small compared with the palpi and
labrum: it is narrow, being about 5/3000ths of an inch in width, but
is produced upwards, so that a considerable length projects freely,
and the rounded, properly external, margin can be traced down for a
length of about 20/3000ths of an inch. In a lateral view of the mouth,
the extreme tip of the mandibular organ could sometimes be seen just
projecting out of the hood. The mandibular organ, when separated and
carefully examined, presents the appearance, represented from a camera
drawing, in (Pl. 24, fig. 2): we here see three groups of teeth; of
these the lower set (_c_) consists of blunter teeth, placed more
transversely, and easily separated from the others, and altogether
clearly appears like a distinct organ. I do not feel nearly so sure
regarding the other two sets; my first impression was strongly that
they were distinct organs, closely united laterally together,--one
(_a_) probably representing the mandibles, and formed into a single
large tooth; the other (_b_) formed of three teeth, and probably
representing the outer maxillæ; the first-mentioned set of teeth, which
seemed to me to arise from between the other two sets, being the inner
maxillæ. If this view (and it must be remembered how excessively minute
the parts are) be not, as I now suspect, correct, we must suppose
that the outer maxillæ are aborted, and we have seen some tendency
towards this in other cirripedes; the compounded organ being formed
only of the mandibles (having on this view four teeth) and the inner
maxillæ. As far as the mandibles are concerned, their existence, I may
remark, is plainly shown by the presence of the palpi, which in all
cases belong to and form part of the mandibles. The ventral surface
of the mouth, immediately beneath the free portion of the compounded
mandibular organ, consists of a triangular projection, but I could
see no appearances to make me suppose that this part represented
the outer maxillæ. The compound organ--in general shape, and in the
oblique manner in which the front part is cut off and terminates in
ligamentous apodemes, to which muscles are attached,--presents an
unmistakable likeness to a mandible. It is hollow within, and muscles
appear to extend some way up, perhaps to the transversely toothed
portion, which represents, as I believe, the inner maxillæ: these two
groups of teeth, anyhow, seemed to have some power of sliding over each
other, and altered their positions during the course of dissection. On
each side of the mouth, there is a muscle attached by its lower end
to the basal edge of the labrum, and two others, one above the other,
attached by their lower ends to about the middle of the labrum; these
muscles, which are distinctly striated or voluntary, I infer, from
analogy, run up to the ligamentous apodemes of the compound mandibles.
There appeared to be other more delicate muscles attached to the basal
articulation of the mouth on the ventral face, and these, I presume,
would run to the supposed inner maxillæ.
The mouth in forming a prominence separated by a distinct articulation
from the body, and in the union of the palpi and labrum (though here
carried to excess), is constructed so far on the cirripedial type; but
how are we to account for the extraordinary reversed position of the
united mandibles and maxillæ, with their backs almost touching each
other, and their toothed edges twisted round so as to face outwards
in a manner unexampled, I believe, in any other articulate animal? It
might, perhaps, be at first suspected, that the compounded mandible had
not really been twisted round, but that the teeth had been abnormally
developed on the outer convex margin: this view, however, certainly
cannot be admitted, for the properly outer convex margin can be traced
running far down the mouth, in a manner utterly inexplicable, if this
were really the inner side; and equally inexplicable on this view would
be the position of the ligamentous apodemes. Hence I cannot doubt
that this compounded mandibular organ has really rotated on its axis;
and if the course of development could be followed, I suspect that
the twisting would be seen to be effected as follows: we know in all
cirripedes that the outer and inner maxillæ, and to a certain extent
the mandibles, instead of facing each other, are directed towards the
labrum; they therefore have already been twisted round a quarter of a
circle, as may be seen in the diagram (Pl. 24, fig. 4), copied from the
mouth of Ibla. Now let us drive inwards the front of the mouth, along
a narrow medial line; these organs would then (fig. 5) be compelled to
turn round a quarter of a circle more, and so face directly outwards.
In this process, the integument between the lower and outer part of
the mandible and the base of the palpus, which normally are in close
contact, would have to be greatly stretched. By a movement of this
order, the mandibles would come to stand posteriorly or exteriorly to
the other gnathites; and as far as I could make out (previously to my
having any theory) the large single toothed portion of the compound
organ which most resembles a mandible, did really stand outside the
other toothed portion.
With respect to the action of this singularly constructed mouth; if
its ventral and oblique surface were applied to any yielding object,
as the adjoining soft prosoma of the Alepas, the compound mandibles
would be worked within an absolutely closed chamber. The action of
these mandibles would be to make a transverse slit, and subsequently
to serve as a grapnel to keep the mouth closely adpressed to its
prey: the other teeth might act in keeping the wound open. When the
mouth was thus closely adpressed over a wound, the great power of
shortening the whole body which the animal possesses (the œsophagus
being closed), would, by the subsequent action of the elasticity of the
outer membrane, almost certainly create suction, and thus cause the
nutritious juices of the Alepas to flow into the body of the parasite.
Hence I have called the mouth suctorial.
_Body._--This, as already stated, consists of eleven segments, of
which the three posterior (abdominal) smaller segments can hardly be
distinguished, without dissection, as separate from each other. The
body is mainly occupied by a vast ovarian sack (_e_, _e_, fig. 7),
filled by innumerable ova: and the three posterior segments by small
testes and their vesiculæ seminales (_i_): but I shall return to
the internal anatomy. The outer membrane, lined by delicate corium,
is thin, transparent, elastic, and covered by groups of excessively
minute blunt little points. The segments can be plainly distinguished
by their outlines, especially on the ventral surface; but they are
rendered unmistakably distinct by the attachment of the muscles; they
can also be perceived when the external membrane is perfectly cleaned,
by yellowish lines. The muscular system is highly symmetrical and
simple: along all eleven segments, there is a narrow, medial, ventral
and dorsal clear space; on both sides of which space there is a band
of longitudinal muscles, which, though encroaching on the two sides,
and rather largely on the dorso-lateral sides, may be called the
ventral and dorsal muscles. These muscles are striæ-less, which is
the case with the homologous posterior thoracic muscles in some other
cirripedes: on the dorsal surface (lower surface in fig. 7) they are
more spread out, and consist, on each side of the medial line, of four
ribbons: this seems to be the case on the ventral side, but the ribbons
are here much more confluent: in the seventh and eighth segments, the
ribbons become broader; but in the ninth, tenth, and eleventh, or
three posterior segments, they become much narrower, and some of the
fasciæ disappear, so that these muscles can hardly be seen from the
outside. Each separate ribbon expands a little at its two ends, which
are attached to the articulations separating the successive segments:
I carefully observed that they did not pass over at either end to the
adjoining segments: hence their action must be either simply to shorten
and arch each segment separately; or when acting together, to shorten
the whole body, or perhaps the ventral or dorsal surface by itself.
In the first segment, and in the three posterior segments, these
longitudinal muscles alone occur; but on the seven segments, from the
second to the eighth inclusive, there are other oblique latero-ventral
muscles. These muscles lie within the longitudinal muscles, and
adhere pretty firmly to the coat (_e_, _e_, fig. 7) of the great
ovarian sack. At their ventral extremities they are attached, near the
anterior margin of each segment, beneath the point of attachment of
the longitudinal fasciæ, and thence they run posteriorly in an oblique
line to the anterior margin of the next succeeding segment, where
they are attached: so that these muscles run obliquely from segment
to segment. The first of these oblique muscles, lying chiefly within
the second segment of the body, is thinner and longer than the others:
those within the third and fourth segments are short: those within the
fifth and succeeding segments extend, at their dorsal (or lower in
fig. 7) extremities, as far as the outer dorsal longitudinal fasciæ:
those within the seventh segment are broad and short, and cross the
longitudinal muscles at only a small angle. In the eighth segment,
there is an oblique lateral muscle, like that in the seventh segment,
running from the ventral surface towards the dorsal surface; but there
is in addition a second oblique lateral muscle, rising from the dorsal
surface, and running towards the ventral surface. This muscle does
not occur in the other segments, but in the fourth segment, at the
dorsal end of the oblique latero-ventral muscle, there may be seen
a small branch of fibres, at right angles, which seems to represent
a muscle homologous with that just mentioned in the eighth segment:
obscure traces, moreover, of similar fibres, can be detected in some
of the other segments: had these oblique latero-dorsal muscles been as
fully developed in the seven anterior segments of the body, as on the
eighth segment, the whole muscular system would have been perfectly
symmetrical. The oblique latero-ventral muscle in the sixth segment is
distinctly striated transversely; but this is not the case with most
of the other muscles, if with any of them; I cannot account for this
difference. The muscles of the gnathites are the only other voluntary
muscles in the animal's body.
_Homologies of the Body._--It will hereafter be, I think, clearly
shown, that when the shell and integuments of the pupa of Proteolepas
are shed, no carapace or general covering for the body is formed; the
three anterior segments of the head, the backward prolongation of
which (as has been elsewhere explained) certainly forms the carapace
of ordinary cirripedes, being here almost absolutely aborted. In every
cirripede the mouth is formed of three pairs of gnathites, which, no
one will doubt, rise from the fourth, fifth, and sixth segments of
the head: here in Proteolepas, the mouth, even on the view of the
mandibular organ on each side being compounded of only two gnathites,
sufficiently resembles the ordinary cirripedial type to make it very
probable, that if examined in the earliest stage of its development,
three pairs of gnathites would be discovered. In accordance with
this conclusion, the segment succeeding the mouth (_i. e._, the
first segment of the body in fig. 7) homologically is the seventh,
or last cephalic segment. The succeeding seven segments, of course,
are the seven thoracic segments, and the three posterior segments
are abdominal; the latter are not developed in ordinary cirripedes
when mature, but are present during their pupal condition. Now this
conclusion, which is, in fact, deduced from what we know of the front
part of the head in other cirripedes, both larval and mature, appears
to me most satisfactorily confirmed by the differences in the muscular
system of the segments in Proteolepas. In no other way, I believe,
can it be explained, why the last cephalic segment and the three
abdominal segments should differ from the seven thoracic segments,
in the entire absence of the oblique lateral muscles. The abdominal
segments, moreover, differ a little in shape, in the indistinctness
of their articulation, in the thinness of the longitudinal muscles,
and even in their contents. With respect to the two threads enclosing
the cement-ducts, which spring from the second segment of the body
(or first of the thorax), and which terminate within the prehensile
antennæ of the pupa, we shall hereafter see that their apparently most
anomalous position, and even the flattened shape of the dorsal surface
of the two anterior segments of the body, all accord perfectly with the
homologies just given.
_Alimentary Canal._--The œsophagus is thin, and for a cirripede short,
for it extends only half-way down the first segment (_i. e._ last
cephalic) of the body; the lower end, which is slightly dilated, nearly
touches the anterior end of the great ovarian sack. At its upper end,
it is surrounded by delicate, striæ-less constrictor muscles; and there
are others radiating outwards, evidently serving to open it: the lower
part of the œsophagus, differently from other cirripedes, is destitute
of muscles, and is only coated by a thin layer of corium, which would
serve to produce a new œsophagus at each exuviation. Strange as
the fact may be, I am prepared to assert that there is no stomach,
rectum, or anus. As I was able to trace so distinctly the œsophagus,
and likewise the generally far smaller orifice and ducts of the male
generative organs, I consider it quite impossible that I could have
missed the stomach. The rectum and anus are absent in Alcippe: and the
absence of a stomach is here in some degree the less surprising, as the
structure of the mouth shows that Proteolepas must live on the already
elaborated fluids of the Alepas, to which, being a cirripede, it is
allied. It is of some importance to observe, that the œsophagus is
fitted with muscles simply for shutting and opening it, the wave-like
swallowing action of which other cirripedes are capable, being,
apparently, here impossible; but the contraction of the body and its
subsequent expansion, the œsophagus being opened, would allow the blood
of its prey to flow inwards.
The nervous system must be much atrophied, for I could not detect
it, and the small size of the animal is not sufficient to account
for this: I wish I could have seen this system, for then I should
almost certainly have beheld an articulate animal without a trace of a
supra-œsophageal ganglion. There is no eye, but such could hardly be
expected, as the anterior cephalic segments are aborted. There are
no branchiæ. I may state that within the abdomen, along the dorsal
surface, there was either a lacuna or a delicate vessel, apparently of
a circulatory nature, of considerable diameter, which, near the extreme
posterior end of the body, gave out branches.
_Female Reproductive Organs._--The eight anterior segments of the body,
with the exception of a small space at the two ends, are occupied by
an immense (_e_, _e_), opaque, ovarian sack. The tissue forming it
is delicate, and presents a peculiar cellular aspect: it is slightly
attached to the corium on the ventral surface of the body, and to the
oblique latero-ventral muscles. Internally, at the anterior end, it is
thickly coated by cellular matter, the cells varying from 4/6000ths to
less than 1/6000th of an inch in diameter, becoming in parts confluent,
and the whole forming a dark orange-coloured mass. In the more central
parts of the sack this cellular matter became aggregated into little
pellets, which, in proceeding towards the posterior end of the sack,
gradually increased in size, from about (4 to 6)/1000ths of an inch in
diameter, and at last appeared as almost mature and perfect ova of a
broadly oval figure. Their size, as we see, is small, and their number
almost infinite. I carefully examined all round this ovarian sack, and
could detect no oviducts; nor from analogy could they be expected: I
have no doubt that the ova burst forth by the rupture, probably, of the
posterior end of the sack and of the overlying corium; and that they
accumulate beneath the external membrane of the body, until this is
moulted, the rupture beneath being in the meantime healed, when they
are freed, or perhaps temporarily protected in the old moulted envelope
of the body.
On each side, within the first two segments of the body, and projecting
a little before the great ovarian sack (_e_), two gut-formed organs
(_f_) may be seen, even from the outside, owing to their opacity and
dark colour. They lie near the external surface; the first pair of
latero-ventral oblique muscles passing between them and the ovarian
sack. They are formed of a branching, grape-like mass of opaque,
orange-coloured cells. They are intimately united, at their posterior
extremities, to the ovarian sack, and I believe open into it; but I
cannot say that I demonstrated this. From their absolute identity in
structure, and similarity in position, namely, on each side of the
lower end of the œsophagus, no doubt is left on my mind that these
bodies answer to the true ovaria, which are situated within the body of
other cirripedes; and that the ovarian sack answers to the inosculating
and branching ovarian tubes and cæca, which fill the peduncle, or cover
the basis in other cirripedes, but here, from the absence of these
parts, necessarily occupying the body.
_Male Organs._--The whole surface of the ovarian sack, the space before
it, even to within the lower parts of the mouth, the posterior half of
the last thoracic segment, and especially the whole three abdominal
segments, are completely netted by branching delicate vessels or ducts
terminating in spherical glands about 1/2000th of an inch in diameter.
These little glands include a brownish pulpy centre, and sufficiently
resemble the testes of other cirripedes in appearance, position, and
connecting ducts, to make me believe that such is their nature. I
may remark that in the more central parts of the abdomen the glands
and ducts seemed to be in process of formation by the confluence of
cellular matter, and in some other cirripedes I have suspected that
the testes are periodically renewed, or at least redeveloped from
an undistinguishable condition. Within the posterior half of the
abdomen, some of the ducts become thicker and unite, others joining in
laterally, so as together to make a dark chord, 7/2000ths of an inch
in diameter. Until dissecting this chord, I thought it was a single
vesicula seminalis, but it separated into several rather thick ducts
or vesiculæ. I was not able to remove from within them the contained
matter, but it appeared very finely and longitudinally flocculent, like
spermatozoa not quite matured. In accordance with the immature state of
the contents of the ovarian sack, in all probability these ducts would
hereafter have become greatly enlarged, and have formed a compound
vesicula seminalis of considerable size. The dark chord, formed
by their union, contracts as it enters the rudimentary penis, and
terminates in a very minute orifice on its apex. The penis consists of
a papilla, only 3/4000ths of an inch in length, situated on the extreme
point of the abdomen, but rather towards the ventral surface.
_Metamorphosis._--In accordance with the general law of the correlation
of parts, it may be inferred, from the description and measurements
of the pupal antennæ immediately to be given, that this abnormal
creature was developed within a pupa of the same general structure,
and of about the size, as the pupæ whence Scalpellum, Alcippe, and
many other cirripedes are developed. As the ova are of remarkably
small size, indeed I have seen no others quite so small, it is certain
that the larvæ, as in the case with all other cirripedes, excepting
Cryptophialus, must undergo several metamorphoses, and increase much in
size, before attaining their pupal condition.
_Attachment._--The animal is attached, as already stated, to the
sack of the Alepas by two threads, rising close together from the
medio-dorsal line, near the posterior end of the second segment of
the body. These threads are attached likewise close together at their
further ends, by the antennæ, into which they enter. They are flattened
and strong, yet quite flexible, with a somewhat sinuous surface: they
were, in this specimen, 42/1000ths of an inch in length, and a little
above 3/1000ths in diameter: where joined to the thoracic segment they
were a little contracted. Their structure in this specimen could be
made out (Pl. 24, fig. 1) with perfect distinctness. Their transparent
outer tunic (_e_, fig. 1) is 1/2000ths of an inch in thickness, and is
continuous with that (_d_) enveloping the whole body, but is abruptly
and considerably thicker than this membrane; and hence a very slight
collar is formed outside, round the line of junction of each thread
with the body. The delicate corium (_c_) lining the external membrane
of the body runs, at least someway, down these threads. It was likewise
indisputably evident that the membrane (_b_), for I separated it by
dissection, forming the great ovarian sack, together with the cellular
contents of this sack (_a_), entered and extended down both threads.
It should, also, be particularly observed, that the coarsely cellular
matter within the ovarian sack, immediately that it entered the tube
formed by the membrane of the ovarian sack, suddenly changed its
appearance into a homogeneous, stiff, pulpy matter, which retained the
same appearance all down the threads to within the antennæ. This finer
matter readily separated from the coarser cellular matter within the
sack, but was not divided from it by any septum or membrane. Some way
within the threads, the corium, the membrane of the ovarian sack, and
the contents appear (_e′_), as seen from the outside, to become, and
perhaps really are, blended together. These threads could not have
been originally formed of their present length, and must therefore
have been added to during the growth of the animal; but from their
entering the not-moulted antennæ, and from the animal being permanently
attached by them, they cannot have grown, by means of the moulting of
their integuments; hence I conclude that at each period of growth and
exuviation they have been added to only at their upper ends, where
there is a sort of collar, or line of growth; and where, I may remark,
the lining corium is alone well developed. We shall presently see the
bearing of these remarks.
These threads contract to about half their former diameter as they
enter the old prehensile antennæ of the pupa, within which they are
firmly attached. Each thread, with its three tunics apparently blended
together, can be traced to the extremity of the disc-segment (_g_),
where the included matter seems to have burst forth. The whole disc and
the terminal segment of both antennæ are enveloped, close together,
in cement, formed into two almost separate little capsules, by which
they adhere very firmly to the integuments of the Alepas. The cement
required to be removed before the antennæ could be plainly seen. The
cement presented all the usual characters, namely, its homogeneous
laminated structure and its yellowish colour. The cement in the case
of the male Ibla, which is parasitic within the sack of the female
Ibla, affects the corium and fibrous matter beneath the chitine-tunic,
and causes them to adhere together, and thus prevents the male from
being cast off each time that the inner tunic of the sack of the female
is moulted: exactly so has the cement of the Proteolepas affected
the integuments of the Alepas. The only difference between ordinary
cement-ducts and the two threads here described is, that the ducts,
in both cases formed by the prolongation of the coat of an ovarian
receptacle, are here protected by a thick outer membrane, lined, at
least in the upper part, by corium; whereas, in the Lepadidæ the two
ducts are included within the peduncle, and are therefore protected by
one common membrane, lined of course by corium; and this membrane, we
shall presently see, is homologous with that separately investing the
two threads.
The antennæ differ remarkably little, considering the anomalous
character of the mature animal, from the same organ in other genera;
they come nearest, perhaps, to the antennæ of Ibla. The length of the
disc (_g_, fig. 1) and great succeeding segment (_f_) together is
40/6000ths of an inch. The lower segment has its basal articulation
only slightly oblique, showing that, as in Alcippe and Ibla, it was
probably articulated near the anterior end of the pupal shell: it is
of nearly the same width throughout.[154] The disc (_g_) is remarkable
from its great proportional length; it is hoof-shaped, with the outer
side rather protuberant, and the end pointed. The ultimate segment
(_h_) is of moderate size: as in Ibla, it has a shoulder or notch on
its inner side near its end, bearing two long spines; and probably
there were originally three or four spines on the square broad upper
end, but these have been broken. This segment is articulated unusually
near to the end of the disc.
[154] As I have given the measurements of the antennæ in so many
genera, I will give these: second (_f_, fig. 1) segment, 24/6000ths
of an inch in length, and (8-9)/6000ths in width. Disc, 16/6000ths
in length, and 8/6000ths in width. Ultimate segment, 6/6000ths (?)
in length, and 10/20,000ths (?) in breadth.
The foregoing remarks on the two threads by which Proteolepas is
attached, are, independently of their relation to this individual
animal, of considerable interest. In my volume on the Lepadidæ, I have
stated, after repeated and rigorous examinations (for I was well aware
how singular the facts were), that in _Conchoderma aurita_ and in
some other genera, the cement-ducts, which entered the pupal antennæ,
could be traced till they joined a gland, the coat of which gland was
absolutely continuous with the coats of the adjoining and continuous
ovarian tubes, of which it was only a modified portion; and what was
still more remarkable, that the matter within the gland was continuous
with, and differed only from, the cellular matter within the ovarian
tubes and cæca (from which ova were in the act of formation), by being
more homogeneous and more coherent. Furthermore, I have shown, that
in Ibla an ovarian tube, becomes by a very small change, namely, by
a double flexure and slight thickening of its coat, converted into a
gland, and thus acquires the power of affecting the cellular ovarian
matter and changing it into cement. Now, in Proteolepas, the great
ovarian sack replaces the ovarian tubes and cæca; and we here see
the very same relations even still more plainly; for the coat of the
ovarian sack is indisputably continuous with that investing or forming
the two cement-ducts within the two threads; and immediately that
the coarse cellular matter, which within the ovarian sack is being
converted into ova, enters the upper contracted end of the cement-duct,
by some power, we must suppose, inherent in its coat, it is converted
into cement, which debouches with all its usual properties through the
pupal antennæ. I may venture to reaffirm that nothing could be plainer
than this structure, or be in more striking conformity with my previous
observations, given in the introduction to the Lepadidæ.
* * * * *
I can hardly express the perplexity which I felt when I first examined
Proteolepas, and when I naturally mistook the mouth for the entire
head, for I saw, as I thought, the antennæ in direct connection with
the second segment of the body, posteriorly to the mouth! It was quite
as monstrous and incredible an inversion of the laws of nature, as
those fabulous half-human monsters, with an eye seated in the middle
of their stomachs. After a time, I perceived that the following
considerations removed all difficulty, and brought Proteolepas into the
type of other cirripedes.
_Firstly_: in ordinary cirripedes, the two cement-ducts can be traced
up from the cemented antennæ to the glands, formed by a part of the
ovarian branching cæca; and the latter can be traced to where they
enter, as two simple tubes, the body of the animal, at a medio-dorsal
point, a little anteriorly to the prosoma, or second thoracic
segment.[155] From what is actually seen in the complemental male
of _Scalpellum Peronii_, and from what may be inferred from the
structure of these parts in the pupæ of all cirripedes, there can be
no doubt that if the ovarian cæca were in any case not developed, the
cement-ducts would enter the body at the spot where the two simple
ovarian tubes, which serve to unite the ovarian cæca with the true
ovaria, do enter. Now if we look at the drawing (Pl. 25, fig. 7) of
Proteolepas, we shall see that the cement-ducts enter the body at a
medio-dorsal point, a little anteriorly to the second thoracic segment,
and therefore in the normal position.
[155] This may be partially seen in the section, fig. 1, of
Balanus, on the same plate (25) with the figure of Proteolepas;
here (bearing in mind that Balanus is a much modified form)
(_z_) shows the pupal antennæ, within which, whilst young, the
cement-ducts are included, and are directly continuous with the
layer of branching ovarian cæca (_g_), which are prolonged up
to the ovaria as a pair of simple tubes (only one being here
represented), entering the body above the upper margin of the
prosoma (_c_). The prosoma of Balanus, I may add, answers to
the segment ^2 _t_ in fig. 7 of Proteolepas; (_e_) the mouth
in Balanus, of course corresponding with (_m_) the mouth of
Proteolepas; the segment ^1 _c_ and ^2 _t_ of the latter, are in
Balanus aborted or confluent, at least on the ventral surface; and,
lastly, the whole great shell of Balanus, the sack with its muscles
and the branchiæ, and the opercular valves with their muscles, are
all represented in Proteolepas merely by the outer membrane of the
two threads (_g_), which enter the pupal antennæ!
_Secondly_: the external membrane of the two threads, investing the two
cement-ducts, it should be remembered, is not moulted, and is added to
during growth (being lined internally by corium), only round the upper,
collar-like edge.
_Thirdly_: the external covering or carapace of every young cirripede,
at the period of its metamorphosis, enters, at its lower end, the
cemented antennæ, in the form of two short tubular prolongations,
by which alone, at first, the cirripede adheres to the surface of
attachment; within these prolongations the cement-ducts are included. I
have, moreover, seen instances, as in _Conchoderma aurita_ and in the
male of Ibla and Alcippe, in which these tubular prolongations, lined
internally by corium, were increased a little in length, so as to form
a trouser-like termination to the peduncle. That the forked extremity
should be a little more developed, and so be converted into a pair of
short tubular threads, cannot be considered as very improbable.
_Fourthly_: in the male Ibla the capitulum is so much atrophied that it
does not enclose the thorax or mouth, but still an elongated support or
peduncle is left. But it would be no very violent assumption to imagine
the peduncle, which does not essentially differ from the capitulum, to
become likewise rudimental,--to grow smaller and smaller, and shorter
and shorter, till the merest remnant was left at the spot where it
entered the cemented antennæ. And in the last paragraph it has been
shown that it would be no violent assumption to imagine this lower end
of the peduncle, where it enters the antennæ, developed into two short
thread-like prolongations.
_Lastly_: it is certain, from the existence of the prehensile antennæ,
that Proteolepas was developed within a pupa, probably differing in no
very essential respect from the pupæ of other cirripedes. Therefore, in
accordance with all analogy, we may believe that the position[156] of
the young Proteolepas (probably much coiled up, with a deep fold close
under the mouth) within the pupa, the general form and structure of
the latter, and the course of the cement-ducts, did not _essentially_
differ from the imaginary figure given, Pl. 25, fig. 6. Now, at the
period of the metamorphosis, let us imagine that no general covering
or carapace was formed, except a small portion on the ventral surface,
round the cemented antennæ. Let us further suppose this remnant to be
specially developed (as in the case of some cirripedes) into a short
trouser-like prolongation, entering the antennæ; and subsequently, in
accordance with the almost universal laws of growth in cirripedes, that
this portion was never moulted, but continued to be added to, during
growth, only at its upper end. By this means we should produce every
leading peculiarity of the _Proteolepas bivincta_. As this parasite
lives within the sack of another cirripede, and is protected by the
capitulum of the latter, we can understand, in accordance with the
usual admirable economy of nature, the absence of any general covering
for its body. We can now, also, understand the structure and manner of
growth of the two threads by which it is bound to its prey; and the
connection, at first so strange and perplexing, between the old pupal
antennæ and the second segment of the thorax. I am convinced that no
other explanation than that here given, will accord with the relations
of the several parts and organs of Proteolepas. Consequently, I fully
believe that we here see an articulate animal in which the whole of
the three anterior segments of the head have been, during the act
of metamorphosis, absolutely aborted, with the exception of a mere
rudiment on the ventral surface, near the anterior end, round the old
antennæ, and which rudiment has been specially developed as a covering
for the two cement-ducts. As the pupal antennæ are, homologically, the
second pair of antennæ, we may further infer that this modified remnant
of the carapace, investing the two threads, belongs to the third
cephalic segment.
[156] Any one who has not specially attended to the metamorphoses
of ordinary cirripedes, who looks at the imaginary figure of
the young Proteolepas, will feel much surprise at the relative
positions of the parts; for the mouth and the first and even
second segments of the body stand posteriorly (_i. e._ above in
the figure) to the succeeding segments of the body, in relation to
the carapace of the pupa; but this is only in accordance with the
remarkable change in position (as explained in the introduction,
p. 123, pl. 30, fig. 2), amounting almost to inversion, which the
whole thorax of every young cirripede undergoes within the pupa,
whilst the anterior cephalic portions and general covering are
developed conformably with the pupal carapace, whence it arises
that the _dorsal_ surface of that part of the thorax immediately
succeeding the mouth becomes attached to the _ventral_ internal
surface of the carapace. I believe that the peculiar flattened
dorsal outline of the first two segments of the body of Proteolepas
is due to these parts having been formed in contact (as represented
in pl. 25, fig. 6) with the straight ventral surface of the
carapace of the pupa. To place the young Proteolepas, and at the
same time the carapace of the pupa, with all the parts in proper
homological sequence, it would be necessary to seize the posterior
end of the abdomen (_a_), and pull till the dorsal surfaces of the
first and second segments of the body, separated from the ventral
internal surface of the carapace, and stood posteriorly (_i. e._
above in figure) to the mouth, which latter would thus also have to
rotate a quarter of a circle, so that the orifice would come to be
directed outwards. Then every part would stand, in accordance with
the archetype crustacean structure, in due order; but the three
confluent anterior cephalic segments, forming the front part and
carapace of the pupa, would, as in the case of all cirripedes, be
of disproportionately large size in relation to the rest of the
body.
SYNOPSIS
ET
INDEX SYSTEMATICUS.
_Ordinum, Familiarum, et Generum Cirripediorum et recentium et
fossilium._
CLASSIS CRUSTACEA.
Sub-classis CIRRIPEDIA.
Crustacea ex anteriore capitis parte defixa, cæmento in hunc usum ex
ovariorum portione ad id specialiter modificatâ emisso. Archetypus è
segmentis 17 compositus, quorum 3 priora magna, in carapacem sæpissimè
conformata, quæ non omnino exuitur et varios motus efficit: antennæ
nullæ: oculi rudimentarii: os prominens, formatum è labro, palpis,
mandibulis, et duobus maxillarum paribus, quæ omnia partim confluunt:
thorax ad superficiem internam sternalem carapacis affixus, plerumque
cum membrorum captantium biramorum, multiarticulorum paribus 6:
abdomen plerumque rudimentarium: branchiæ, si quæ adsunt, ad inferiora
carapacis latera affixæ: plerumque bisexualia; in unisexualibus, mares
fœminis parasitice inserti: penis unicus, plerumque probosciformis,
ad posteriorem abdominis extremitatem situs: oviductus nulli:
metamorphoses multiplices.
ORDO I. THORACICA. (Darwin, 'Balanidæ,' p. 30.)
Cirripedia quibus pro carapace est aut capitulum pedunculatum, aut
testa operculata cum basi. Corpus è 6 thoracicis segmentis, ferè cum
6 cirrorum paribus, constat. Abdomen rudimentarium, sed sæpe cum
appendiculis caudalibus. Oris labrum motus proprios non efficit. Larva
primò monocula cum 3 crurum paribus, postremò binocula cum 6 crurum
thoracicorum paribus.
Familia 1. BALANIDÆ. (Darwin, 'Balanidæ,' p. 33.)
Cirripedia sine pedunculo: scuta et terga musculis depressoribus
instructa: reliquæ testæ valvæ inter se immobiliter conjunctæ.
Sub-Familia 1. BALANINÆ. (Darwin, 'Balanidæ,' p. 175.)
Rostrum cum radiis, sed sine alis; valvæ testæ laterales omnes, ex uno
latere alis, ex altero radiis instructæ: parietes ferè aut porosi aut
ad interiorem superficiem longitudinaliter costati.
[_Sectio_ †.]
Scutum et tergum inter se articulata aut mutuò interclusa: branchiarum
unaquæque ex unicâ plicâ constat.
1. _Genus_--BALANUS, _Auctorum_. (Darwin, 'Balanidæ,' p. 177.)
Valvæ testæ 6; basis calcarea aut membranacea; valvæ operculares
subtriangulares.
2. _Sub-Genus_--ACASTA, _Leach_. (Darwin, 'Balanidæ,' p. 302.)
Valvæ testæ 6; parietes et basis non porosa; basis calcarea,
cyathiformis, non elongata. Spongiis, aut rarò Isidis cortici, affixa.
3. _Genus_--TETRACLITA, _Schumacher_. (Darwin, 'Balanidæ,' p. 321.)
Valvæ testæ 4; interdum inter se externè confluentes: parietes poris
perforati, multis plerumque seriebus; basis plana, irregularis,
calcarea aut membranacea.
4. _Genus_--ELMINIUS, _Leach_. (Darwin, 'Balanidæ,' p. 345.)
Valvæ testæ 4; parietes non porosi; basis membranacea.
5. _Genus_--PYRGOMA, _Leach_. (Darwin, 'Balanidæ,' p. 354.)
Valvæ testæ in unam confluentes; basis cyathiformis aut subcylindrica,
coraliis affixa.
6. _Sub-Genus_--CREUSIA, _Leach_. (Darwin, 'Balanidæ,' p. 375.)
Valvæ testæ 4, radiis instructæ; basis cyathiformis, coraliis affixa.
7. _Genus_--CHELONOBIA, _Leach_. (Darwin, 'Balanidæ,' p. 382.)
Valvæ testæ admodum crassæ, 6; sed ex iis, valva rostralis intùs è
tribus valvis rudimentariis conjunctis constat; basis membranacea;
scuta angusta, tergis cristâ articulari corneâ conjuncta.
[_Sectio_ ††.]
Scutum et tergum (ubi ambo adsunt) non inter se articulata; basis
membranacea; parietes sæpe profundè plicati, laminâ exteriore, ad basin
versâ, plerumque imperfectâ; branchiarum unaquæque e duobus plicis
constat. Testa vertebratis vivis affixa.
8. _Genus_--CORONULA, _Lamarck_. (Darwin, 'Balanidæ,' p. 397.)
Valvæ testæ 6; æquali latitudine; parietes tenues, profundè plicati,
plicis cavitates infrà solùm apertas efficientibus; valvæ operculares
orificio testæ multo minores. Cetaceis affixa.
9. _Genus_--PLATYLEPAS, _J. E. Gray_. (Darwin, 'Balanidæ,' p. 424.)
Valvæ testæ 6; unaquæque bilobata et intùs producta, ita ut 6 medias
costas longitudinales efficiant, quæ basin membranaceam extrorsùs
convexam sustinent.
10. _Genus_--TUBICINELLA, _Lamarck_. (Darwin, 'Balanidæ,' p. 430.)
Valvæ testæ 6, æquali latitudine; testa subcylindrica, orificio latiore
quam basis, pluribus cristis transversis virgata. Cetaceis affixa.
11. _Genus_--XENOBALANUS, _Steenstrup_. (Darwin, 'Balanidæ,' p. 438.)
Testa pæne rudimentaria, stelliformis, è valvis 6 formata, è quarum
medio corpus longum pedunculiforme exoritur; valvæ operculares absunt.
Cetaceis affixus.
Sub-Familia 2. CHTHAMALINÆ. (Darwin, 'Balanidæ,' p. 446.)
Rostrum cum alis sed sine radiis; valvæ rostro-laterales utrinque sine
alis; parietes non porosi.
12. _Genus_--CHTHAMALUS, _Ranzani_. (Darwin, 'Balanidæ,' p. 447.)
Valvæ testæ 6; basis membranacea, sed interdum ad speciem calcarea,
ideò quod parietes inflectuntur.
13. _Genus_--CHAMÆSIPHO, _Darwin_. ('Balanidæ' p. 470.)
Valvæ testæ 4, suturis sæpe admodùm obliteratis; basis membranacea.
14. _Genus_--PACHYLASMA, _Darwin_. ('Balanidæ,' p. 475.)
Valvæ testæ, conchâ recenter natâ, 8; adultâ aut 6, aut ad speciem 4,
ideò quod valvæ laterales arctè conjunguntur; basis calcarea.
15. _Genus_--OCTOMERIS, _G. B. Sowerby_. (Darwin, 'Balanidæ,' p. 482.)
Valvæ testæ 8; radii marginibus crenatis; basis membranacea.
16. _Genus_--CATOPHRAGMUS, _G. B. Sowerby_. (Darwin, 'Balanidæ,' p.
485.)
Valvæ testæ interiores 8, cum pluribus exterioribus parvarum
supplementalium valvarum verticillis; basis aut membranacea aut
calcarea.
Familia 2. VERRUCIDÆ. (Darwin, 'Balanidæ,' p. 495.)
Cirripedia sine pedunculo; scuta et terga, musculis depressoribus non
instructa, ex uno latere tantum mobilia, ex altero cum carinâ et rostro
in testam asymmetricam immobiliter conjuncta.
1. _Genus_--VERRUCA, _Schumacher_. (Darwin, 'Balanidæ,' p. 496.)
Familia 3. LEPADIDÆ. (Darwin, 'Lepadidæ,' p. 8, et 'Balanidæ,' p. 526.)
Cirripedia pedunculo flexili, musculis instructo; scuta et terga (si
qua adsunt) musculis depressoribus non instructa; reliquæ valvæ (si quæ
adsunt) in annulum immobilem non conjunctæ.
1. _Genus_--LEPAS, _Linn._ (Darwin, 'Lepadidæ,' p. 67.)
Valvæ 5, approximatæ; carina sursùm inter terga extensa, deorsùm
aut furcâ infossâ aut disco externo terminata; scuta subtriangula,
umbonibus ad angulum rostralem positis.
2. _Genus_--PŒCILASMA, _Darwin_. ('Lepadidæ,' p. 99.)
Valvæ 3, 5, aut 7, approximatæ; carina solùm ad basales apices
tergorum extensa, termino basali aut truncato aut in discum profunde
infossum producto; scuta pænè ovalia, umbonibus ad angulum rostralem
positis.
3. _Genus_--DICHELASPIS, _Darwin_. ('Lepadidæ,' p. 115.)
Valvæ 5, quæ ferè pro septem haberi possent, scuto in segmenta planè
duo, ad angulum autem rostralem conjuncta, diviso; carina plerumque
sursum inter terga extensa, deorsum aut disco infosso aut furcâ aut
calyce terminata.
4. _Genus_--OXYNASPIS, _Darwin_. ('Lepadidæ,' p. 133.)
Valvæ 5, approximatæ; scutorum umbones in medio marginis occludentis
positi; carina rectangulè flexa, sursùm inter terga extensa, termino
basali simpliciter concavo.
5. _Genus_--CONCHODERMA, _Olfers_. (Darwin, 'Lepadidæ,' p. 136.)
Valvæ 2 ad 5, minutæ, inter se remotæ; scuta bi- aut tri-lobata,
umbonibus in medio marginis occludentis positis; carina arcuata,
terminis utrinque pæne similibus.
6. _Genus_--ALEPAS, _S. Rang_. (Darwin, 'Lepadidæ,' p. 156.)
Capitulum aut sine valvis, aut scutis corneis, pæne abditis.
7. _Genus_--ANELASMA, _Darwin_. ('Lepadidæ,' p. 169.)
Capitulum sine valvis; aperturâ amplâ; pedunculus fimbriatus,
sub-globosus, infossus.
8. _Genus_--ALCIPPE, _Hancock_. (Darwin, 'Balanidæ,' p. 529.)
(Fœm.) Capitulum sine valvis, aperturâ spinosâ; pedunculus ad
basalem extremitatem crescit; superficie rostrali depressâ et disco
corneo tectâ; capitulum et pedunculus in cavitate conduntur, ab ipso
cirripedio formatâ.
9. _Genus_--IBLA, _Leach_. (Darwin, 'Lepadidæ,' p. 180.)
(Herm. et Fœm.) Valvæ 4, corneæ; pedunculus spinis corneis,
persistentibus vestitus.
10. _Genus_--SCALPELLUM, _Leach_. (Darwin, 'Lepadidæ,' p. 215, et
'Lepad. Foss.' p. 13.)
(Herm. et Fœm.) Valvis 12 ad 15; lateribus verticilli inferioris
quatuor vel sex, lineis incrementi plerumque convergentibus;
sub-rostrum rarissime adest; pedunculo squamifero, rarissime nudo.
CHARACTERES VALVARUM IN SPECIEBUS FOSSILIBUS.
Carina angusta, introrsùm arcuata, ab apice ad marginem basalem
paululum dilatata; carinæ parietes valde inflexi, costis manifestis
a tecto plerumque disjuncti; in multis speciebus intra-parietibus
instructa; intraparietes nonnunquam supernè producti ultra umbonem
carinæ, qui fit inde subcentralis; carinæ parietum lineæ incrementi
perobliquæ. Scuta plerumque subconvexa et tenuia, trapezoidea;
marginibus tergalibus lateralibusque angulo insigni disjunctis.
11. _Genus_--POLLICIPES, _Leach_. (Darwin, 'Lepadidæ,' p. 293, et
'Lepad. Foss.' p. 42.)
Valvæ ab 18 usque ad 100 et amplius; lateribus verticilli inferioris
multis; lineis incrementi deorsùm ordinatis; subrostrum semper adest;
pedunculus squamiferus.
CHARACTERES VALVARUM IN SPECIEBUS FOSSILIBUS.
Carina ab apice ad marginem basalem multum dilatata, apice plerumque
liberè prominente; carinæ parietes à tecto non distincté separati,
lineis incrementi parietum parum obliquis. Scuta plerumque subsolida,
convexa, subtrigonalia, margine tergo-laterali plus minusve eminente,
sed non angulo in margines duos discreto.
12. _Genus_--LITHOTRYA, _G. B. Sowerby_. (Darwin, 'Lepadidæ,' p. 332.)
Valvæ 8, si inter eas parvum (sæpe rudimentale) rostrum et duo parva
latera numerentur; incrementi lineis concinnè crenatis; pedunculus
squamis calcareis parvis vestitus, in verticillis superioribus
crenatis, aut calyci basali calcareo aut discorum ordini affixus.
13. _Genus_--LORICULA, _G. B. Sowerby, jun._ (Darwin, 'Lepad. Foss.' p.
81.)
Capitulo decem (fortasse) valvis instructo. Pedunculo seriebus decem
squamarum lævium calcarearum instructo; sex lateralibus multum
transversè elongatis; quatuor terminalibus angustis; secundùm pedunculi
margines rostralem et carinalem decurrit sutura medialis recta, squamis
non intersecantibus.
ORDO II. ABDOMINALIA. (Darwin, 'Balan.' p. 563.)
Cirripedia quibus carapax lageni-formis est; corpus ex 1 cephalico,
7 thoracicis, 3 abdominalibus segmentis constat; quorum abdominalia
tribus cirrorum paribus muniuntur; thoracica membris carent. Oris
labrum longè producitur et motus proprios efficit; œsophagi inferior
extremitas dentibus munitur. Larva, primò ovöides, sine externis
membris, sine oculo; postremò binocula, thoracicis cruribus nullis.
1. _Genus_--CRYPTOPHIALUS, _Darwin_. ('Balanidæ,' p. 566.)
ORDO III. APODA. (Darwin, 'Balan.' p. 587.)
Cirripedia quibus carapax ad duo separata fila (quæ defigendo
inserviunt) redactus est. Corpus ex 1 cephalico, 7 thoracicis, 3
abdominalibus segmentis constat, quæ omnia cirris carent. Os suctorium,
mandibuli et maxillæ (dorsis inter se appositis) cucullo includuntur,
qui è labro et palpis confluentibus formatur. Metamorphoses incognitæ.
1. _Genus_--PROTEOLEPAS, _Darwin_. ('Balanidæ,' p. 589.)
SYNOPSIS
ET
INDEX SYSTEMATICUS SPECIERUM,
_Et recentium, et fossilium_.[157]
[157] In hâc portione Synopsis, Auctorum nomina referunt solummodo
ad species.
ORDO I. THORACICA.
Fam. BALANIDÆ.
Sub-Fam. BALANINÆ.
[_Sectio_ †.]
1. _Genus_--BALANUS.
_Sectio_ A.
Parietes et basis et radii poris perforati.
_Sectio_ B.
Parietes et basis interdum poris perforati; radii nunquam; testæ axis
rostro-carinalis elongatus; basis cymbiformis, Gorgoniis et Milleporis
affixa.
_Sectio_ C.
Parietes et basis semper poris perforati; radii nunquam.
_Sectio_ D.
Parietes semper poris perforati; basis et radii nunquam.
_Sectio_ E.
Basis membranacea.
_Sectio_ F.
Parietes et radii nunquam, basis interdum, poris perforata; basis
interdum admodum tenuis, adeo ut vix distingui possit.
[_Sectio_ A.]
1. _Balanus tintinnabulum_, Linn. (Darwin, 'Balanidæ,' p. 194, Tab. 1
et 2, fig. 1.)
B. testâ à roseâ ad atropurpuream variante, sæpe longitudinaliter
virgatâ et costatâ. Orificio plerumque integro, interdum dentato. Scuti
cristâ articulari latâ et reflexâ. Tergi margine basali plerumque in
contrariis calcaris partibus rectam lineam formante.
_Hab._--In tepidis et torridis ubique maribus. _Foss._ in Europâ.
2. _Balanus tulipiformis_, Ellis. (Darwin, 'Balanidæ,' p. 204, Tab. 2,
fig. 2.)
B. testâ, obscurè roseâ, interdum purpurascente; orificio dentato.
Scuto externè admodum lævi membranâ tecto. Tergi musculorum depressorum
cristis distinctis.
_Hab._--In Europâ meridionali et Madeirâ.
3. _Balanus psittacus_, Molina. (Darwin, 'Balanidæ,' p. 206, Tab. 2,
fig. 3.)
B. testâ pallidè roseâ, sordidâ; orificio hexagonali. Scuti cristâ
articulari minimâ, quæ cum adductoris cristâ, admodum prominente,
confluit; musculo depressore laterali tubulari cavitate locato, quæ
sursùm ad valvæ apicem extendit. Tergi apice producto, aculeato,
purpureo; calcare ab angulo basi-scutali propiùs quam suâ ipsius
latitudine distante.
_Hab._ et _foss._ in Americâ meridionali.
4. _Balanus Capensis_, Ellis. (Darwin, 'Balanidæ,' p. 209, Tab. 2, fig.
4.)
B. testâ colore lucidè-roseo, umbratâ et sæpe longitudinaliter virgatâ.
Scuto ut in B. psittaco. Tergi apice producto aculeato, albo; calcare
ab angulo basi-scutali suâ ipsius latitudine distante.
_Hab._--In Africâ meridionali.
5. _Balanus nigrescens_, Lamarck. (Darwin, 'Balanidæ,' p. 210, Tab. 2,
fig. 5.)
B. testâ cinereâ, pallidè-cæruleo aut atro-cæruleo, aut albo tinctâ.
Scuti cristâ articulari parvâ, deorsùm in parvum, acutum aculeum
desinente; adductoris cristâ prominente. Tergi apice producto, aculeato.
_Hab._--In Australiâ.
6. _Balanus decorus_, Darwin. ('Balanidæ,' p. 212, Tab. 2, fig. 6.)
B. parietibus pallidè-roseis: radiis aliquanto intensioribus. Scuti
cristâ articulari parvâ. Tergi sulco longitudinali admodum tenui et
aperto; marginibus basalibus utrinque ad calcar declivibus.
_Hab._--In Novâ Zealandiâ.
7. _Balanus vinaceus_, Darwin. ('Balanidæ,' p. 213, Tab. 2, fig. 7.)
B. testâ fusco-purpureâ; parietum laminâ interiore cancellatâ. Scuto
longitudinaliter et tenuiter striato. Tergi sulco longitudinali tenui
et aperto: marginibus basalibus utrinque ad calcar declivibus.
_Hab._--In occidentali littore Americæ merid.
8. _Balanus Ajax_, Darwin. ('Balanidæ,' p. 214, Tab. 3, fig. 1.)
B. testâ globoso-conicâ, sæpe in axe rostro-carinali elongatâ,
pallidè-roseâ, lævi, admodum crassâ: tubis parietalibus prope marginem
basalem, cylindricis et minimis. Scuti cristâ articulari latâ, reflexâ.
_Hab._--In Arch. Philippino.
[_Sectio_ B.]
9. _Balanus stultus_, Darwin. ('Balanidæ,' p. 216, Tab. 3, fig. 2.)
B. parietibus et basi porosis: testâ albâ aut purpurâ leviter tinctâ.
Scuti margine basali in medio prominente. Tergi sulco longitudinali
supernè clauso; calcare angulo basi-scutali non approximato.
_Hab._--In Indiâ occident. et orient.
10. _Balanus calceolus_, Pallas. (Darwin, 'Balanidæ,' p. 218, Tab. 3,
fig. 3.)
B. parietibus et basi porosis. Scuto musculi depressoris lateralis
fossâ parvâ, profundâ.
_Hab._--In Africâ occidentali et Indiâ. _Foss._ in Angliâ.
11. _Balanus galeatus_, Linn. (Darwin, 'Balanidæ,' p. 220, Tab. 3, fig.
4.)
B. parietibus non porosis: basi porosa. Tergi apice, propter cristæ
articularis magnitudinem, quadrato.
_Hab._--In Americâ sept. et Indiâ occident.
12. _Balanus cymbiformis_, Darwin. ('Balanidæ,' p. 221, Tab. 3, fig. 5.)
B. parietibus et basi non porosis. Scuti et tergi cristis articularibus
minimis. Tergo lato, pæne æquilaterali.
_Hab._--In Indiâ.
13. _Balanus navicula_, Darwin. ('Balanidæ,' p. 221, Tab. 3, fig. 6.)
B. parietibus et basi non porosis: valvis testæ carino-lateralibus
admodùm angustis, latitudine à vertice ad imum pæne æquâ. Radiorum
marginibus suturalibus lævibus. Scuto externè longitudinaliter striato.
_Hab._--In Indiâ.
[_Sectio_ C.]
14. _Balanus trigonus_, Darwin. ('Balanidæ,' p. 223, Tab. 3, fig. 7.)
B. parietibus costatis, purpureo-rubris, maculatis: orificio
lato, trigonali, vix dentato. Scuto crasso, fossularum seriebus
longitudinalibus 1 ad 6. Tergo sine sulco longitudinali; calcare
truncato ad plenùm 1/3 valvæ latitudine.
_Hab._--In Arch. Indiæ Orient.: Australiâ: Californiâ: Peruviâ.
15. _Balanus spongicola_, Brown. (Darwin, 'Balanidæ,' p. 225, Tab. 4,
fig. 1.)
B. parietibus plerumque lævibus, interdum longitudinaliter plicatis;
roseis; orificio dentato: scuto longitudinaliter striato: tergum,
apice producto, sine sulco longitudinali; calcare truncato 1/3 valvæ
latitudine.
_Hab._--In Europâ meridionali et mediâ, Africâ meridionali; _var._ in
Indiâ occidentali. _Foss._ in Angliâ, in "Miocenâ Formatione."
16. _Balanus lævis_, Bruguière. (Darwin, 'Balanidæ,' p. 227, fig. 2.)
B. testâ aut fuscâ membranâ testâ, aut nudâ et albâ, aut
pallide-purpureâ: orificio parvo: radiis minimis: scuto 1 aut 2 sulcis
longitudinalibus profundis.
_Hab._--In littore occidentali Americæ utriusque.
17. _Balanus perforatus_, Bruguière. (Darwin, 'Balanidæ,' p. 231, Tab.
4, fig. 3, et Tab. 5, fig. 1.)
B. testâ sordidâ, pallidè-purpureâ aut albâ aut cinereâ, lævi aut
propter corrosionem longitudinaliter tenuiter costatâ; vaginâ purpureâ;
orificio plerumque parvo: radiis plerumque angustis aut nullis: scuto
introrsus cristâ brevi minutâ sub cristam adductoris prominentem
proximè et parallelè positâ: tergi apice aliquantum producto.
_Hab._--In Europâ meridionali et mediâ; in Africâ occidentali.
18. _Balanus concavus_, Bronn. (Darwin, 'Balanidæ' p. 235, Tab. 4, fig.
4.)
B. testâ, albo cum roseo aut obscurè purpureo longitudinaliter striatâ;
interdum pure albâ: scuto longitudinaliter tenuiter striato; internè
adductoris cristâ admodum aut modicè prominente.
_Hab._--In Peruviâ, Californiâ, Arch. Philippino, Australiâ. Et _foss._
in Europâ, Americâ septent. et meridionali.
19. _Balanus amphitrite_, Darwin. ('Balanidæ,' p. 240, Tab. 5, fig. 2.)
B. testâ purpureo aut roseo longitudinaliter striatâ; striis interdum
confluentibus; interdum pure albâ: scutum internè adductoris cristâ
prominente.
_Hab._--In tepidis et torridis ubique maribus.
20. _Balanus pœcilus_, Darwin. ('Balanidæ,' p. 246, Tab. 5, fig. 3.)
B. testâ obscurè rubrâ albo maculatâ scuto internè adductoris cristâ
nullâ tergi calcare præcisè truncato, ferè 1/3 valvæ latitudine.
_Hab._--In littore occident. Americæ merid.
21. _Balanus eburneus_, A. Gould. (Darwin, 'Balanidæ,' p. 248, Tab. 5,
fig. 4.)
B. testâ flavescente albâ: scuto longitudinaliter tenuiter striato:
tergi calcare truncato, margine basi-carinali ferè admodùm excavato;
margine carinali supernè prominente.
_Hab._--In Americâ septent. et Indiâ occident.
22. _Balanus improvisus_, Darwin. ('Balanidæ,' p. 250, Tab. 6, fig. 1.)
B. testâ albâ: radiis angustis, marginibus superioribus lævibus,
leniter arcuatis, admodùm obliquis: tergi sulco longitudinali;
calcaris, termino rotundato.
_Hab._--In Europâ, et Americæ utriusque littoribus orient. et occident.
23. _Balanus nubilus_, Darwin. ('Balanidæ,' p. 253, Tab. 6, fig. 2.)
B. testâ albâ, irregulari: basi alicubi imperfecte porosâ: scuti cristâ
articulari minutâ; adductoris cristâ prominente, fossam profundam
musculo depressori præbente: tergo introrsus purpureâ maculâ majore
notato; apice producto, purpureo.
_Hab._--In Californiâ.
24. _Balanus corrugatus_, Darwin. ('Balanidæ,' p. 254, Tab. 6, fig. 3.)
B. testâ albâ longitudinaliter plicatâ: radiis angustis: scuto internè
sine adductoris cristâ.
_Foss._ in Italiâ.
[_Sectio_ D.]
25. _Balanus porcatus_, Da Costa. (Darwin, 'Balanidæ,' p. 256, Tab. 6,
fig. 4.)
B. testâ albâ, plerumque longitudinaliter acutè costatâ: radiorum
marginibus superioribus pæne basi parallelis: scuto longitudinaliter
striato; tergi apice producto, purpureo.
_Hab._ et _foss._ in Europâ, et Americâ septent. et regionibus Arcticis.
26. _Balanus patellaris_, Spengler. (Darwin, 'Balanidæ,' p. 259, Tab.
6, fig. 5.)
B. testâ depressâ fuscâ, plerumque obscuro-violaceo longitudinaliter
striatâ: radiorum marginibus superioribus (in adultis speciminibus)
rotundatis, superficie sulcis tenuibus basi parallelis: basi interdùm
poris imperfectis perforatâ: scuto internè cum adductoris cristâ.
_Hab._--In Indiâ et Arch. Philippino.
27. _Balanus crenatus_, Bruguière. (Darwin, 'Balanidæ,' p. 261, Tab. 6,
fig. 6.)
B. testâ albâ: radiorum marginibus superioribus obliquis, asperis,
rectis: scuto sine adductoris cristâ: tergi calcare rotundato.
_Hab._ et _foss._ in Europâ, et Americâ septent. et regionibus
Arcticis, et Indiâ occident. et Africâ meridionali.
28. _Balanus glandula_, Darwin. ('Balanidæ,' p. 265, Tab. 7, fig. 1.)
B. testâ albâ; parietum laminâ internâ longitudinaliter et fortiter
costatâ, poris imperfectis et minutis, interdùm alicubi nullis: radiis
angustis marginibus superioribus rotundatis: scuto cum adductoris
cristâ: tergi calcare truncato, rotundato.
_Hab._--In Californiâ, et Oceano Pacifico meridion.
[_Sectio_ E.]
29. _Balanus balanoides_, Linn. (Darwin, 'Balanidæ,' p. 267, Tab. 7,
fig. 2.)
B. parietibus aut solidis, aut cancellatis, aut (rarò) unicâ pororum
serie formatis: tergi calcare obtuso aut acuto.
_Hab._--In Europâ et Americâ septent. et regionibus Arcticis.
30. _Balanus cariosus_, Pallas. (Darwin, 'Balanidæ,' p. 273, Tab. 7,
fig. 3.)
B. parietibus crassis, pluribus seriebus pororum inæqualium formatis;
tergo angusto, apice rostrato, calcare acuminato.
_Hab._--In littore occident. Americæ septent. et Freto Behringi et
Insulis Kuriliis.
31. _Balanus declivis_, Darwin. ('Balanidæ,' p. 275, Tab. 7, fig. 4.)
B. parietibus solidis: rostro, carinâ et valvis carino-lateralibus pæne
duplo longiore, basi igitur obliquâ: tergi calcare truncato, 1/2 valvæ
latitudine.
_Hab._--In Indiâ occident.
[_Sectio_ F.]
32. _Balanus Hameri_, Ascanius. (Darwin, 'Balanidæ,' p. 277, Tab. 7,
fig. 5.)
B. testâ albâ: radiorum marginibus superioribus obliquis, lævibus,
arcuatis; aciebus suturalibus lævibus: basi solidâ: scuto
longitudinaliter, debiliter striato: tergi calcare angusto.
_Hab._ et _foss._ in Europâ septent. et Americâ septent.
33. _Balanus amaryllis_, Darwin. ('Balanidæ,' p. 279, Tab. 7, fig. 6.)
B. testâ subroseo-purpureo striatâ, aut obnubilatâ; interdùm purè albâ:
radiis angustis, marginibus superioribus obliquis lævibus, arcuatis:
basi porosâ: scuto longitudinaliter planè striato: tergi calcare
angusto.
_Hab._--In Indiâ, et Arch. Indiæ orient. et Australiâ septent.
34. _Balanus allium_, Darwin. ('Balanidæ,' p. 281, Tab. 7, fig. 7.)
B. testâ pallidè purpuratâ, radiis latis, marginibus superioribus non
obliquis: basi porosâ: scuto lineis incrementi crenatis: tergi calcare
admodum brevi, truncato, 1/2 valvæ latitudine.
_Hab._--In Australiâ septent.
35. _Balanus cepa_, Darwin. ('Balanidæ,' p. 283, Tab. 7, fig. 8.)
B. testâ sordidè rubro-purpureâ, abruptè conicâ: radiis angustis:
basi obscurè porosâ: scuto lineis incrementi crenatis: tergi calcare
truncato, dimidiâ valvæ latitudine, et infrà angulum basi-scutalem
dependente, usque ad 1/2 sui ipsius latitudinem.
_Hab._--In Japaniâ.
36. _Balanus quadrivittatus_, Darwin. ('Balanidæ,' p. 284, Tab. 8, fig.
1.)
B. testâ abruptè conicâ, quatuor vittis longitudinalibus transversè
positis: radiorum marginibus superioribus obliquis: basi tenui, solidâ:
scuto, lineis incrementi lævibus; musculi depressoris lateralis fossâ
distinctâ nullâ: terga ut in _B. cepâ_.
_Hab._--In Archipel. Indiæ orient.
37. _Balanus terebratus_, Darwin. ('Balanidæ,' p. 285, Tab. 8, fig. 2.)
B. testâ albâ, costis longitudinalibus fortibus, margine basali in
longos aculeos producto: basi concavâ non porosâ, lineis radiantibus
fortiter costatâ; spatio intercostali foraminibus minutis rotundatis
sæpe duplici serie, cribrosè perforato.
_Hab._--(?)
38. _Balanus vestitus_, Darwin. ('Balanidæ,' p. 286, Tab. 8, fig. 3.)
B. testâ subroseo-purpureâ aut albâ, membranâ aurantiacâ vestitâ:
radiorum loco, meris fissuris: basi solidâ: scuto, cristâ adductoris
acutâ curvatâ; musculi depressoris cristæ adsunt: tergi calcare brevi,
truncato, 1/3 valvæ latitudine.
_Hab._--In Australiâ et Novâ Zealandiâ.
39. _Balanus imperator_, Darwin. ('Balanidæ,' p. 288, Tab. 8, fig. 4.)
B. testâ internè imperatorio-purpureâ: parietibus crassis margine
interno basali spinis et cristis irregularibus aspero: radiis angustis:
basi admodùm tenui, solidâ: scuto cum musculorum depressorum rostralium
et lateralium cristis: tergi calcare ad finem rotundato.
_Hab._--In Australiâ.
40. _Balanus flosculus_, Darwin. ('Balanidæ,' p. 290, Tab. 8, fig. 5.)
B. testâ purpureâ aut sordide-albâ, parietum margine basali spinis et
cristis irregularibus asperâ: radiis angustis aut nullis: basi quam
tenuissimâ, ad speciem nullâ: scuto cum musculi depressoris lateralis
cristis: tergo admodùm angusto, calcare acuminato.
_Hab._--In Americæ merid. littore occid.
41. _Balanus bisulcatus_, Darwin. ('Balanidæ,' p. 293, Tab. 8, fig. 6.)
B. radiorum marginibus superioribus obliquis, lævibus; aciebus
suturalibus lævibus: basi poris magnis perforatâ: scuto angusto,
sulcis longitudinalibus 2 ad 4: tergi calcare brevissimo dimidiâ valvæ
latitudine.
_Foss._ in Angliâ, et Galliâ et Belgio. "Miocenâ Formatione."
42. _Balanus dolosus_, Darwin. ('Balanidæ,' p. 295, Tab. 8, fig. 7.)
B. radiorum marginibus superioribus obliquis lævibus: aciebus
suturalibus item lævibus: basi poris magnis perforatâ: tergi calcare
non admodum brevi, 1/3 valvæ latitudine.
_Foss._ in Angliâ. "Pliocenâ, et miocenâ Formationibus."
43. _Balanus unguiformis_, J. de C. Sowerby. (Darwin, 'Balanidæ,' p.
296, Tab. 8, fig. 8.)
B. parietibus tenuibus, interdum poris perforatis: radiorum marginibus
superioribus obliquis; aciebus suturalibus tenuissimè crenatis: basi
solidâ: tergi calcare angusto, obtuso.
_Foss._ in Angliâ et Belgio. "Eocenâ Formatione."
44. _Balanus varians_, Darwin. ('Balanidæ,' p. 298, Tab. 8, fig. 9.)
B. parietibus modicè crassis; radiorum marginibus superioribus
perobliquis; aciebus suturalibus pæne lævibus aut tenuiter crenatis:
basi tenuiter porosâ: tergi calcare, parvo, angusto, obtuso.
_Foss._ in Patagoniâ.
45. _Balanus inclusus_, Darwin. ('Balanidæ,' p. 299, Tab. 8, fig. 10.)
B. testâ rufo-fuscâ: radiis latis, marginibus superioribus aut non
obliquis aut modicè; aciebus suturalibus cum septis planè denticulatis:
basi porosâ: scuto sine adductoris cristâ: tergi calcare subangusto.
_Foss._ in Angliâ et Germaniâ. "Miocenâ Formatione."
2. _Sub-Genus_--ACASTA.
1. _Acasta spongites_, Poli. (Darwin, 'Balanidæ,' p. 308, Tab. 9, fig.
1.)
A. parietibus carino-lateralibus ferè 1/6 parietum lateralium
latitudine: superficie internâ parietum plerumque debiliter costatâ:
scuti cristâ articulari ad extremitatem inferiorem abruptè præcisâ:
tergi calcare truncato rotundato, ferè 1/3 valvæ latitudine.
_Hab._--In Europâ et Africâ meridion.
2. _Acasta sulcata_, Lamarck. (Darwin, 'Balanidæ,' p. 310, Tab. 9, fig.
2.)
A. parietibus carino-lateralibus ferè 1/6 parietum lateralium
latitudine: parietum superficie internâ plerumque fortiter costatâ:
basis acie fortiter crenatâ: testæ orificio sub-exiguo: tergi calcare
plerumque truncato, pæne 1/2 valvæ latitudine.
_Hab._--In Australiâ.
3. _Acasta cyathus_, Darwin. ('Balanidæ,' p. 312, Tab. 9, fig. 3.)
A. parietibus carino-lateralibus ferè 1/4 parietum lateralium
latitudine: radiis latioribus quam parietes: basi pæne planâ, exiguâ:
tergi calcare truncato, 1/2 valvæ latitudine.
_Hab._--In Madeirâ et Indiâ occid.
4. _Acasta undulata_, Darwin. ('Balanidæ,' p. 313, Tab. 9, fig. 4.)
A. testâ, ad speciem, ut in "A. spongites," sed majore: scuto externe
striis longitudinalibus, sæpe binis, signato, sulcis intermediis
latioribus: tergi calcare, pæne 1/2 valvæ latitudine.
_Foss._ in Angliâ. "Miocenâ Formatione."
5. _Acasta glans_, Lamarck. (Darwin, 'Balanidæ,' p. 314, Tab. 9, fig.
5.)
A. parietibus internè per-lævibus, marginibus lateralibus uniuscujusque
testæ valvæ intùs prominentibus: basis acie rarò crenatâ, sed 6
dentibus introrsùs prominentibus instructâ: scuto longitudinaliter
fortiter striato.
_Hab._--In Australiâ.
6. _Acasta lævigata_, J. E. Gray. (Darwin, 'Balanidæ,' p. 315, Tab. 9,
fig. 6.)
A. parietibus intùs per-lævibus, marginibus lateralibus uniuscujusque
testæ valvæ internùs prominentibus: basis acie fortiter crenatâ, et 6
dentibus introrsùs prominentibus instructâ: scuto aut longitudinaliter
tenuiter striato aut lævi.
_Hab._--In Mari Rubro et Archipel. Philippino.
7. _Acasta fenestrata_, Darwin. ('Balanidæ,' p. 316, Tab. 9, fig. 7.)
A. testâ subrufâ, foraminibus magnis membranâ-tectis inter suturas
suprà basin proximis, 6; parietibus carino-lateralibus 1/2 parietum
lateralium latitudine: intùs, parietibus et basis acie lævibus: tergi
cristâ articulari brevi prominente; calcare acuminato.
_Hab._--In Archipel. Philippino.
8. _Acasta purpurata_, Darwin. ('Balanidæ,' p. 318, Tab. 9, fig. 8.)
A. testâ obscurè cæruleo-purpureâ, foraminibus parvis, membrana-tectis
inter suturas suprà basin proximis, 6: tergi cristâ articulari
per-brevi prominente; calcare admodum lato rotundato.
_Hab._--In Sumatrâ et Archipel. Philippino.
9. _Acasta sporillus_, Darwin. ('Balanidæ,' p. 319, Tab. 9, fig. 9.)
A. testâ purpureo-fuscâ, parietibus intùs fortiter costatis et
reticulatis: testæ valvis carino-lateralibus angustissimis, ad basin
non extendentibus.
_Hab._--In Sooloo Insulis, in Archipelago Indico orient.
3. _Genus_--TETRACLITA.
1. _Tetraclita porosa_, Linn. (Darwin, 'Balanidæ,' p. 328, Tab. 10,
fig. 1.)
T. radiis (si qui rarò adsunt) angustis; etiam suturis sæpe nullis:
testâ abruptè conicâ, superficie plerumque corrosâ, stalactitiferam
speciem exhibente.
_Hab._--In torridis et tepidis ubique maribus.
2. _Tetraclita serrata_, Darwin. ('Balanidæ,' p. 333, Tab. 10, fig. 2.)
T. testâ obscurè viridi-griseâ, costis longitudinalibus serratis
angustis: radiis nullis: scuti cristis (articulari et adductoris)
cavitatem usque ad apicem valvæ extendentem formantibus.
_Hab._--In Africâ merid.
3. _Tetraclita rosea_, Krauss. (Darwin, 'Balanidæ,' p. 335, Tab. 10,
fig. 3.)
T. testâ sordidè albâ, subroseo tinctâ; parietibus è serie unicâ
magnarum tubarum formatis: radiis plerumque angustis: tergi calcare
sub-brevi, lato.
_Hab._--In Australiâ, et Africâ meridion.
4. _Tetraclita purpurascens_, W. Wood. (Darwin, 'Balanidæ,' p. 337,
Tab. 11, fig. 1.)
T. testâ depressâ, pallidè purpureâ aut sordidè albâ, superficie
longitudinaliter costatâ, aut corrosâ et granulatâ: radiis, aut etiam
suturis, nullis, aut radiis planè formatis et latis, marginibus
superioribus basi parallelibus: basi membranaceâ: scuto transversè
elongato: tergo exiguo, calcare per-brevi, rotundato.
_Hab._--In Australiâ.
5. _Tetraclita costata_, Darwin. ('Balanidæ,' p. 339, Tab. 11, fig. 2.)
T. testâ depressâ sub-albâ, plerumque costis longitudinalibus admodùm
prominentibus, 10: radiis latis, marginibus superioribus basi
parallelis: basi calcareâ: scuto externè longitudinaliter striato:
tergi calcare brevi rotundato.
_Hab._--In Archipel. Philippino.
6. _Tetraclita vitiata_, Darwin. ('Balanidæ,' p. 340, Tab. 11, fig. 3.)
T. testâ albâ, parte superiore plerumque roseo tinctâ; superficie
irregulari; tubis parietalibus admodum irregularibus; radiis modicè
latis, marginibus superioribus obliquis; alarum aciebus suturalibus
percrassis crenatis: tergi calcare cum angulo basi-scutali non juncto;
calcaris termino æquabiliter rotundato.
_Hab._--In Australiâ septentr. et orient. et Archipel. Philippino.
7. _Tetraclita cœrulescens_, Spengler. (Darwin, 'Balanidæ,' p. 342,
Tab. 11, fig. 4.)
T. testæ parte superiore viridi-cæruleo tinctâ, longitudinaliter
costatâ; radiis modice latis, marginibus superioribus obliquis: scuto
adductoris cristâ exiguâ, articulari admodùm prominente; ambabus ita
conjunctis ut cavitatem parvam subcylindricam efficiant: tergi calcare
cum angulo basi-scutali non juncto.
_Hab._--In Archipel. Philippino et Oceano Pacifico.
8. _Tetraclita radiata_, De Blainville. (Darwin, 'Balanidæ,' p. 343,
Tab. 11, fig. 5.)
T. testâ albâ costis longitudinalibus approximatis multis; radiis
latis, marginibus superioribus leviter obliquis, intus porosis:
tergi cristâ articulari suprà modum prominente; calcare cum angulo
basi-scutali non juncto.
_Hab._--In Indiâ occid. et Australiâ: sæpe navigiis suffixa.
4. _Genus_--ELMINIUS.
1. _Elminius Kingii_, J. E. Gray. (Darwin, 'Balanidæ,' p. 348, Tab. 11,
fig. 6.)
E. testâ lævi; griseâ aut sordidè albâ; radiis latis, marginibus et
aciebus lævibus: scuto, sine adductoris cristâ: tergi calcare ab angulo
basi-scutali distincto: scuto et tergo interdum inter se calcifactis.
_Hab._--In Americâ merid.
2. _Elminius modestus_, Darwin. ('Balanidæ,' p. 350, Tab. 12, fig. 1.)
E. testâ longitudinaliter plicatâ, subviridi aut albâ; radiis modicè
latis, marginibus et aciebus lævibus: scuto sine adductoris cristâ:
tergo angusto, calcare cum angulo basi-scutali confluente.
_Hab._--In Australiâ.
3. _Elminius plicatus_, J. E. Gray. (Darwin, 'Balanidæ,' p. 351, Tab.
12, fig. 2.)
E. testâ longitudinaliter profundè plicatâ, corrosâ, alicubi
aurantiacâ; radiis per-angustis, aciebus sinuosis, leviter dentatis:
scuto cum adductoris cristâ.
_Hab._--In Novâ Zealandiâ.
4. _Elminius simplex_, Darwin. ('Balanidæ,' p. 353, Tab. 12, fig. 3.)
E. testâ longitudinaliter costatâ, sordidè albâ; radiis angustissimis,
marginibus et aciebus lævibus: scuto cum adductoris cristâ.
_Hab._--In Australiâ.
5. _Genus_--PYRGOMA.
1. _Pyrgoma Anglicum_, Leach. (Darwin, 'Balanidæ,' p. 360, Tab. 12,
fig. 4.)
P. testâ abruptè conicâ, purpureo-rubrâ; orificio ovato, angusto; basi
porosâ, plerumque è coralio exsertâ: scuto et tergo subtriangularibus.
_Hab._--In Magnâ Britanniâ et Europâ meridionali, et Insulis "Cape de
Verde."
2. _Pyrgoma Stokesii_, Darwin. ('Balanidæ,' p. 361, Tab. 12, fig. 6.)
P. testâ modicè conicâ, pallidè purpureo-rubrâ; orificio ovato;
basi non-porosâ, in coralium profunde conditâ: scuto et tergo
subtriangularibus.
_Hab._--In Indiâ occid.
3. _Pyrgoma cancellatum_, Leach. (Darwin, 'Balanidæ,' p. 362, Tab. 12,
fig. 5.)
P. testâ circumferentiâ plerumque lobatâ: scuto elongato, adductoris
cristâ longè infrà marginem basalem descendente, et ad rostralem finem
in acumen quadratum producto: tergi calcare valvæ partem superiorem
longitudine quadruplo superante.
_Hab._--In Archip. Indico. orient. (?)
4. _Pyrgoma conjugatum_, Darwin. ('Balanidæ,' p. 364, Tab. 12, fig. 7.)
P. testâ pæne planâ, cristis approximatis radiantibus: scuto et tergo
inter se sine suturâ calcifactis; scuto, adductoris cristâ infrà
marginem basalem descendente, et ad terminum rostralem in acumen
producto; tergi calcare superioris valvæ partis longitudinem ferè
æquante.
_Hab._--In Mari Rubro.
5. _Pyrgoma grande_, G. B. Sowerby, jun. (Darwin, 'Balanidæ,' p. 365,
Tab. 13, fig. 1.)
P. testâ modicè convexâ, pæne lævi: scuto et tergo inter se sine suturâ
calcifactis; scuto limbo occludente parvo instructo, adductoris cristâ
infrà marginem basalem descendente: tergo quadrato, calcare nullo.
_Hab._--In Archip. Indico orient.
6. _Pyrgoma milleporæ_, Darwin. ('Balanidæ,' p. 367, Tab. 13, fig. 2.)
P. testæ orificio angustè-ovato; vaginâ obscurè purpureâ: scuto multùm
elongato: tergo triangulo, convexo, calcare nullo.
_Hab._--In Archip. Philippino.
7. _Pyrgoma dentatum_, Darwin. ('Balanidæ,' p. 369, Tab. 13, fig. 3.)
P. scuto multùm elongato, cum prominentiâ articulari dentiformi: tergo
convexo, irregulariter triangulo, interdum cum calcare imperfecto, et
in superficie internâ dente introrsùs prominente; scuto et tergo limbo
occludente instructis.
_Hab._--In Mari Rubro.
8. _Pyrgoma crenatum_, G. B. Sowerby. (Darwin, 'Balanidæ,' p. 370. Tab.
13, fig. 4.)
P. scuto multùm elongato, adductoris cristâ infra marginem basalem
reflexum descendente: tergi calcare lato depresso; scuto et tergo limbo
occludente instructis.
_Hab._--In Archip. Ind. orient et Philippino.
9. _Pyrgoma monticulariæ_, J. E. Gray. (Darwin, 'Balanidæ,' p. 372.
Tab. 13, fig. 5.)
P. testâ irregulari, margine exteriore aspero; orificio minuto rotundo:
scuto et tergo multum elongatis, inter se sine suturâ calcifactis,
limbo occludente lato instructis.
_Hab._--In Archip. Indico orient.
6. _Sub-Genus_--CREUSIA.
1. _Creusia spinulosa_, Leach, (Darwin, 'Balanidæ,' p. 376, Tab. 13,
fig. 6, et Tab. 14, fig. 6.)
_Hab._--In Maribus torridis.
7. _Genus_--CHELONOBIA.
1. _Chelonobia testudinaria_, Linn. (Darwin, 'Balanidæ,' p. 392, Tab.
14, fig. 1 et fig. 5, Tab. 15, fig. 1).
Ch. testâ conicâ, depressâ, gravi; radiis subangustis, depressis
plerumque utrinque scrobiculatis.
_Hab._--In tepidis et torridis ubique Maribus; testudinibus affixa.
2. _Chelonobia caretta_, Spengler. (Darwin, 'Balanidæ,' p. 394, Tab.
14, fig. 2.)
Ch. testâ globoso-convexâ, crassissimâ, gravissimâ; parte superiore
tritâ, substriatâ; radiis aut non formatis aut admodum angustis;
parietibus sine cavitatibus sursùm inter septa interrupta radiantia
extendentibus.
_Hab._--In Africâ occid. et Australiâ septent.; testudinibus affixa.
3. _Chelonobia patula_, Ranzani. (Darwin, 'Balanidæ,' p. 396, Tab. 14,
fig. 3).
Ch. testâ abruptè conicâ, admodum lævi et tenui; orificio magno, ferè
dimidium diametri basalis testæ superante; radiis latis, lævibus,
leviter depressis.
_Hab._--Mediterraneo; Africâ occid.; Indiâ occid.; Australiâ Crustaceis
et molluscarum testis lævibus affixa.
[_Sectio_ ††.]
8. _Genus_--CORONULA.
1. _Coronula balænaris_, Linn. (Darwin, 'Balanidæ,' p. 415, Tab. 15,
fig. 2, Tab. 16, fig. 3 et 5.)
B. testâ multùm depressâ, costis longitudinalibus planatis, aciebus
earum simplicibus; orificio hexagono-rotundato; radiis testæ ipsius
crassitudinem pænè æquantibus; valvis opercularibus, 4.
_Hab._--In Oceano meridionali.
2. _Coronula diadema_, Linn. (Darwin, 'Balanidæ,' p. 417, Tab. 15, fig.
3.)
C. testâ coroniformi, costis longitudinalibus convexis, aciebus earum
crenatis; orificio hexagono; radiis modicè crassis, admodum latis;
tergis nullis aut rudimentariis.
_Hab._--In Oceano septentrionali.
3. _Coronula reginæ_, Darwin. ('Balanidæ,' p. 419, Tab. 15, fig. 5.)
C. testâ globoso-conicâ aut depressâ, costis longitudinalibus multùm
planatis, aciebus earum crenatis, superficie striatâ, granulatâ;
orificio hexagono; radiis tenuibus 1/5 testæ valvæ crassitudinem non
superantibus; tergis nullis.
_Hab._--In Oceano Pacifico.
4. _Coronula barbara_, Darwin. ('Balanidæ,' p. 421, Tab. 15, fig. 6).
C. testâ (probabiliter) coroniformi, costis longitudinalibus convexis,
aciebus earum crenatis, superficie internâ et externâ cristis
transversis asperâ; radiis modicè crassis; spatio inter radios et alas
solidè impleto.
_Foss._--In Angliâ, "Red Crag Formatione."
9. _Genus_--PLATYLEPAS.
1. _Platylepas bissexlobata_, De Blainville. (Darwin, 'Balanidæ,' p.
428, Tab. 17, fig. 1.)
P. testâ lineis transversis incrementi conspicuis; parietibus porosis;
vaginâ ad dimidium parietum vix descendente.
_Hab._--In Mediterraneo; Africâ occid.; Indiâ occid.; Australiâ
septentrionali.
2. _Platylepas decorata_, Darwin. ('Balanidæ,' p. 429, Tab. 17, fig. 2.)
P. testâ costis longitudinalibus tenuibus, parte earum inferiore
globulis minutis decoratâ; parietibus non porosis; membranâ basali pari
cum testâ convexitate.
_Hab._--In Oceano Pacifico.
10. _Genus_--TUBICINELLA.
1. _Tubicinella trachealis_, Shaw. (Darwin, 'Balanidæ,' p. 431, Tab. 17,
fig. 3).
_Hab._--In Oceano Pacifico meridionali.
11. _Genus_--XENOBALANUS.
1. _Xenobalanus globicipitis_, Steenstrup. (Darwin, 'Balanidæ,' p. 440,
Tab. 17, fig. 4.)
_Hab._--In Oceano Atlantico septentrionali.
Sub-Fam. CHTHAMALINÆ.
12. _Genus_--CHTHAMALUS.
1. _Chthamalus stellatus_, Poli. (Darwin, 'Balanidæ,' p. 455, Tab. 18,
fig. 1.)
Ch. testâ albâ aut griseâ, plerumque multùm corrosâ et punctâ; radiis
(si qui adsunt) angustis, aciebus suturalibus tenuissimè crenatis;
tergo, musculi depressoris cristis infrà marginem basalem vix
dependentibus.
_Hab._--In Europâ; Americâ sept.; Braziliâ; Mari Rubro; Chinâ;
Californiâ septent.
2. _Chthamalus antennatus_, Darwin. ('Balanidæ,' p. 460, Tab. 18, fig.
2.)
Ch. testâ conicâ, plerumque lævi; nisi profundè corrosâ, pallidè
sordidè carneâ; suturis semper distinctis; radiis, si qui adsunt,
aciebus suturalibus per-lævibus.
_Hab._--In Australiâ.
3. _Chthamalus cirratus_, Darwin. ('Balanidæ,' p. 461, Tab. 18, fig. 4.)
Ch. testâ albâ aut griseâ; vaginâ et valvis opercularibus membranâ
fimbriatâ plerumque vestitis; tergo, angulo basi-carinali dependente et
acuminato.
_Hab._--In Chiliâ, Peruviâ.
4. _Chthamalus fissus_, Darwin. ('Balanidæ,' p. 462, Tab. 18, fig. 6.)
Ch. testâ subfuscâ, plicatâ; orificio longitudine duplo majore quam
latitudo; tergo triangulo, æquilaterali, marginibus basalibus et
carinalibus leviter prominentibus.
_Hab._--In Californiâ.
5. _Chthamalus dentatus_, Krauss. (Darwin, 'Balanidæ,' p. 463, Tab. 18,
fig. 3).
Ch. testâ sordidè albâ, aut subfuscâ; suturis, dentibus inter se
concurrentibus formatis; tergi margine carinali prominente.
_Hab._--In Africâ.
6. _Chthamalus Hembeli_, Conrad. (Darwin, 'Balanidæ,' p. 465, Tab. 18,
fig. 5.)
Ch. testâ obscurè rubro-purpureâ; suturis aut obliteratis aut
dentibus inter se concurrentibus formatis; basi interdum per parietum
inflectionem calcareâ factâ; scuto, sulcis duobus vel tribus ad mediam
valvam deorsùm extendentibus.
_Hab._--In Californiâ.
7. _Chthamalus intertextus_, Darwin. ('Balanidæ,' p. 467, Tab. 19, fig.
1.)
Ch. testâ (si benè conservetur) violaceo-purpureâ; suturis aut
obliteratis aut ex interplicatis laminis obliquis formatis; basi
membranaceâ, sed cingente projecturâ ex parietum marginibus basalibus
inflexis formatâ: scuto et tergo inter se prorsus calcifactis.
_Hab._--In Archip. Philippino.
8. _Chthamalus scabrosus_, Darwin. ('Balanidæ,' p. 468, Tab. 19, fig.
2.)
Ch. testâ (si benè conservetur) obscurè purpureo-fuscâ; suturis ex
interplicatis laminis obliquis (sed raro benè) formatis; tergo, cum
musculi depressoris fossâ profundâ angustâ, ad angulum basi-carinalem.
_Hab._--In Americæ merid. littore merid. et occid.
13. _Genus_--CHAMÆSIPHO.
1. _Chamæsipho columna_, Spengler. (Darwin, 'Balanidæ,' p. 470, Tab.
19, fig. 3.)
Ch. suturis, nisi in recenter natis, plerumque et internè et externè
obliteratis; tergo, cum depressoris musculi fossis parvis.
_Hab._--In Australiâ.
2. _Chamæsipho scutelliformis_, Darwin. ('Balanidæ,' p. 472, Tab. 19,
fig. 4.)
Ch. rostro per-exiguo, elongato, triangulo: testæ valvarum lateralium
unâquæque cum foramine, et carinâ cum duobus similibus foraminibus; quæ
4 in columnas tubulares testaceas conducunt.
_Hab._--In Chinâ (?)
14. _Genus_--PACHYLASMA.
1. _Pachylasma giganteum_, Philippi. (Darwin, 'Balanidæ,' p. 477, Tab.
19, fig. 5.)
P. testâ et operculo sordide albis; testæ valvâ laterali et
carino-laterali alis similibus instructâ.
_Hab._--In Mari Mediterraneo.
2. _Pachylasma aurantiacum_, Darwin. ('Balanidæ,' p. 480, Tab. 20, fig.
1.)
P. testâ aurantiaco tinctâ; ad speciem externam, quasi è 4 solùm testæ
valvis formatâ, ideò quòd laterales, et carino-laterales valvæ utrinque
obscurâ fissurâ solùm separantur.
_Hab._--In Australiâ.
15. _Genus_--OCTOMERIS.
1. _Octomeris angulosa_, G. B. Sowerby. (Darwin, 'Balanidæ,' p. 483,
Tab. 20, fig. 2.)
O. testâ sordidè albâ, asperâ, solidâ; alis crassis, aciebus
suturalibus obtusè crenatis.
_Hab._--In Africâ merid.
2. _Octomeris brunnea_, Darwin. ('Balanidæ,' p. 484, Tab. 20, fig. 3.)
O. testâ rubro-fuscâ depressâ, tenui, longitudinaliter sulcis tenuibus
signatâ; tergi margine basali leviter in angulum flexâ.
_Hab._--In Archip. Philippino.
16. _Genus_--CATOPHRAGMUS.
1. _Catophragmus polymerus_, Darwin. ('Balanidæ,' p. 487, Tab. 20, fig.
4.)
C. basi membranaceâ: sine appendicibus caudalibus.
_Hab._--In Australiâ.
2. _Catophragmus imbricatus_, G. B. Sowerby. (Darwin, 'Balanidæ,' p.
490.)
C. basi calcareâ: cum appendicibus caudalibus.
_Hab._--In Indiâ occident.
2. Fam. VERRUCIDÆ.
1. _Genus_--VERRUCA.
1. _Verruca Strömia_, Müller. (Darwin, 'Balanidæ,' p. 518, Tab. 21,
fig. 1.)
V. scuto mobili, cristâ articulari inferiore dimidiam brevis cristæ
articularis superioris latitudinem non æquante: testâ plerumque
longitudinaliter sulcatâ.
_Hab._--In Europâ; Mari Rubro.
2. _Verruca lævigata_, G. B. Sowerby. (Darwin, 'Balanidæ,' p. 520, Tab.
21, fig. 3.)
V. scuto mobili, cristâ articulari inferiore latiore quam brevis crista
articularis superior; tergo mobili, latitudine majore quam altitudo,
cristâ articulari superiore in terminum subacutum producto.
_Hab._--In Americâ merid.
3. _Verruca Spengleri_, Darwin. ('Balanidæ,' p. 521, Tab. 21, fig. 2.)
Scuto mobili, cum adductoris musculi cristâ acutâ, rectâ, mediali;
scuto immobili non majore quam immobile tergum.
_Hab._--In Insulâ Madeirâ.
4. _Verruca nexa_, Darwin. ('Balanidæ,' p. 522, Tab. 21, fig. 5.)
V. testâ sub-rubrâ; scuto mobili cum cristis externis longitudinalibus
fortiter prominentibus, præter cristas articulares; scuto immobili
majore quam carina, adductoris musculi cristâ vel laminâ distinctâ
nullâ.
_Hab._--In Indiâ occid.
5. _Verruca prisca_, Bosquet. (Darwin, 'Balanidæ,' p. 525, Tab. 21,
fig. 4.)
V. testâ lævi; scuti mobilis cristâ articulari inferiore aliquanto
latiore quam superior.
_Foss._--In Cretaceâ Formatione.
3. Fam. LEPADIDÆ.
1. _Genus_--LEPAS.
1. _Lepas anatifera_, Linn. (Darwin, 'Lepadidæ,' p. 73, Tab. 1, fig. 1,
_var._)
L. valvis aut lævibus aut delicate striatis; è duobus scutis, dextro
solùm dente interno umbonali instructo; pedunculi parte superiore fuscâ.
_Hab._--In Oceano, pæne ubique.
2. _Lepas Hillii_, Leach. (Darwin, 'Lepadidæ,' p. 77, Tab. 1, fig. 2.)
L. valvis lævibus; scutorum dentibus internis umbonalibus nullis;
carinâ à cæteris valvis, furcâ etiam a scutorum basali margine,
paululum distante; pedunculi parte superiore aut pallidâ aut aurantiacâ.
_Hab._--In Oceano, pæne ubique.
3. _Lepas anserifera_, Linn. (Darwin, 'Lepadidæ,' p. 81, Tab. 1, fig.
4.)
L. valvis approximatis leviter sulcatis (tergis præcipuè); scuto dextro
dente forti interno umbonali, lævo aut dente exiguo, aut merâ cristâ
instructo; margine occludente arcuato, prominente; pedunculi parte
superiore aurantiacâ.
_Hab._--In Oceano, pæne ubique.
4. _Lepas pectinata_, Spengler. (Darwin, 'Lepadidæ,' p. 85, Tab. 1,
fig. 3.)
L. valvis tenuibus, crassè sulcatis, sæpe pectinatis; scutorum cristâ
prominente ab umbone ad apicem, juxta marginem occludentem, pertinente;
furcæ carinalis cruribus inter angulos 135° et 180° divergentibus.
_Hab._--In Oceano Atlantico, sept. et meridionali.
5. _Lepas australis_, Darwin. ('Lepadidæ,' p. 89, Tab. 1, fig. 5.)
L. valvis glabris, tenuibus, fragilibus; scutorum dentibus umbonalibus
utrinque internis; carinæ parte superiore latâ, planâ, suprà furcam
valdè constrictâ; furcæ cruribus latis, planis, tenuibus, acuminatis,
intermedio margine non reflexo.
_Hab._--In Oceano meridionali.
6. _Lepas fascicularis_, Ellis. (Darwin, 'Lepadidæ,' p. 92, Tab. 1,
fig. 6.)
L. valvis glabris, tenuibus, pellucidis; carinâ rectangulè flexâ, parte
inferiore in discum planum oblongum expansâ.
_Hab._--In Oceano torrido et tepido, pæne ubique.
2. _Genus_--PŒCILASMA.
1. _Pœcilasma Kæmpferi_, Darwin. ('Lepadidæ,' p. 102, Tab. 2, fig. 1.)
P. valvis 5; carinæ basi truncatâ et cristatâ: scutorum dentibus
internis umbonalibus fortibus: tergorum acumine basali truncato,
margini occludenti pæne parallelo.
_Hab._ in Japaniâ.
2. _Pœcilasma aurantia_, Darwin. ('Lepadidæ,' p. 105, Tab. 2, fig. 2.)
P. valvis 5; carinæ basi truncatâ: scutis ovatis, margine basali
per-brevi, dentibus parvis, internis, umbonalibus instructo: tergorum
acumine basali perobliquè truncato.
_Hab._--In Insulâ Madeirâ.
3. _Pœcilasma crassa_, J. E. Gray. (Darwin 'Lapadidæ,' p. 107, Tab. 2,
fig. 3.)
P. valvis 5; carinæ termino basali in discum parvum infossum producto:
scutis convexis, dentibus internis umbonalibus nullis: tergis pæne
rudimentalibus, vix carinâ latioribus.
_Hab._--In Insulâ Madeirâ.
4. _Pœcilasma fissa_, Darwin. ('Lepadidæ,' p. 109, Tab. 2, fig. 4.)
P. valvis 7; scuto utroque è duobus juxtapositis segmentis formato;
segmento altero intus dentato: tergis brevibus, ter aut quater carinâ
latioribus: carinæ termino basali in discum parvum angustum infossum
producto.
_Hab._--In Archipel. Philippino.
5. _Pœcilasma eburnea_, Hinds. (Darwin, 'Lepadidæ,' p. 112, Tab. 2,
fig. 5.)
P. valvis 3; scutis acuminatis, ovatis; ad pedunculum pæne transversè
spectantibus; dentibus internis umbonalibus fortibus: tergis nullis
carinæ termino basali in discum amplum oblongum infossum producto.
_Hab._--In Novâ Guineâ.
3. _Genus_--DICHELASPIS.
1. _Dichelaspis Warwickii_, J. E. Gray. (Darwin, 'Lepadidæ,' p. 120,
Tab. 2, fig. 6.)
D. scutorum segmento basali duplo latiore quam segmentum occludens:
tergorum parte inferiore paulò, latiore quam occludens scutorum
segmentum.
_Hab._--In Archipel. Indico orientali.
2. _Dichelaspis Grayii_, Darwin. ('Lepadidæ,' p. 123, Tab. 2, fig. 9.)
D. scutorum segmento basali angustiore quam segmentum occludens;
longitudine pæne dimidiâ: tergis bipenniformibus, margine crenato,
spinâ posticâ, manubrio angustiore quam occludens scutorum segmentum.
_Hab._--In Oceano torrido, aut Indico, aut Pacifico.
3. _Dichelaspis pellucida_, Darwin. ('Lepadidæ,' p. 125, Tab. 2, fig.
7.)
D. valvarum singularum acuminibus superioribus et inferioribus vix
intersecantibus: scutorum segmento basali multo angustiore quam
segmentum occludens; longitudine ferè dimidia: tergis bipenniformibus,
margine integro, manubrii acumine ad carinam flexo.
_Hab._--In Oceano Indico.
4. _Dichelaspis Lowei_, Darwin. ('Lepadidæ,' p. 128. Tab. 2, fig. 8.)
D. scutorum segmento basali angustiore quam occludens segmentum,
longitudine ferè 4/5: tergorum parte inferiori duplo latiore quam
occludens scutorum segmentum.
_Hab._--In Insulâ Madeirâ.
5. _Dichelaspis orthogonia_, Darwin. ('Lepadidæ,' p. 130, Tab. 2, fig.
10.)
D. scutorum basali segmento angustiore quam occludens segmentum;
longitudine ferè dimidiâ; duorum segmentorum junctione calcareâ:
tergorum prominentiis marginalibus inæqualibus 5: carinâ deorsum in
parvo calyce lunato terminatâ.
_Hab._--(?)
4. _Genus_--OXYNASPIS.
1. _Oxynaspis celata_, Darwin. ('Lepadidæ,' p. 134, Tab. 3, fig. 1.)
_Hab._--Insulâ Madeirâ.
5. _Genus_--CONCHODERMA.
1. _Conchoderma aurita_, Linn. (Darwin, 'Lepadidæ,' p. 141, Tab. 3,
fig. 4.)
C. capitulo duobus tubularibus quasi-auribus instructo, pone terga
rudimentalia (sæpe nulla) positis: scutis bilobatis: carinâ nullâ, aut
omnino rudimentali: pedunculo longo, a capitulo distincte separato.
_Hab._--In Oceano, pænè ubique.
2. _Conchoderma virgata_, Spengler. (Darwin, 'Lepadidæ,' p. 146, Tab.
3, fig. 2.)
C. scutis trilobatis: tergis intùs concavis, apicibus introrsùm leviter
curvatis: carinâ modicâ, leviter curvatâ: pedunculo in capitulum
coalescente.
_Hab._--In Oceano, pænè ubique.
3. _Conchoderma Hunteri_, Owen. (Darwin, 'Lepadidæ,' p. 153, Tab. 3,
fig. 3.)
C. valvis angustis: scutis trilobatis, prominentiâ laterali non latiore
quam inferior: tergorum parte superiore pæne rectangulè secundùm
aperturæ marginem flexâ: carinâ valde arcuatâ: pedunculo brevi, in
capitulum coalescente.
_Hab._--In Oceano torrido, aut Indico, aut Pacifico.
6. _Genus_--ALEPAS.
1. _Alepas minuta_, Philippi. (Darwin, 'Lepadidæ,' p. 160, Tab. 3, fig.
5.)
A. aperturâ non prominente, capituli longitudinis vix tertiam partem
æquante: scutis corneis, pæne absconditis: longitudine totâ ad quartam
unciæ partem.
_Hab._--In Mari Mediterraneo.
2. _Alepas parasita_, S. Rang. (Darwin, 'Lepadidæ,' p. 163.)
A. aperturâ non prominente, capituli longitudinis 2/3 æquante: scutis
corneis: longitudine totâ ad 2 uncias.
_Hab._--In Mari Mediterraneo et Oceano Atlantico septentrionale.
3. _Alepas cornuta_, Darwin. ('Lepadidæ,' p. 165, Tab. 3, fig. 6.)
A. aperturâ parvâ, leviter prominente: scutis nullis: capitulo
plerumque tribus, parvis, compressis eminentiis secundum carinalem
marginem instructo.
_Hab._--In Insulis Indicis occidentalibus.
4. _Alepas tubulosa_, Quoy et Gaimard. (Darwin, 'Lepadidæ,' p. 169.)
A. aperturâ parvâ prominente et tubulosâ: scutis et prominentiis
secundùm marginem carinalem, nullis.
_Hab._--In Novâ Zealandiâ.
7. _Genus_--ANELASMA.
1. _Anelasma squalicola_, Lovén. (Darwin, 'Lepadidæ,' p. 170, Tab. 5,
fig. 1-7.)
_Hab._--In Mari septentrionali, squalis affixa.
8. _Genus_--ALCIPPE.
1. _Alcippe lampas_, Hancock. (Darwin, 'Balanidæ,' p. 530, Tab. 22, et
Tab. 23, fig. 16-19.)
_Hab._--Littoribus Anglicis, molluscis condita.
9. _Genus_--IBLA.
1. _Ibla Cumingii_, Darwin. ('Lepadidæ,' p. 183, Tab. 4, fig. 8.)
I. (fœm.) valvarum marginibus lateralibus, et superficie interiore,
cæruleis; pedunculi spinis plerumque annulis cæruleo-fuscis.
_Hab._--In Archipel. Philippino, et littore Burmah.
2. _Ibla quadrivalvis_, Cuvier. (Darwin, 'Lepadidæ,' p. 203, Tab. 4,
fig. 9.)
I. (Herm.) valvis et pedunculi spinis sub-flavis; basali tergorum
angulo, introrsùm spectanti, hebete, quia margo carinalis inferior
longiùs quam mango scutalis prominet.
_Hab._--In Australiâ meridionali.
10. _Genus_--SCALPELLUM.
_Sectio † Carinæ umbone sub-centrali._
_Sectio †† Carinæ umbone ad apicem posito._
[_Sectio_ †.]
1. _Scalpellum vulgare_, Leach. (Darwin, 'Lepadidæ,' p. 222, Tab. 5,
fig. 15.)
S. (Herm.) valvis 14, si rostrum pæne rudimentale includatur;
sub-carinâ nullâ; lateribus superioribus inæqualiter ovatis.
_Hab._--In maribus Europæ.
2. _Scalpellum magnum_, Darwin. ('Lepadidæ Fossiles,' p. 18, Tab. 1,
fig. 1.)
S. valvis probabiliter 14, sub-carinâ nullâ; laterum carinalium et
rostralium umbonibus liberè (sicut cornua) prominentibus, dimidiam seu
tertiam partem longitudinis valvarum æquantibus.
_Foss._--In "Coralline Crag Formatione," Angliâ.
3. _Scalpellum ornatum_, J. E. Gray. (Darwin, 'Lepadidæ,' p. 244, Tab.
6, fig. 1.)
S. (Fœm.) valvis 14, sub-rufis; sub-carinâ nullâ; lateribus
superioribus quadranti-formibus, arcu crenâ profundâ notato.
_Hab._--In Africâ meridionali.
4. _Scalpellum rostratum_, Darwin. ('Lepadidæ,' p. 259, Tab. 6, fig. 7.)
S. (Herm.) valvis 15; rostro per-magno; sub-carinâ præsente; laterum
paribus quatuor; pari superiore pentagono.
_Hab._--In Archipel. Philippino.
5. _Scalpellum Peronii_, J. E. Gray. (Darwin, 'Lepadidæ,' p. 264, Tab.
6, fig. 6.)
S. (Herm.) valvis 13; sub-carinâ præsente; laterum paribus tribus; pari
superiore multùm elongato; pedunculi squamis calcareis nullis.
_Hab._--In Australiâ.
6. _Scalpellum Darwinianum_, Bosquet. ('Monograph. Crust. Foss. de
Limbourg,' Tab. 4, fig. 1-5.)
S. carinâ ad angulum 115° flexâ, umbone centrali, costis tenuibus ab
umbone utrinque radiantibus; carinæ tectis et superioris et inferioris
partis rectis, planis, lævibus.
_Foss._--In Cretaceâ superiore Formatione.
7. _Scalpellum Hagenovianum_, Bosquet. (Idem, Tab. 4, fig. 13-16.)
S. carinæ umbone ad 1/6 totius ab apice longitudinis posito; carinæ
tecto plano, utrinque sulcato; parietum et intra-parietum confluentium
parte superiore solummodò cristis instructo ab umbone radiantibus.
_Foss._--In Cretaceâ superiore Formatione.
8. _Scalpellum radiatum_, Bosquet.[158] (Idem, Tab. 4, fig. 17-18.)
[158] Scalpelli 4 aliæ species fossiles, ad secundam generis
sectionem pertinentes, à Dom. Bosquet descriptæ et nominatæ
sunt. Jam cum equidem, ubi valvæ disjunctæ reperiuntur, nomina
specifica non nisi carinis (et interdùm scutis) tribuere soleam,
_S. pulchellum_, Bosq., hic inserere non possum, cujus scuti aut
carinæ fragmenta valdè imperfecta solùm reperta sunt. _Scalpellum
gracile_, _S. pygmæum_, et _S. radiatum_, Bosq., omnia _Scalpello
maximo_ et ejus varietatibus tam propè associantur, ut, cum ipse
specimina non viderim, characteres breves diagnosticos hic exhibere
non audeam. Veruntamen, cum Dom. Bosquet se hujus rei penitus
peritum se probaverit, longissimè abest ut in dubium vocare velim
quin hæ species reverà distinctæ existant.
S. scuti superficie cristis tectâ, inter se aliquantùm distantibus,
ab umbone radiantibus; scuti umbone propè medium marginis occludentis
posito; termino superiore truncato, lato, non acuminato.
_Foss._--In Cretaceâ superiore Formatione.
[_Sectio_ ††.]
9. _Scalpellum rutilum_, Darwin. ('Lepadidæ,' p. 253, Tab. 6, fig. 2.)
S. (Fœm. an Herm.) valvis 14 sub-rufis; sub-carinâ nullâ; carinæ tecto
plano, utrinque cristâ rotundatâ instructo, margine basali truncato.
Lateribus superioribus latitudine duplo longioribus.
_Hab._--(?)
10. _Scalpellum villosum_, Leach. (Darwin, 'Lepadidæ,' p. 274, Tab. 6,
fig. 8.)
S. (Herm.) valvis 14; sub-rostro et sub-carinâ præsentibus; carinâ pæne
rectâ; laterum paribus tribus, pari superiore triangulo.
_Hab._--In maribus orientalibus.
11. _Scalpellum quadratum_, Dixon. (Darwin, 'Lepad. Foss.' p. 22, Tab.
1, fig. 3.)
S. tecto parietibusque carinæ planis, lævibus, simplicibus, margine
basali fere rotundato. Lateribus superioribus quinque-lateralibus,
lævibus.
_Foss._--In "Eocenâ Formatione" Anglicâ.
12. _Scalpellum fossula_, Darwin. ('Lepad. Foss.' p. 24, Tab. 1, fig.
4.)
S. carinâ intra-parietibus instructâ; tecto utrinque costis magnis,
tumidis, superne planatis, marginato; margine basali obtusè acuminato.
Lateribus superioribus quinque-lateralibus, costis duabus modicis ab
apice ad marginem basalem continuatis.
_Foss._--In Cretaceâ superiore Formatione.
13. _Scalpellum maximum_, J. Sowerby. (Darwin, 'Lepad. Foss.' p. 26,
Tab. 2, fig. 1-10.)
S. carinâ intra-parietibus instructâ; tecto subangulari vel
subcarinato; margine basali rectangulariter acuto; totâ valvâ plus
minusve introrsùm arcuatâ, sed margine interno ferè-recto; tecto
transversè plus minusve convexo; superficie pænè lævi, striis
paucis obsoletis longitudinalibus elevatis; tectum, parietes, et
intra-parietes inter se costis plus minusve prominentibus separantur.
_Var. typicum_, (Tab. 2, fig. 1, 4, 5, 8.)
S. carinâ introrsùm leviter arcuatâ, latitudine valvæ altitudinem
superante; tecto transversè leniter arcuato; parietibus
intra-parietibusque angustis, superficie pæne lævi.
_Var. cylindraceum_, (Tab. 2, fig. 2.)
S. parte superiore carinæ liberè prominente, parte interiore
intra-parietibus rotundatis, inflexis, itâ repletâ, ut pæne cylindrica
fiat; superficie externâ lævi, tecto parietibusque pæne confluentibus.
_Var. sulcatum_, (Tab. 2, fig. 3.)
S. carinâ introrsùm valde arcuatâ, sub-carinatâ; valvæ latitudine
circà dimidium altitudinis æquante, tecto transversè præruptè arcuato;
parietibus intra-parietibusque latiusculis. Apice solidè repleto,
liberè paululùm prominente; superficie externâ striis paucis,
rotundatis, ad alterum vel utrumque latus costarum duarum tectum et
parietes separantium.
_Foss._--In Cretaceâ a superiore Formatione.
14. _Scapellum lineatum_, Darwin. ('Lepad. Foss.' p. 35, Tab. 2, fig.
11, 12.)
S. superficie totâ carinæ lineis tenuibus, rotundatis,
longitudinalibus, proximis, microscopicis obtectâ; cristæ centralis
costâ crassiore; costis duabus vel tribus tectum et parietes
separantibus; latitudine valvæ circa dimidium altitudinis æquante;
intra-parietibus latiusculis, nullâ costâ conspicuâ a parietibus
separatis; apice solidè repleto, aliquantulum liberè prominente.
_Foss._--In Cretaceâ inferiore Formatione.
15. _Scapellum hastatum_, Darwin. ('Lepad. Foss.' p. 37, Tab. 2, fig.
13.)
S. carinâ intra-parietibus, intrà paululùm positis, instructâ; valvâ
totâ introrsùm valdè arcuatâ, margine interno non recto; margine basali
acuto, lanceolato; valvâ tenui, lævi, tecto transversè leniter arcuato;
parietibus à tecto vix disjunctis.
_Foss._--In "Grey Chalk, Dover."
16. _Scalpellum angustum_, Dixon. (Darwin, 'Lepad. Foss.' p. 37, Tab.
1, fig. 2.)
S. carinâ angustâ, introrsùm valdè arcuatâ; tecto à parietibus
rectangulè inflexis costâ, (ut videtur) disjuncto; intra-parietibus
usque ad dimidium valvæ pertinentibus, deinde obliquè et abruptè
truncatis; margine basali acutè cuspidato.
_Foss._--In Cretaceâ Formatione.
17. _Scalpellum quadricarinatum_, Reuss. (Darwin, 'Lepad. Foss.' p. 38.)
S. carinâ intra-parietibus latis (ut videtur) instructâ; tecto
transversè plano, lævi, costâ prominente utrinque marginato; margine
basali abruptè truncato.
_Foss._--In "Untern Pläner-Kalke, Bohemia."
18. _Scalpellum trilineatum_, Darwin. ('Lepad. Foss.' p. 38, Tab. 1,
fig. 5.)
S. carinæ tecto transversè leniter arcuato, subcarinato, costâ centrali
et costis duabus lateralibus, rotundatis, tumidis; parietibus angustis
leviter concavis, rectangulè inflexis.
_Foss._--In "Grey Chalk, Dover."
19. _Scalpellum simplex_, Darwin. ('Lepad. Foss.' p. 39, Tab. 1, fig.
9.)
S. carinâ lævi; parietibus angustissimis, rectangulè inflexis; tecto
subcarinato, transversè mediocriter arcuato; margine basali rectangulè
acuminato.
_Foss._--In "Lower Greensand."
20. _Scapellum arcuatum_, Darwin. ('Lepad. Foss.' p. 40, Tab. 1, fig.
7.)
S. valvarum lineis angustis elevatis ab apice radiantibus: carinæ tecto
transversè leniter arcuato, et parietibus rectangulè inflexis, leniter
concavis, lævibus.
_Foss._--In "Gault."
21. _Scalpellum solidulum_, Steenstrup. (Darwin, 'Lepad. Foss.' p. 42,
Tab. 1, fig. 8.)
S. valvarum lineis latiusculis elevatis ab apice radiantibus. Carinæ
parte superiori liberè prominente, et cristâ centrali, internâ,
longitudinali instructâ.
_Foss._--In Cretaceâ superiore Formatione; Scaniâ.
22. _Scalpellum tuberculatum_, Darwin. ('Lepad. Foss.' p. 43, Tab. 1,
fig. 10.)
S. valvarum lineis tenuibus, tuberculatis, elevatis, ab apice
radiantibus: carinæ tecto transversè leniter arcuato, et parietibus
striatis: scuti umbone prope in medio marginis occludentis posito,
costis duobus ab umbone ad angulum basi-lateralem, et ad basalis
marginis medium decurrentibus.
_Foss._--In Cretaceâ Formatione.
23. _Scalpellum semiporcatum_, Darwin. ('Lepad. Foss.' p. 44, Tab. 1,
fig. 6.)
S. carinâ ignotâ: scuti costis duobus ab umbone ad angulum
basi-lateralem et ad marginis basalis medium decurrentibus;
superficie inter hanc costam et marginem occludentem lineis tenuibus,
longitudinalibus, elevatis instructâ.
_Foss._--In Cretaceâ superiore Formatione; Scaniâ.
24. _Scalpellum_ (?) _cretæ_, Steenstrup. (Darwin, 'Lepad. Foss.' p.
45, Tab. 1, fig. 11.)
S. valvis lævibus tenuissimis: scuti umbone propè medium marginis
occludentis posito; costis tribus obscuris ab umbone ad angulos
tergo-lateralem et basi-lateralem, et ad medium marginis basalis
decurrentibus: carinæ apice et margine basali acutis; distincti
parietes absunt.
_Foss._--In Cretaceâ Formatione.
11. _Genus_--POLLICIPES.
†. _Scuta, aut lævia aut lineis incrementi tenuibus solùm notata._
A. _Scuta, costâ ab apice ad centrum marginis basalis non decurrente._
B. _Scuta, costâ, nonnunquam subobsoletâ, ab apice ad centrum marginis
basalis decurrente._
††. _Scuta, aut longitudinaliter aut transversè (i. e. secundum lineas
incrementi) costata._
[_Sectio_ †, A.]
1. _Pollicipes cornucopia_, Leach. (Darwin, 'Lepadidæ,' p. 298, Tab. 7,
fig. 1.)
P. capitulo, valvarum duobus aut pluribus sub rostro verticillis
instructo: valvis albis, aut glaucis: pedunculo, squamarum densis
verticillis symmetricè dispositis.
_Hab._--In Europâ.
2. _Pollicipes elegans_, Lesson. (Darwin, 'Lepadidæ,' p. 304.)
P. capitulo, valvarum duobus aut pluribus sub rostro verticillis
instructo: valvis et pedunculi squamis rufo-aurantiacis: squamarum
verticillis densis symmetricè dispositis.
_Hab._--In Peruviâ et Mexico.
3. _Pollicipes polymerus_, G. B. Sowerby. (Darwin, 'Lepadidæ,' p. 307,
Tab. 7, fig. 2.)
P. capitulo, valvarum duobus, tribus, aut pluribus sub rostro
verticillis instructo; valvis sub-fuscis; lateribus à supremo ad
infimum gradatim quoad magnitudinem positis; carinæ margine basali
(introrsùm spectanti) ad medium excavato; pedunculi squamarum
verticillis densis, symmetricè dispositis.
_Hab._--In Californiâ et Oceano Pacifico meridionali.
4. _Pollicipes spinosus_, Quoy. (Darwin, 'Lepadidæ,' p. 324, Tab. 7,
fig. 4.)
P. capitulo valvarum uno aut pluribus sub rostro verticillis instructo:
laterum pari superiore vix inferioribus longiore: membranâ valvas
tegente (post desiccationem) subfuscâ flavescente: pedunculi squamis
inæqualibus, non symmetricis: verticillis longiusculè distantibus.
_Hab._--Novâ Zealandiâ.
5. _Pollicipes sertus_, Darwin. ('Lepadidæ,' p. 327, Tab. 7, fig. 5.)
P. capitulo valvarum uno aut pluribus sub rostro verticillis instructo:
laterum pari superiore vix inferioribus longiore: membranâ valvas
tegente (post desiccationem) fusco rufescente obscuro: rostro dimidiam
carinæ longitudinem æquante, superficiei internæ altitudine latitudinem
plus duplo superante: pedunculi squamis inæqualibus, non symmetricis:
verticillis longiusculè distantibus.
_Hab._--Novâ Zealandiâ.
6. _Pollicipes concinnus_, J. Morris. (Darwin, 'Lepad. Foss.' p. 50,
Tab. 3, fig. 1.)
P. scutis pæne quadratis, margine basali propè rostrum subconcavo,
segmento tergo-laterali, è lineis incrementi, ut videtur, reflexis
formato, lato, rotundato et prominente: tergis latis, pæne quadratis:
carinæ margine basali, ut videtur, acuto.
_Foss._--In 'Oxford Clay.'
7. _Pollicipes ooliticus_, Buckman. (Darwin, 'Lepad. Foss.' p. 51, Tab.
3, fig. 2.)
P. scutis triangulis; superficie undulatâ; margine basali rectangulè
ad marginem rectum tergo-lateralem posito; segmentum tergo-laterale
è lineis reflexis incrementi formatum deest. Carinâ pæne rectâ,
semi-cylindricâ, margine basali quadrato.
_Foss._--In 'Stonesfield Slate.'
8. _Pollicipes Nilssonii_, Steenstrup. (Darwin, 'Lepad.' Foss. p. 52,
Tab. 3, fig. 11.)
P. scutis triangulis, planis: margine basali cum margine occludente
angulum pæne rectum, cum margine recto tergo-laterali, angulum
aliquanto minorem formante: deest segmentum tergo-laterale, è lineis
incrementi reflexis formatum. Carinâ introrsùs admodùm arcuatâ, crassâ;
marginis basalis mucrone obtuso.
_Foss._--In Cretaceâ superiore Formatione; Scaniâ.
9. _Pollicipes Hausmanni_, Koch. (Darwin, 'Lepad. Foss.' p. 53, Tab. 3,
fig. 3.)
P. scutis subtriangulis, angulo basi-laterali valde rotundato; apice
producto; margine basali cum margine occludente angulum pæne rectum
formante; internâ apicis superficie prominente, margineque tergali
sulcato.
_Foss._--In 'Hilsthon,' Germaniâ.
10. _Pollicipes politus_, Darwin. ('Lepad.' Foss. p. 54, Tab. 3, fig.
4.)
P. scutis ferè rhombicis, lævissimis; margine basali cum margine
occludente angulum recto majorem formante; margine occludente
projecturâ parietali, lineari, minutâ instructo; internâ apicis
superficie concavâ.
_Foss._--In 'Gault.' (?)
11. _Pollicipes elongatus_, Steenstrup. (Darwin, 'Lepad. Foss.' p. 55,
Tab. 3, fig. 5.)
P. scutis pæne quadratis; margine occludente et parte inferiori
marginis tergo-lateralis rectangulè ad marginem basalem positis; apice
obtuso.
_Foss._--In Cretaceâ Formatione albâ. Daniâ.
12. _Pollicipes acuminatus_, Darwin. ('Lepad. Foss.' p. 56, Tab. 3,
fig. 6.)
P. scutis elongatis, triangulis; margine basali cum margine occludente
angulum recto longè minorem formante; internâ apicis superficie concavâ.
_Foss._--In Cretaceâ Formatione.
13. _Pollicipes Angelini_, Darwin. ('Lepad. Foss.' p. 56, Tab. 3, fig.
7.)
P. scutis elongatis, triangulis, margine basali propè angulum rostralem
in prominentiam obliquè rotundatam producto; internâ apicis superficie
prominente, margine occludente sulcato.
_Foss._--In Cretaceâ superiore Formatione.
14. _Pollicipes reflexus_, J. Sowerby. (Darwin, 'Lepad. Foss.' p. 58,
Tab. 3, fig. 8.)
P. scutis tenuibus, subovalibus; margine basali cum margine occludente
angulum recto longè majorem formante; costâ obsoletâ rotundatâ ab
apice ad angulum basi-lateralem decurrente, valvam in duas ferè
æquales partes separante. Carinâ lineari, transversè abruptè arcuatâ;
margine basali multum producto, apice truncato. Lateribus superioribus
sub-pentagonis.
_Foss._--In Angliâ: 'Eocenâ Formatione.'
[_Sectio_ †, B.]
15. _Pollicipes carinatus_, Philippi. (Darwin, 'Lepad. Foss.' p. 60,
Tab. 3, fig. 9.)
P. scutis crassiusculis ad formam trianguli æquianguli accedentibus;
margine occludente externè costâ, humili firmato; costâ firmâ ab apice
ad centrum marginis basalis decurrente; margine basali recto; angulo
basi-laterali truncato, brevi; segmentum tergo-laterale ex lineis
incrementi reflexis formatum, deest. Carinâ externè valdè carinatâ cum
sulco laterali ad utrumque latus.
_Foss._--In Form. Tertiariâ, Siciliâ.
16. _Pollicipes glaber_, Roemer. (Darwin, 'Lepad. Foss.' p. 61, Tab. 3,
fig. 10.)
P. scutis subtenuibus, latiusculis, ad formam trianguli æquianguli
accedentibus; margine basali non prorsus recto; tergo-laterali segmento
e zonis incrementi reflexis formato, ubi (latissimum), reliquæ valvæ
dimidium æquante: margine tergali apicis intùs sulcato. Carinæ margine
basali obtusè acuminato. Lateribus superioribus triangulis, tertiam
partem longitudinis tergorum æquantibus. Lateribus anticis inferioribus
singulis costâ propè terminum marginis basalis decurrente.
_Foss._--In Cretaceâ, super. et inferiore Formatione.
17. _Pollicipes unguis_, J. Sowerby. (Darwin, 'Lepad. Foss.' p. 64,
Tab. 4, fig. 1.)
P. scutis incognitis, verisimiliter ut in P. glabro. Margine basali
carinæ obtusè acuminato, sub-rotundato. Lateribus superioribus
elongatis triangularibus, dimidium longitudinis tergorum superantibus.
Lateribus anticis inferioribus costâ subcentrali instructis.
_Foss._--In 'Gault.'
18. _Pollicipes validus_, Steenstrup. (Darwin, 'Lepad. Foss.' p. 68,
Tab. 4, figs. 2 and 3.)
P. scutis crassis, angustis; margine occludente externè costâ rotundatâ
forti firmato; intus prominentiâ rostrali infra marginem rectum basalam
dependente; costâ ab apice ad marginem basalem propiùs ad rostralem
quam ad basi-lateralem angulum accedente. Carinâ lævissimâ, transversè
semicylindricâ; parte superiori liberè prominente, internè aut planâ
aut cristâ centrali instructâ.
_Foss._--In Cretaceâ sup. Formatione, Scaniâ et Maestricht.
19. _Pollicipes Darwinianus_, Bosquet. (Monographie Crust. Foss. de
Limbourg, Tab. 1, fig. 8-16.)
P. scuti marginibus basali et occludente angulum solum 35° formantibus;
intùs prominentiâ rostrali infra marginem basalem dependente; costâ ab
apice ad marginem basalam propemodùm ad angulum rostralem accedente.
Tergo pæne pentagono, intùs prominentiâ à medio marginis scutalis
surgente.
_Foss._--In Cretaceâ superiore Formatione.
20. _Pollicipes dorsatus_, Steenstrup. (Darwin, 'Lepad. Foss.' p. 69,
Tab. 4, fig. 4.)
P. scutis crassis; ad formam trianguli æquianguli accedentibus; margine
occludente externè costâ rotundatâ firmato; angulo basi-laterali
latè truncato, dimidium longitudinis marginis basalis veri, æquante;
tergo-laterali segmento ex lineis incrementi reflexis formato,
angustissimo. Tergorum costâ rectâ, latâ, proclivi, ab apice ad angulum
basalem decurrente; angulo basali scutum versùs, obliquè truncato.
_Foss._--In Cretaceâ superiore Formatione, Daniâ.
[_Sectio_ ††.]
21. _Pollicipes mitella_, Linn. (Darwin, 'Lepadidæ,' p. 316, Tab. 7,
fig. 3.)
P. capitulo valvarum unico sub rostro verticillo instructo: laterum
pari superiore (introrsum spectanti) inferiorum magnitudinem ter
aut quater superante: lateribus inferioribus utrinque obtegentibus:
pedunculi squamarum verticillis densis, symmetricè dispositis.
_Hab._--In Insulis Indicis orientalibus.
22. _Pollicipes striatus_, Darwin. ('Lepad. Foss.' p. 70, Tab. 4, fig.
5.)
P. valvis longitudinaliter striatis: scutis sub-triangulis, margine
tergo-laterali valdè arcuato et prominente; costâ obscurâ, rotundatâ,
ab apice ad angulum basi-lateralem decurrente; internâ apicis
superficie solidè repletâ, sine sulco manifesto.
_Foss._--In Cretaceâ superiore Formatione, Angliâ.
23. _Pollicipes semilatus_, Darwin. ('Lepad. Foss.' p. 72, Tab. 4, fig.
6.)
P. valvis longitudinaliter et transversè costatis: scutorum margine
basali brevi, recto, cum margine occludente angulum rectum formante;
costâ parietali tenuissimâ ab apice, ad angulum prominentem
basi-lateralem decurrente; hæc valvam in duas partes inæquales dividit,
è quibus portio tergo-lateralis latior est.
_Foss._--In Cretaceâ Formatione.
24. _Pollicipes rigidus_, J. Sowerby. (Darwin, 'Lepad. Foss.' p. 73,
Tab. 4, fig. 7.)
P. valvis transversè costatis: scutorum margine basali recto, cum
margine occludente angulum recto majorem formante; costâ angustissimâ,
parietali, ab apice ad angulum basi-lateralem decurrente: tergis,
costâ curvâ, parietali, ad angulum basalem decurrente instructis;
apice basali in prominentiam parvam terminante, lateribus prominentiæ
parallelis.
_Foss._--In 'Gault.'
25. _Pollicipes fallax_, Darwin. ('Lepad. Foss.' p. 75, Tab. 4, fig. 8.)
P. valvis transversè costatis: scutis margine basali non recto, angulum
pæne rectum cum margine occludente formante; costâ, parietibus
obliquis, ab apice ad angulum basi-lateralem decurrente: tergis, costâ
curvatâ, parietibus obliquis, ad angulum basalem latum, rotundatum,
decurrente.
_Foss._--In Cretaceâ superiore Formatione, Angliâ, Scaniâ, Hanoveriâ.
26. _Pollicipes elegans_, Darwin. ('Lepad. Foss.' p. 76, Tab. 4, fig.
9.)
P. valvis longitudinaliter et transversè striatis: scutorum margine
basali recto, cum margine occludente angulum recto paulo majorem
formante; costâ parietali, latiore quam pro solitâ incrementorum
linearum latitudine, ab apice ad angulum basi-lateralem decurrente:
tergorum costâ parietali, rectâ, ad apicem basalem, acuminatum
decurrente.
_Foss._--In Cretaceâ superiore Formatione, Scaniâ, Daniâ.
[_Species aliquot quas, scutis incognitis, auctores a valvis insignibus
nominârunt, hìc pro tempore solummodo iterum describuntur._]
27. _Pollicipes Bronnii_, F. Roemer. (Darwin, 'Lepad. Foss.' p. 77,
Tab. 4, fig. 10.)
P. carinâ lævi, subcarinatâ, margine basali arcuato et turgido; totâ
valvâ vel extrorsùm arcuatâ vel pæne rectâ; internè, costis duabus
elevatis ad utrumque latus partis superioris liberè prominentis.
_Foss._--In 'Upper Greensand,' et 'Hils-conglomerat.'
28. _Pollicipes planulatus_, J. Morris. (Darwin, 'Lepad. Foss.' p. 27,
Tab. 4, fig. 11.)
P. tergis subrhombicis, latis, lævibus, apice basali latè truncato,
latitudine dimidium longitudinis marginis occludentis æquante; apice
basali truncato angulum rectum cum margine scutali formante; parte
superiori marginis scutalis cuspidem latam, rotundatam, leviter
prominentem formante.
_Foss._--In 'Oxford Clay.'
12. _Genus_--LITHOTRYA.
1. _Lithotrya dorsalis_, G. B. Sowerby. (Darwin, 'Lepadidæ,' p. 351,
Tab. 8, fig. 1 _a′_.)
L. scutis terga angustè obtegentibus: carinâ intùs concavâ; rostro,
duorum aut trium squamarum subjacentium latitudinem æquante: lateribus,
squamarum quinque subjacentium longitudinem æquantibus, superficie
internâ angustè ellipticâ: pedunculi squamis superioribus verticillum
secundum minus duplo superantibus.
_Hab._--Insulis Indicis occidentalibus.
2. _Lithotrya cauta_, Darwin. ('Lepadidæ,' p. 356, Tab. 8, fig. 3.)
L. scutis terga amplè obtegentibus: carinâ intus concavâ: rostro,
squamarum subjacentium latitudinem vix æquante: lateribus, squamas
subjacentes sesquitertio superantibus, superficie internâ latè
ellipticâ: pedunculi squamis superioribus verticillum secundum pæne
quadruplo superantibus.
_Hab._--In Australiâ.
3. _Lithotrya Nicobarica_, Reinhardt. (Darwin, 'Lepadidæ,' p. 359, Tab.
8, fig. 2.)
L. scutis terga angustè obtegentibus: carinæ cristâ internâ tenui in
parte superiore positâ: rostro conspicuo, squamarum sex subjacentium
latitudinem æquante: lateribus, superficie internâ triangulâ squamarum
septem subjacentium latitudinem æquantibus.
_Hab._--In Insulis Nicobaricis et Timor.
4. _Lithotrya rhodiopus_, J. E. Gray. (Darwin, 'Lepadidæ,' p. 363, Tab.
8, fig. 4.)
L. scutis terga amplè obtegentibus: carinæ cristâ internâ tenui, in
parte superiore positâ: lateribus, superficie internâ symmetricè et
latè ovatâ, carinæ latitudinis plus quam tertiam partem æquantibus:
tergorum basali apice tenui, et angulo carinali producto: rostro et
pedunculo ignotis.
_Hab._--(?)
5. _Lithotrya truncata_, Quoy. (Darwin, 'Lepadidæ,' p. 366, Tab. 9,
fig. 1.)
L. scutis in profundam tergorum plicam insertis: carinæ cristâ centrali
prominente et rotundatâ in parte superiore: rostro et lateribus
rudimentalibus, carinæ latitudinis quindecimam fere partem æquantibus.
_Hab._--In Arch. Philippino et Mari Pacifico.
6. _Lithotrya Valentiana_, J. E. Gray. (Darwin 'Lepadidæ,' p. 371, Tab.
8. fig. 5.)
L. scutis in profundam tergorum plicam insertis: tergorum opposito
superiore margine, plicâ alterâ æquè profundâ instructo: carinæ cristâ
prominente centrali, marginibus quadratis, in parte superiore: rostro
rudimentali: lateribus et pedunculo ignotis.
_Hab._--In Mari Rubro.
13. _Genus_--LORICULA.
1. _Loricula pulchella_, G. B. Sowerby, jun. (Darwin, 'Lepad. Foss.' p.
81, Tab. 5.)
_Foss._--In Cretaceâ inferiore Formatione.
ORDO II. ABDOMINALIA.
1. _Genus_--CRYPTOPHIALUS.
1. _Cryptophialus minutus_, Darwin. ('Balanidæ,' p. 566, Tab. 23 et 24.)
_Hab._--In littore occidentali Americæ meridionalis.
ORDO III. APODA.
1. _Genus_--PROTEOLEPAS.
1. _Proteolepas bivincta_, Darwin. ('Balanidæ,' p. 589, Tab. 24 et 25.)
_Hab._--In insulis Indicis occidentalibus.
DESCRIPTION OF PLATES.
PLATE 1.
BALANUS TINTINNABULUM.
Fig.
_a_, _b_, and
_f_ (upper shell), Balanus tintinnabulum,--_var._ communis.
_c_ and _f_ (part of shell), " " " validus.
_d_, " " " " coccopoma.
_e_, " " " " concinnus.
_f_ (part of shell), " " " validus.
_g_, " " " " zebra.
_h_, " " " " crispatus.
_i_, " " " " spinosus.
_k_, " " " " occator.
_l_, " " " " d'Orbignii.
PLATE 2.
_Genus_--BALANUS.
Fig.
1 _a_, Balanus tintinnabulum,--_var._ communis, scutum.
1 _b_, " " " occator "
1 _c_, 1 _d_, and 1 _e_, " " " communis "
1 _f_, " " " coccopoma "
1 _g_, " " " concinnus "
1 _h_, (magnified) " " (_young_) " vesiculosus "
1 _i_, and 1 _k_, " " " communis, tergum.
1 _l_, " " " coccopoma "
1 _m_,and 1 _n_, " " " d'Orbignii "
1 _o_, " " " coccopoma "
2 _a_, 2 _b_, 2 _c_, 2 _d_, Balanus tulipiformis, scutum and
tergum, external and internal views of.
3 _a_, 3 _b_, 3 _c_, 3 _d_, Balanus psittacus, scutum and tergum,
external and internal views of.
4 _a_, 4 _b_, Balanus Capensis, scutum and tergum, internal view of.
5 _a_, 5 _b_, Balanus nigrescens " "
6 _a_, Balanus decorus, scutum, internal view; 6 _b_, tergum,
external view.
7 _a_, Balanus vinaceus, scutum, internal view; 7 _b_, do.
external view; 7 _c_, tergum, external view; 7 _d_, basal
edge of the parietes of one of the compartments, showing the
cancellated structure of the inner lamina.
PLATE 3.
_Genus_--BALANUS.
Fig. 1 _a_, Balanus Ajax; 1 _b_, scutum external; 1 _c_, tergum
external; and 1 _d_, scutum, internal view.
2 _a_, 2 _b_, Balanus stultus, scutum and tergum, external
views; 2 _c_ and 2 _d_, scutum, internal and external views.
3 _a_, Balanus calceolus; 3 _b_, variety; 3 _c_, scutum,
internal view; 3 _d_, tergum, internal view; 3 _e_, spur of
tergum toothed.
4 _a_, Balanus galeatus; 4 _b_, 4 _c_, scutum and tergum,
internal view.
5 _a_, 5 _b_, Balanus cymbiformis, scutum and tergum,
internal view.
6 _a_, Balanus navicula; 6 _b_, 6 _c_, scutum and tergum,
internal view; 6 _d_, scutum, external view.
7 _a_, Balanus trigonus; 7 _b_ and 7 _c_, scutum, external
views, two varieties; 7 _d_, scutum, internal view; 7 _e_
and 7 _f_, tergum, external views, two varieties.
PLATE 4.
_Genus_--BALANUS.
Fig. 1 _a_ and 1 _b_, Balanus spongicola, external views of
scutum and tergum; 1 _c_, _var._ of the tergum, from a
specimen from the Cape of Good Hope; 1 _d_, _var._ of the
tergum from the West Indies.
2, Balanus lævis.
2 _a_, " " fossil, elongated _var._, showing
lower part of basal cup filled
up with a cancellated mass.
2 _b_ and 2 _c_, " " scutum, external and internal view
of.
2 _d_ and 2 _e_, " " scutum (_variety_), internal and
external view of.
2 _f_, " " scutum (_variety_), external view.
2 _g_, " " tergum, external view.
3 _a_, Balanus perforatus, scutum, internal view; 3 _b_,
tergum, internal view; 3 _c_, tergum, _var._ external view.
4 _a_, Balanus concavus (_var._ from Panama), scutum,
internal view; 4 _b_, scutum, external view; 4 _c_, tergum,
external view; 4 _d_, tergum, fossil, Coralline Crag; 4
_e_, tergum, fossil, Italy.
PLATE 5.
_Genus_--BALANUS.
Fig. 1 _a_, Balanus perforatus,--_var._ angustus.
1 _b_, " " " Cranchii.
1 _c_, " " " mirabilis.
1 _d_, " " " fistulosus.
2 _a_, " amphitrite,--_var._ venustus.
2 _b_, " " _var._ cirratus.
2 _c_, " " " pallidus.
2 _d_, " " " Stutchburii.
2 _e_, " " " communis.
2 _f_, " " " niveus.
2 _g_, " " " obscurus.
2 _h_, " " scutum, _var._ communis.
2 _i_, " " " " Stutchburii.
2 _k_, " " tergum, " pallidus.
2 _l_, " " " " communis.
2 _m_, 2 _n_, and 2 _o_," " _var._ Stutchburii.
3 _a_, Balanus pœcilus, scutum, internal view; 3 _b_, tergum,
external.
4 _a_, Balanus eburneus, scutum, external view; 4 _b_,
tergum, external; 4 _c_, scutum, internal; 4 _d_, tergum
(_var._), internal.
PLATE 6.
_Genus_--BALANUS.
Fig. 1 _a_, Balanus improvisus; 1 _b_, scutum, internal; 1 _c_,
tergum, external view.
2 _a_, Balanus nubilus, scutum, internal view; 2 _b_, tergum,
ditto; 2 _c_, tergum, external view.
3 _a_, 3 _b_, Balanus corrugatus, scutum and tergum, internal
view.
4 _a_, Balanus porcatus; 4 _b_ and 4 _c_, scutum and tergum,
internal views; 4 _d_, tergum external; 4 _e_, basal edge
of the parietes of wall of shell.
5 _a_, Balanus patellaris; 5 _b_, scutum, internal; 5 _c_,
tergum, external.
6 _a_, Balanus crenatus.
6 _b_, " " smooth _var._
6 _c_, " " elongated _var._
6 _d_ and 6 _e_, " scutum and tergum, external views.
6 _f_ and 6 _g_, " " " internal views.
PLATE 7.
_Genus_--BALANUS.
Fig. 1 _a_, Balanus glandula, scutum, internal; 1 _b_, tergum,
external.
2 _a_, Balanus balanoides; 2 _b_, portion of basal edge of
parietes of a compartment; 2 _c_, tergum; 2 _d_, tergum,
_var._
3 _a_, Balanus cariosus; 3 _b_, portion of basal edge of
parietes of a compartment; 3 _c_, scutum, internal view of;
3 _d_, tergum, internal; 3 _e_, tergum, external view.
4 _a_, Balanus declivis, lateral view, with basal membrane
removed; 4 _b_, rostrum (to left hand), two lateral
compartments, and carina; 4 _c_ and 4 _d_, scutum and
tergum, external views.
5 _a_, Balanus Hameri; 5 _b_, scutum, internal view; 5 _c_,
tergum, external view.
6 _a_, Balanus amaryllis; 6 _b_, scutum, internal; 6 _c_,
tergum, external view.
7 _a_, Balanus allium; 7 _b_ and 7 _c_, scutum, external and
internal views; 7 _d_, tergum, external.
8 _a_, Balanus cepa, scutum, internal; 8 _b_, scutum,
external; 8 _c_, tergum, external view.
PLATE 8.
_Genus_--BALANUS.
Fig. 1, Balanus quadrivittatus.
2 _a_, Balanus terebratus, part of basis seen from the
outside; 2 _b_, the same, seen from within.
3 _a_, Balanus vestitus, scutum, internal view; 3 _b_,
tergum, external view.
4 _a_, Balanus imperator, scutum, internal view; 4 _b_,
tergum, external view; 4 _c_, portion of basal edge of wall
of the shell.
5 _a_, Balanus flosculus.
5 _b_, " " _var._ sordidus.
5 _c_, " " scutum, internal view.
5 _d_, " " " " _var._
5 _e_ and 5 _f_, " tergum, internal and external views of.
6 _a_, Balanus bisulcatus, scutum, external view; 6 _b_,
tergum, external view; 6 _c_, scutum, internal view.
7, Balanus dolosus, tergum, external view.
8 _a_, Balanus unguiformis, scutum, internal view; 8 _b_,
tergum, do.
9, Balanus varians, tergum, external view.
10 _a_, Balanus inclusus.
10 _b_, " " _var._ that has been attached to a
branch of a coralline.
10 _c_, " " " seen from beneath, showing the
narrow furrowed basis and wide
lateral compartments; the other
four compartments, namely, the
two carino-lateral, the carina,
and rostrum, being of very small
size.
PLATE 9.
_Sub-Genus_--ACASTA.
Fig. 1 _a_, Acasta spongites; 1 _b_, scutum, external view; 1
_c_, scutum, internal; 1 _d_, tergum, internal.
2 _a_, Acasta sulcata; 2 _b_, _var._, lateral compartment of;
2 _c_ and 2 _d_, terga, internal views of two varieties.
3 _a_, Acasta cyathus; 3 _b_ and 3 _c_, scutum and tergum,
internal views.
4, Acasta undulata, scutum, external view.
5 _a_, Acasta glans, basis seen from within; 5 _b_,
compartments of a smaller specimen, seen from within;
carina on the right-hand, cut into two; carino-lateral
and lateral compartments to the left-hand; 5 _c_, tergum,
internal view.
6 _a_, Acasta lævigata, tergum, internal view; 6 _b_, another
_var._
7 _a_, Acasta fenestrata; 7 _b_, scutum, and 7 _c_, tergum,
internal views.
8 _a_, Acasta purpurata; 8 _b_, scutum; 8 _c_, tergum,
internal views.
9 _a_, Acasta sporillus; 9 _b_, carino-lateral compartment
(almost rudimentary) and lateral compartment, seen from
within; 9 _c_ and 9 _d_, scutum and tergum, seen from
within.
PLATE 10.
_Genus_--TETRACLITA.
Fig. 1 _a_, Tetraclita porosa, _var._ communis.
1 _b_, " " " rubescens.
1 _c_, " " " nigrescens.
1 _d_, " " " elegans.
1 _e_, " " " communis, young, with radii
developed.
1 _f_, " " " patellaris.
1 _g_, " " basal edge of wall of shell.
1 _h_, " " basal edge of wall of shell, _var._
elegans, showing also the
structure of one of the
rudimentary radii or sutures.
1 _i_, " " internal view of scutum; 1 _k_, do.
of tergum; 1 _l_, do. of scutum,
_var._; 1 _m_, do. of tergum, _var._
2 _a_, Tetraclita serrata; 2 _b_, portion of external surface
of wall, low down, near basis; 2 _c_ and 2 _d_, scutum and
tergum, internal views.
3 _a_, Tetraclita rosea, scutum; 3 _b_, tergum; 3 _c_, whole
shell; 3 _d_, basal edge of wall of shell.
PLATE 11.
_Genera_--TETRACLITA AND ELMINIUS.
Fig. 1 _a_, Tetraclita purpurascens, _var._ with surface
disintegrated, and with the sutures united and obliterated.
1 _b_, Tetraclita purpurascens, _var._ well preserved, with
the radii developed; 1 _c_ and 1 _d_, _var._ scutum and
tergum, internal views.
2 _a_, Tetraclita costata; 2 _b_ and 2 _c_, scutum and tergum.
3 _a_, Tetraclita vitiata; 3 _b_, basal edge of portion of
wall; 3 _c_ and 3 _d_, scutum and tergum, internal views
of; 3 _e_, tergum, internal view, _var._
4 _a_, Tetraclita cœrulescens; 4 _b_, scutum and tergum
united, external view; 4 _c_ and 4 _d_, scutum and tergum,
internal views.
5 _a_, Tetraclita radiata; 5 _b_, scutum and tergum united,
external view; 5 _c_ and 5 _d_, scutum and tergum, internal
views.
6 _a_, Elminius Kingii; 6 _b_ and 6 _c_, scutum and tergum,
internal views of; 6 _d_, tergum, _var._ external view of;
6 _e_, tergum, another _var._ internal view of.
PLATE 12.
_Genera_--ELMINIUS AND PYRGOMA.
Fig. 1 _a_, Elminius modestus; 1 _b_, scutum, internal view;
1 _c_, tergum, internal view; 1 _d_, do., tergum, _var._;
1 _e_, do., tergum, _var._
2 _a_, Elminius plicatus; 2 _b_, basal edge of portion of
wall of shell; 2 _c_ and 2 _d_, scutum and tergum, internal
views of; 2 _e_ and 2 _f_, scutum and tergum, internal
views of, _var._
3, Elminius simplex; shell.
4 _a_, Pyrgoma Anglicum; 4 _b_ and 4 _c_, scutum and tergum,
internal views.
5 _a_, Pyrgoma cancellatum, shell, seen from above; 5 _b_,
shell seen on the under side; 5 _c_ and 5 _d_, scutum and
tergum, external views; 5 _e_ and 5 _f_, scutum and tergum,
internal views of.
6, Pyrgoma Stokesii, shell imbedded in coral.
7 _a_, Pyrgoma conjugatum, shell, external view of; 7 _b_,
scutum and tergum, calcified together, seen from the
outside; 7 _c_, do., internal view of.
PLATE 13.
_Genera_--PYRGOMA AND CREUSIA.
Fig. 1 _a_, Pyrgoma grande; 1 _b_, scutum and tergum, calcified
together, external view of; 1 _c_, scutum and tergum,
internal view of; 1 _d_, the two scuta and terga, external
view, seen from above, having their natural position, when
closed.
2 _a_, Pyrgoma milleporæ, shell viewed from above; 2 _b_,
internal view of, showing the sheath; 2 _c_, scutum and
tergum, united, external view of; 2 _d_, scutum, _var._
external view of; 2 _e_, scutum, internal view of; 2 _f_,
tergum, internal view of.
3 _a_, Pyrgoma dentatum shell, external view of; 3 _b_,
scutum, _var._ 1, external view of; 3 _c_, tergum, _var._
1, external view of; 3 _d_, tergum, _var._ 2, external
view of; 3 _e_, scutum, _var._ 3, external view of; 3 _f_,
tergum, _var._ 3, external view of; 3 _g_, scutum and
tergum, _var._ 1, joined together, internal view of.
4 _a_, Pyrgoma crenatum, scutum, external view of, showing
the great depending adductor plate and large occludent
ledge; 4 _b_, external view of.
5 _a_, Pyrgoma monticulariæ, external view of; 5 _b_, another
specimen, internal view of; 5 _c_, another specimen,
external view of; 5 _d_, portion of circumference of shell,
viewed externally, much magnified, showing the outer
lamina, and the irregular, longitudinal septa; 5 _e_,
another specimen of the same, with the longitudinal septa
more regular; 5 _f_, scutum and tergum completely calcified
together, external view of.
6 _a_, Creusia spinulosa, shell, seen from above.
6 _b_, 6 _c_, Creusia spinulosa, _var._ 1, scutum and tergum,
internal view of; 6 _d_, scutum and tergum, joined
together, external view of.
6 _e_, 6 _f_, 6 _g_, Creusia spinulosa, _var._ 2, 6 _e_,
scutum, internal view; 6 _f_, tergum, internal view of;
6 _g_, tergum, of another specimen, internal view of.
6 _h_, Creusia spinulosa, _var._ 3, scutum and tergum joined
together, external view of.
PLATE 14.
_Genera_--CREUSIA AND CHELONOBIA.
Fig. 6 _i_, 6 _k_, 6 _l_, Creusia spinulosa, _var._ 4, 6 _i_,
tergum, external view; 6 _k_, the same, from another
individual; 6 _l_, the same, internal view.
6 _m_, Creusia spinulosa, _var._ 5, tergum, external view.
6 _n_, 6_o_, 6_p_, 6_q_, Creusia spinulosa, _var._ 6, 6 _n_,
scutum, external view of; 6 _o_, tergum of same individual,
external view of; 6 _p_, scutum of another individual,
external view of; 6 _q_, tergum of latter, external view of.
6 _r_, Creusia spinulosa, _var._ 7, scutum, internal view of.
6 _s_, Creusia spinulosa, _var._ 9, scutum and tergum,
calcified together, external view.
6 _t_, Creusia spinulosa, _var._ 10, scutum and tergum,
calcified together, external view.
6 _U_, Creusia spinulosa, _var._ 11, shell seen from above; 6
_u_, scutum and tergum, calcified together, external view.
Fig. 1 _a_, Chelonobia testudinaria, shell seen from above; a
specimen has been chosen, with some of the radii smooth,
and others more or less notched; 1 _b_, scutum, internal
view of; 1 _c_, tergum, internal view of; 1 _d_, scutum and
tergum, joined together, external view of.
2, Chelonobia caretta, shell seen from above.
3 _a_, Chelonobia patula, shell seen from above; 3 _b_,
tergum, external view of.
4, Carina of Chelonobia patula, seen from within; _a a_, outer
lamina of wall, joined by the longitudinal septa to _b_,
the inner lamina; _e c e_, sheath, descending to the base
of shell, with the middle part largely hollowed out; the
upper part of the sheath, above _d d_, is transversely
marked by the lines of attachment of the opercular
membrane; _d_, _d_, alæ.
5, View of the edge of the radius of the rostrum of Chelonobia
testudinaria.
_a_, _a_, inner portion of radius.
_b_, thick outer lamina of the radius, transversely pitted;
beneath this pitted portion, which forms only the outer
surface of the radius, and distinct from the inner portion
_a_, the arborescent and dentated sutural edges of this
outer lamina are seen.
_c_, outer lamina of wall of compartment.
_d_, the exterior septum, connecting the outer lamina of
the wall to the inner lamina; the basal edge of the inner
lamina of the wall is seen beneath and without the lower
end of the inner portion (_a_) of the radius.
PLATE 15.
_Genera_--CHELONOBIA AND CORONULA.
Fig. 1, Transverse section of shell of Chelonobia testudinaria.
B A B, Compound rostrum: A being the true rostrum.
B B, the rostro-lateral compartments, which normally, in
the Balaninæ, are quite confounded with the true rostrum:
the sutures separating these three compartments are not
continued through the outer lamina.
C, left-hand lateral compartment.
D, left-hand carino-lateral compartment.
E, carina.
S S, Sutures, six in number, separating the six compartments.
_a_, outer lamina of wall of compartment, whence the
radiating, longitudinal septa (left unshaded) arise, and at
the opposite end blend into the indistinct inner lamina,
viz. _b_, (see fig. 4, in last plate.)
_c′_, sheath.
_d_, ala, forming, as usual, part of the sheath.
_e_, inner portion of the radius.
_f_, _f_, outer lamina of the radius (see _b_, in fig. 5,
in last plate), of great thickness, and externally deeply
pitted;--the sharp ridges between the pits, produce in the
section the points, such as that marked by the outer _f_.
2, Coronula balænaris, shell seen from above.
2 _a_, " " section of one of the transverse
circumferential loops, formed by the
folded wall of a compartment. The wall
itself is formed of an outer and inner
lamina, with longitudinal septa. The
internal surfaces of the loop-part are
connected by special shelly plates or
longitudinal septa.
2 _b_, " " scutum and tergum united together by the
opercular membrane (with horny ridges),
seen from the inner side.
3, Coronula diadema, shell, seen from the outside; 3 _a_,
scutum and tergum joined together by the opercular
membrane, seen from the outside; 3 _b_, scutum cleaned and
enlarged, seen from the inside.
4, Section, in a vertical plane, through the skin of a whale,
on which a _Coronula diadema_ had been attached, but has
been wholly removed: the two curved, horn-like projections
occupied two of the eighteen cavities on the under side of
the shell, formed by the folded walls: the blacker part is
the epidermis; the lighter part is the yellowish fibrous
tissue of the skin under the epidermis.
5, Coronula reginæ, shell seen from the outside.
6, Coronula barbara, internal view of the basal margin and
inside of one of the compartments, exhibiting the
circumferential transversely looped ends of the folds of
the wall, with the inner surface transversely wrinkled.
7-9, diagrams, showing how one of the circumferential transverse
loops of the wall becomes divided into two transverse
loops, thus giving rise to another fold in the wall.
10, diagrams, showing how the wall of the young shell in
Coronula, from being (_a_) simply sinuous, becomes deeply
folded (_b_); the folds lastly (_c_) expanding transversely
at their ends, thus giving rise to the circumferential
transverse loops, as in fig. 7.
PLATE 16.
_Genus_--CORONULA.
Fig. 1, rostrum, viewed from the inner side, of _Coronula
diadema_.
_a_, sheath, marked transversely in the upper part by the
attachment of the opercular membrane.
_b_, furrow on each side of (_a_), receiving the edges of the
thick alæ of the adjoining lateral compartments.
_c′′_, special plate, on which the alæ on their outer sides
rest.
_d_, radius, on the edge it may be just seen to consist of an
outer layer (the normal radius), and a much thicker inner
part (the pseudo or complementary radius) formed of oblique
denticulated septa.
_e e′ e′′_, basal edge of wall, which from its commencement
at _e_, or _e′′_ can be followed, folding up to near the
basal edge of the sheath, to its termination at _e′′_ or
_e_.
_f_, serrated lines of junction between the folds of the wall.
2, Lateral compartment of _C. diadema_, seen laterally, on a
smaller scale than last fig., but taken from the same
shell; letters of reference the same: this figure, if the
ala _a′_ were removed, would do for a lateral view of the
rostrum, or fig. 1.
_a_, sheath, much foreshortened.
_a′_, ala (therefore also part of sheath), and the edge of
which fits into _b_ of fig. 1.
_b_, furrow receiving edge of ala of the adjoining
carino-lateral compartment.
_c′′_, special plate, seen edgeways.
_d_, radius, the division into two parts, viz., the thin
outer normal radius, and the under pseudo or complementary
radius here shown (rather exaggerated) plainer than in fig.
1.
_e_, basal edge of wall; to the right hand the three folds at
their inner ends are seen obliquely, one behind the other:
these are seen directly in front in fig. 1.
3, lateral compartment of _C. balænaris_, seen laterally;
letters of reference the same as in last fig., but _c′′_
not introduced for this plate is barely developed, and
only in the lower part, and is attached to the radius: _g
g_ points where fresh folds have been formed in the walls
along the lines of suture, as may be seen in the outline of
the wall in fig. 5.
4, lateral compartment of _C. reginæ_, seen laterally; letters
of reference the same as in the last two figures.
5, Coronula balænaris, outline of the basal margin of the
folded walls: _s_, _s_, the six sutures. The wall can be
continuously followed from one end of a suture to the next
suture.
6, Coronula diadema; small portion of the external surface of
the wall, close to the basal edge, highly magnified; _a_,
outer lamina, not extending down to the basal edge; _b_,
inner lamina; _c_, projecting longitudinal septa.
7, Coronula diadema; transverse section through the upper part
of the shell, showing the rostrum and the two adjoining
lateral compartments (and the alæ of the carino-lateral
compartments), all supposed to be a little separated from
each other, so that the sutures, _s s_, are shown by white
lines.
A A A, rostrum.
C C, lateral compartments.
D D, carino-lateral compartments, of which only the alæ
(_a′_) are shown.
Of the letters in italics, those which occur in figs. 1, 2, 3,
4, 5, refer all to the same parts, viz.:--
_a_, sheath, continuous with (_a′_) the alæ, where such occur.
_c′′_, special plate on which the alæ rest.
_d_, radius, divided into a thin outer normal radius, and the
inner complementary portion, the latter shaded by convex
lines of growth.
_d′_ is a special tube (seen only as a black dot) for a
thread of corium.
_e_, folded walls.
_f_, junction of the folded walls (see _f_, in fig. 1)
leading into the cavities (_f′_) open beneath, and filled
up by the epidermis (see Pl. 15, fig. 4) of the whale.
_h_, inside of the transverse loops of wall, occupied by
corium: in the enlarged section of a loop, in C. balænaris,
in Pl. 15, fig. 2 _a_, the opposed sides are seen to be
connected by shelly longitudinal plates.
_v_, cavity occupied by the ovarian tubes and cæca.
PLATE 17.
_Genera_--PLATYLEPAS, TUBICINELLA, AND XENOBALANUS.
Fig. 1 _a_, Platylepas bissexlobata, shell of, viewed from the
under side; A, rostrum; B, lateral compartment; C,
carino-lateral compartment; E, carina. These letters are
placed opposite the inward folds of each compartment
forming the midribs.
1 _b_, Platylepas bissexlobata, shell of, viewed externally.
1 _c_, " " scutum and tergum, viewed
externally and separated.
1 _d_, " " lateral compartment, viewed
from the inside, showing the
inward folded wall or midrib;
_a′_, ala; _d_, radius.
2 _a_, Platylepas decorata, shell, viewed externally; A,
rostrum; B, lateral compartment; C,
carino-lateral compartment; E,
carina. These letters are placed
opposite the inward folds of each
compartment.
2 _b_, " " minute portion of external
surface of wall, magnified.
3 _a_, Tubicinella trachealis, natural size; _p_, tubular
soft lip leading into sack; _s_, scutum; _t_, tergum.
3 _b_, Tubicinella trachealis, carino-lateral compartment of
a young specimen, natural size; _e_, broken summit of wall;
_d_, radius: the dotted lines, connecting the two figures,
show the size and form which this compartment would have
attained, if it could have grown without the summit
constantly breaking away.
3 _c_, Tubicinella trachealis, scutum and tergum viewed from
within, and connected by layers of thickened membrane; the
scutum is to the left-hand.
4 _a_, Xenobalanus globicipitis, enlarged; _b_, skin of the
porpoise, with the shell imbedded; at _c_ the lower end of
the prosoma of the included animal's body is supposed to be
seen through.
4 _b_, Xenobalanus globicipitis, shell greatly enlarged, seen
from above, with the basal membrane at the bottom; the
peduncle-formed body having been removed.
A, rostrum, on the inner side of this and the adjoining
compartments, the transversely ribbed sheath can be seen.
B, lateral compartment.
C, carino-lateral compartment.
E, carina; these compartments are separated by the sutures
_s s_, which run from the top of the shell to the bottom,
along the ends of the six rays formed by the adjoining
portions of the inwardly folded compartments.
_a_, alæ, of rectangular shape.
_d_, pseudo, or complimentary radii.
4 _c_, Xenobalanus globicipitis, small portion of external
surface of wall of shell, showing the external lamina,
_a a_, which consists only of narrow ledges (expanding
and contracting) on each side of the longitudinal septa,
_c_; _b_, the internal lamina.
PLATE 18.
_Genus_--CHTHAMALUS.
Fig. 1 _a_, Chthamalus stellatus, shell, _var. a_ (communis).
1 _b_, " " " " _e_ (depressus).
1 _c_, " " " " _c_ (communis).
1 _d_, " " " " _d_ (fragilis).
1 _e_, " " scutum and tergum, viewed inside,
_var. c_, British.
1 _f_, " " scutum and tergum, viewed inside,
_var. a_, Madeira.
1 _g_, " " scutum and tergum, viewed inside,
_var. e_, Mediterranean.
1 _h_, " " scutum and tergum, viewed inside,
_var. c_ and _e_, St. Jago,
Cape de Verde Arch.
2, Chthamalus antennatus, shell.
3 _a_, Chthamalus dentatus, shell; 3 _b_, _var._ shell,
attached to a ship's bottom; 3 _c_, tergum, viewed from
within.
4 _a_, Chthamalus cirratus, scutum, viewed on inside; 4 _b_,
tergum, viewed on inside.
5 _a_, Chthamalus Hembeli, shell, old specimen.
5 _b_, Chthamalus Hembeli, younger shell; C, carino-lateral
compartment; _a′_, ala; _d_, radius; E, carina; _a′_, the
two alæ.
5 _c_, Chthamalus Hembeli, scutum, external view of; 5 _d_,
tergum, external view of; _e_, two of the crests, to which
the tergal depressor muscles are attached, much enlarged
and viewed from vertically beneath, showing the lateral
denticuli or sub-crests.
6 _a_, Chthamalus fissus, shell; 6 _b_, scutum and tergum,
separated, internal views of.
PLATE 19.
_Genera_--CHTHAMALUS, CHAMÆSIPHO, AND PACHYLASMA.
Fig. 1 _a_, Chthamalus intertextus, shell; 1 _b_, scutum and
tergum, calcified together, with the suture obliterated in
the upper part, internal view of.
2 _a_, Chthamalus scabrosus, shell; 2 _b_, scutum, internal
view of; 2 _c_, tergum, internal view, _p_, plate forming
one side of the cavity in which the depressor muscle is
attached; 2 _d_, tergum, as seen from almost vertically
beneath; _p_, pit for the depressor muscle; _q_, special
pit for the lateral, properly scutal depressor muscle; _r_,
articular furrow.
3 _a_, Chamæsipho columna, shell; A, rostrum; B, lateral
compartment, probably formed by fusion of the
rostro-lateral and lateral compartments; _E_, carina;
3 _b_, 3 _c_ scutum and tergum, internal views of.
4 _a_, Chamæsipho scutelliformis, shell much enlarged, seen
from above; A, rostrum, separated by imperfect sutures from
the (B) lateral compartments, which are formed probably by
the fusion of the rostro-lateral and lateral compartments;
E, carina; B stands opposite one of the inward folds of
the wall, leading into one of the tubular apertures, which
runs (generally with an irregular spiral curve) through
the whole thickness of the shell down to the surface of
attachment. The carina has two of these apertures.
4 _b_, Chamæsipho scutelliformis, another specimen, seen from
the under side, letters of reference the same.
4 _c_, Chamæsipho scutelliformis, rostrum, disarticulated,
enlarged, showing the alæ.
4 _d_, Chamæsipho scutelliformis, scutum and tergum, internal
views of.
5 _a_, Pachylasma giganteum; 5 _b_, rostrum of an extremely
young specimen, much enlarged, showing its compounded
nature, viz., of a true rostrum and rostro-lateral
compartments; 5 _c_, scutum and tergum, external views of;
5 _d_, scutum and tergum, internal views of.
PLATE 20.
_Genera_--PACHYLASMA, OCTOMERIS, AND CATOPHRAGMUS.
Fig. 1 _a_, Pachylasma aurantiacum, shell, lateral view; A,
rostrum, separated only by the finest suture from B,
the rostro-lateral compartment, which latter has a mere
rim-like radius on the side facing the lateral (C)
compartment. The latter (C) has a great ala, and is
separated from the (D) carino-lateral compartment, by the
finest suture; D has a rim-like radius; E, carina, with
very large alæ (_a_).
1 _b_, Pachylasma aurantiacum, inside view of about half the
shell, showing the compound rostrum, consisting of the true
rostrum (A) and the two rostro-lateral (B, B) compartments;
of the latter, that to the left has a shoulder, receiving
the ala of the lateral compartment which has been removed;
the ala of the lateral compartment extends, in the
upper part, over the whole width of the rostro-lateral
compartment, as may be seen on the right-hand, B; C,
lateral compartment; D, carino-lateral compartment, with a
shoulder (_b_) to receive the ala of the carina, which has
been removed.
1 _c_, 1 _d_, Pachylasma aurantiacum, scutum and tergum,
external views of.
2 _a_, Octomeris angulosa, shell seen from above; 2 _b_,
scutum and tergum, internal views of.
3 _a_, Octomeris brunnea, shell seen from above; 3 _b_,
scutum and tergum, external views of.
4 _a_, Catophragmus polymerus, shell seen from above.
4 _b_, " " external view of one of the
scales or valves, from the second
whorl, counting from the inside.
4 _c_, " " outline of the basal edges of all
the valves of a shell; A,
rostrum; B, rostro-lateral; C,
lateral; D, carino-lateral
compartments; E, carina.
4 _d_, " " external view of a portion of the
shell of a much corroded and worn
down specimen, serving almost as
a transverse section; C, lateral
compartment, D, carino-lateral
compartment.
4 _e_, " " scutum and tergum, internal views
of.
PLATE 21.
_Genus_--VERRUCA.
N.B.--_Letters of reference the same throughout the Plate._
A, rostrum.
B, carina.
S, moveable scutum, S′, scutum fixed and modified so as to
form part of the shell.
T, moveable tergum, T′, tergum fixed and modified so as to
form part of the shell.
In S, and S′, _a_ is the occludent margin; _b_, the basal
margin; _m_, the plate to which the adductor muscle is
fixed; _o_, a ledge formed during continued growth, in
the upper part of the fixed scutum, in order to keep the
orifice perfectly closed; this ledge is seen distinctly
only in fig. 5.
In S and S′ the tergal margin is marked by small dashes; (′)
being the upper articular ridge, and (′′), the second or
lower articular ridge: in S′ (′′) is called the parietal
portion of the valve.
In T and T′, the scutal margin is marked by small dashes; (′)
being the first and upper articular ridge, hardly distinct
from the occludent margin, and called in T′′ the occludent
rim; (′′) is the second or middle, and (′′′) the lower or
third articular ridge, called in T′′ the parietal portion
of the valve; _x_ is the carinal margin, called in T′ the
carinal rim, and _z_ the basal margin; _o_ is the occludent
margin: in T′, a ledge is formed across the under side
close to the apex, which ledge is necessary to keep the
orifice neatly closed, owing to the apex of the moveable
tergum coming to project freely during growth.
Fig. 1 _a_, Verruca Strömia, specimen with the left hand scutum
and tergum fixed and modified into part of the shell.
1 _b_, Verruca Strömia, (with left hand scutum and tergum
fixed), with the valves and compartments separated from
each other; the homologous parts of the opercular valves
are marked by corresponding letters and dashes.
1 _c_, Verruca Strömia, left hand scutum and tergum fixed,
viewed from the under side.
1 _d_, Verruca Strömia, _smooth var._, reversed specimen, _i.
e._, with the right hand scutum and tergum fixed and
modified into part of the shell.
1 _e_, Verruca Strömia, shell seen from the under side; the
right hand scutum and tergum (as in 1 _d_) being the fixed
pair: the moveable scutum and tergum (S, T) are seen in the
shade at the bottom of the shell, and their shape will be
best understood by looking at fig. 1 _f_. The fixed scutum
and tergum S′, T′, differ a little in shape, and in the
form of their line of junction, from the same two valves
(reversed) in fig. 1 _c_.
1 _f_, Verruca Strömia, moveable scutum and tergum, seen from
the under side, taken from a specimen, in which (as in 1
_d_ and _e_) the right hand scutum and tergum were the
fixed pair.
2, Verruca Spengleri, scutum seen from the under side, showing
the medial, prominent adductor ridge; taken from a
specimen, in which the right hand scutum and tergum were
fixed.
3 _a_, Verruca lævigata, scutum and tergum, external view of,
from a specimen in which the left hand pair was fixed; 3
_b_, scutum and tergum, internal view of, from a specimen,
in which the right hand pair was fixed.
4, Verruca prisca, scutum and tergum, external view of, from a
specimen, in which left hand pair was fixed.
5, Verruca nexa, with the valves separated, the left hand
scutum and tergum being the fixed and modified pair.
6, Portion of ribbed shell of a Venus, to which a _Verruca
Spengleri_ had been attached, showing the peculiar form of
the excavation.
PLATE 22.
ALCIPPE LAMPAS.
Fig. 1, Entire animal (female and two males) greatly magnified,
being an unusually symmetrical specimen, partly copied from
Mr. Hancock's Plate ('Annal. and Mag. Nat. Hist.' ser. 2,
vol. 4, Pl. VIII). H, horny disc and surface of attachment;
_a_, projection formed by the lower end of the lip of the
orifice leading into the sack; _m_, a pair of males, of
their proper proportional sizes, attached in their ordinary
position.
2, Entire animal, much distorted.
3, Small portion of a Fusus (copied from Mr. Hancock's Plate),
perforated by the Alcippe; the darker curved marks are the
slit-like orifices, leading into the chambers; the latter
are seen from the outside, owing to a difference of tint in
the shell of the mollusc where worn thin over the chambers;
they are represented by the fan-shaped shaded patches.
4, One of the orifices, leading into a chamber, much enlarged;
_a_, the curved narrow end, which was open and used when
the animal was young, but has since been closed externally
by sand or shelly matter, and internally by the upward
prolongation of the horny disc; _b_, rims of an inorganic,
calcareous deposit, by which the narrow end of the orifice
is kept of the due degree of narrowness.
5, Longitudinal section through the outer envelopes of a very
symmetrical specimen, giving a lateral view of the included
body.
_a_, point at the lower end of the orifice of the capitulum,
leading into the sack.
_b_, end of adductor muscle.
_c_, mass of branching ovarian cæca, much developed and
protuberant on the under side.
_d_, basal point of the quasi-peduncle, projecting beyond the
level of the horny disc (H, H).
_e_, sack or open cavity: of the two branchiæ or ovigerous
fræna, one has been removed with the outer envelopes, the
other is hidden by the projection caused by the medial
distended mass of the ovarian cæca.
_e′_, lateral line of junction of the body to the outer
envelopes; which latter have been here cut through in
removing the near half of the capitulum and peduncle.
_f_, notch separating the capitulum or upper part from the
peduncle or lower part of the external covering: this notch
varies much in depth.
_g_, the end (homologically the carinal end), of the orifice
leading into the sack, where the cutting of the outer
envelopes has commenced.
H H, horny disc, cut longitudinally down the middle.
_h_, first pair of cirri.
_i_, prosoma (homologically the second thoracic segment).
_k_, the thoracic segment, which would have borne the second
pair of cirri, had such existed.
_l_, thoracic segment, which should have borne the third pair.
_m_, thoracic segment, which should have borne the fourth
pair.
_n_, thoracic segment, very small, bearing the fifth pair of
cirri.
_n′_, fifth pair of cirri, only one cirrus on the near side
being represented.
_o′_, sixth cirrus, borne on the last thoracic segment, too
small to be shown. _p_, caudal appendages.
6, one side or lip of the orifice leading into the sack,
greatly enlarged, seen on the inner side, formed of an
inner membrane, _b_, (on which the long hairs and an S-like
band of spines, too fine to be plainly represented, are
placed), and an outer membrane _c_, studded with short,
thick spines, the corium between these two membranes having
been removed; _a_, projection at lower end of orifice;
_g_, upper end of orifice, showing the point where the
corresponding side or lip of the orifice has been cut away.
7, small portion of the external membrane, showing the
star-shaped, hard, projecting points of chitine; but they
are here placed too closely together.
8, front view of mouth, greatly enlarged, and with the
gnathites rather unnaturally separated from each other;
_h_, the first pair of cirri; in front, the outer maxillæ
appear like a bilobed lower lip; the inner maxillæ (with
their singular membranous swelling behind, see fig. 15),
can be distinguished by their long apodemes or horny
imbedded bars; behind are seen the one-toothed mandibles,
with a swelling behind, probably representing the palpi;
all above the mandibles consists of the immensely developed
labrum, with the foreshortened ends of the row of long
hairs on each side; the fold, or articulation, separating
the mouth and body, is seen crossing above the basal ends
of the apodemes of the maxillæ.
9, Lateral view of the coriaceous button or cushion, _c′_, on
one corner of the upper segment, _b_, of the pedicel of
the sixth cirrus; _c_, being the lower part of the lower
segment of the one ramus of this same cirrus: the hooked
hairs are rather distorted.
10, The same, viewed from the inner side.
11, Lateral view of the labrum, with the mandible, _m_,
attached to it, which latter, from overhanging the
œsophagus, shows the position of the mouth; _h h_, is the
first cirrus on the near side; _b_, is the medial crest of
labrum, on the side of which extends a long row of hairs;
_a a a_, delicate membrane of side of body, attached to the
margins of the labrum.
12, Diagram showing the probable manner in which the young
Alcippe bores into the shell of molluscs; _a_, pupa
attached by the antennæ to the outer surface of shell; _b_,
outline of young Alcippe soon after its metamorphosis; the
anterior or lower end has increased considerably in length,
so as to project beyond the point whence the antennæ
rise, and it has now penetrated the shell, being attached
to the roof of its incipient chamber by its horny disc,
represented by a thick black line; _c_, is the Alcippe
after further growth, when it has succeeded in burying
itself; the horny disc is now attached parallel to the
surface of the shell; the horny disc of its former state,
now lines the narrow end of the slit-like orifice leading
into the shelly chamber: the above changes in position
are supposed to have been effected quite gradually. The
diagram, _c_, I may add, represents the position of the
Alcippe for the rest of its life, the chamber being added
to at both ends, sides, and bottom.
13, Lateral view of the posterior extremity of the thorax, much
enlarged, with its articulated appendages represented only
on one side; the four thoracic segments, _l_, _m_, _n_,
_o_, correspond with those represented in fig. 5. In one
monstrous specimen, segment _n_, bore a single cirrus. The
posterior thoracic segment, _o_, bearing the sixth pair of
cirri, _o′_, is very small and obscure, and can be seen
only by separating the fifth and sixth pairs of cirri;
or by longitudinally bisecting the thorax, and viewing
the inner side; _n′_, fifth cirrus; _a_, lower segment,
_b_, upper segment of pedicel; _c_, lower, and _d_, upper
segment of the one ramus, the other ramus being represented
by the coriaceous boss _c′_; _o′_, sixth cirrus, with
similar segments; _p_, caudal appendage.
14, First cirrus.
15, Maxillæ, with the lower end of the apodeme cut off, showing
the curious membranous swelling on the side towards the
mandible.
PLATE 23.
_Genera_--ALCIPPE AND CRYPTOPHIALUS.
ALCIPPE LAMPAS.
Fig. 16, Pupa of male Alcippe lampas, viewed laterally; _a_,
antennæ, _c′_, eyes attached to the apodemes; above these
the single eye of the mature male may be distinguished.
17, Abdomen of the same; _p_, posterior end of thorax; _q_,
abdomen; _r_, caudal appendages.
18, Male of Alcippe lampas immediately after the metamorphosis,
the ventral surface being uppermost, but the specimen has
been in some way distorted, as the lobe (_i_) ought to
project on the opposite side; _a_, antennæ of the pupa;
_g_, _h_, lateral lobes; internal organs not yet formed;
orifice at upper end (_i_) not visible; scale same as for
the pupa, fig. 16.
19, Male of Alcippe lampas, when fully mature; scale same as in
last figure (18) and as in fig. 16.
_a_, antennæ of pupa.
_b_, vesicula seminalis.
_c_, _eye_, (seen in fig. 16 above the eyes of the pupa,
_c′_.)
_d_, testis.
_e_, lower transverse muscle.
_f_, upper transverse muscle.
_g_, _h_, lateral lobes of the external covering, answering
to the sides of the peduncle of the female.
_i_, terminal lobe on the ventral side of the orifice,
probably corresponding with _a_ in fig. 1, Pl. 22.
_k_, orifice of the tubular sack.
_l_, oblique muscles on the ventral side of the sack.
_m_, probosciformed penis, shown exserted; _m′_ portion
within the tubular sack.
CRYPTOPHIALUS MINUTUS.
Fig. 1, Cryptophialus minutus (female with an attached male)
viewed laterally, but with the disc (H) rather turned
towards the beholder, much enlarged; _a_, orifice leading
into the sack; H, thin horny disc by which the animal is
attached within its cavity; H′, upper free worn edge of the
disc; _z_, male attached in its usual position to the edge
of the disc of the female.
2, Cryptophialus minutus, natural size, within a half-inch
circle; this represents the largest specimen which I have
ever seen.
3, Half the orifice, with a large portion of the external
membrane and of the membrane lining the sack.
_a_, _a_, dentated horny rim of orifice.
_b_, _b_, external membrane, with the underlying corium or
true skin removed.
_b′_, a horny bar, expanding at its lower end into a toothed
plate, and at the upper end connected with the horny rim
round the orifice.
_c_, _c_, membrane lining the sack.
_c′_, horny bar by which this membrane is strengthened.
_c′′_, thickened membrane, or bar, expanding into fibrous
sheets for the attachment of a muscle.
_d_, delicate lateral lip, within the outer dentated horny
rim.
4, Orifice of sack, cut off, widely open, viewed from above;
_a_, horny dentated rim; _b′_, top of horny bar, where
united to the rim; _d_, inner lateral lip folded backward;
_d′_, lip, with coarser hairs, at the carinal end of the
orifice.
5, Animal with the outer tunics removed, viewed laterally.
_a_, dentated horny rim of orifice, continuous with
_b_, outer membrane, here cut off.
_c_, inner membrane of sack, continuous with
_c_, _c_, the reflexed membrane of the body, by which the
body is united to the disc and outer envelopes.
_e_, lancet-shaped labrum.
_e′_, projection behind the labrum.
_f_, palpi.
_g_, outer maxillæ, between which and the palpi lie the inner
maxillæ and mandibles; these latter are rather exaggerated
in size.
_h_, first maxilliped, in a rudimentary condition, close
above which is the articulation separating the whole mouth
(_o_) from the first (^1) segment of the body.
_i_, _k_, tapering appendages springing from the second and
third segments of the body.
_l_, pedicels of the three pairs of abdominal cirri.
_m_, cirri, three pairs of.
_o_, mouth, of great size, formed by the confluence of the
lower segments of the gnathites.
^1, first segment of body, homologically the last or seventh
cephalic segment.
^2, second segment of body, homologically the first thoracic
segment.
^3, third segment of body, homologically the second thoracic
segment.
^4, fourth segment of body, homologically the third thoracic
segment.
^5, fifth segment of body, homologically the fourth thoracic
segment.
^6, sixth segment of body, homologically the fifth thoracic
segment.
^7, seventh segment of body, homologically the sixth thoracic
segment.
^8, eighth segment of the body, homologically the seventh
thoracic segment.
6, Front of mouth; _g_, outer maxillæ; _g′_, an articulation
separating the mouth from the membrane of the first segment
of the body; _h_, first pair of maxillipeds; whole figure
on same scale with the labrum, fig. 9.
7, Maxillæ, drawn on thrice the scale as the outer maxillæ;
_a_, apodeme.
8, Mandibles, drawn on thrice the scale as the outer maxillæ.
9, Labrum, on same scale with the outer maxillæ, fig. 6, and
one third of the scale of the mandibles and inner maxillæ;
_e_, labrum; _f_, palpi.
PLATE 24.
_Genera_--CRYPTOPHIALUS AND PROTEOLEPAS.
CRYPTOPHIALUS MINUTUS.
Fig. 10, Lower enlarged end of œsophagus, where entering the
stomach, viewed from one of the sides bearing the discs
of teeth; _a_, upper cut off end of œsophagus; _b_, the
lower end and natural opening; _c_, thickened rim, having a
somewhat beaded structure.
11, The same; diagram giving a transverse section of the
above in the line of the discs of teeth; _c c c c_, four
thickened beaded edges; _e e_, two other but narrower and
less prominent beaded edges; _d d_, discs of teeth; _f_,
rows of fine bristles; the almost double row of bristles on
each side under (_d_) is not of course seen in this section.
12, Lower end of œsophagus viewed from a position at right
angles to that in fig. 10; or from the side _e_, of the
diagram, fig. 11; the two discs of teeth are consequently
shown in profile; _a_, upper cut off end of œsophagus; _b_,
lower natural end; _c_, _c_, thickened edges; _e_, narrower
thickened edge, projecting towards the beholder; _d_, disc
of teeth seen in profile.
13, Portion of one of the cirri; _a_, lower segment of pedicel;
_b_, upper segment, supporting the two rami.
14, Segments of one of the rami much enlarged; _a_, thickened
shield-like portion of membrane.
15, Ovum, (thirty-five times the natural size.)
16, Egg-like larva in the first stage, on same scale.
17, " " in the second stage, on same scale.
18, Larva in the last or pupal stage, on four times the scale
of fig. 15-17; _a_, antennæ; _b_, apodeme and eyes; _c_,
abdominal bristles. A male would have been developed from
this pupa.
19, Male on same scale as the pupa, fig. 18; _a_, three
terminal segments of the antennæ of the pupa; _d_, orifice
of the sack.
PROTEOLEPAS BIVINCTA.
Fig. 1, Antenna of the pupa (three terminal segments of), with
a portion of one of the two threads, enclosing the
cement-ducts, by which the body of the Proteolepas is
attached; the upper portion of the thread, and the
adjoining part of the body, are represented in section.
_a_, part of the great mass of cellular matter within the
ovarian sack, in process of development into ova, and
changing its character as soon as it enters the tube or
cement-duct.
_b_, membrane forming the ovarian sack and the cement-ducts,
the latter enclosed within the threads of attachment.
_c_, corium lining the outer membrane of the body, and the
upper part of the threads.
_d_, outer membrane of body.
_e_, " " becoming suddenly thicker where
forming the outer membrane of the thread.
_e′_, outer membrane of the thread in the lower part, here
not shown in section.
_f_, main or second segment of the pupal antenna, the basal
segment having, as in all cases, been moulted, with the
carapace of the pupa.
_g_, disc-segment, apparently with a small orifice for the
issuing of the cement.
_h_, terminal segment with the shorter spines broken off.
2, Compounded mandibular organ, sketched by the camera; _a_,
supposed mandible; _b_, perhaps portion of the mandible;
_c_, maxilla; _d_, ligamentous fibres giving attachment to
muscles.
3, Mouth seen from the ventral side, sketched by the camera;
_r_, articulation separating the mouth from the body;
_c_, compounded mandibular organ; _d_, palpus, united to
the opposite palpus and to the crest of the labrum; the
latter forming the back of the hollow in which the compound
mandibles work.
4, Diagram of the mouth of an ordinary cirripede, seen from
above; _a_, outer maxilla; _b_, maxilla; _c_, mandible;
_d_, labrum.
5, Diagram, illustrating the supposed changes in _position_ of
the gnathites in Proteolepas, causing them to stand back
to back; _a_, outer maxilla; _b_, maxilla; _c_, mandibles;
_d_, labrum.
PLATE 25.
_Genera_--PROTEOLEPAS AND BALANUS.
PROTEOLEPAS BIVINCTA.
Fig. 6, Proteolepas bivincta: diagram, showing the probable
position of the young animal, just before its
metamorphosis, within the carapace of the supposed pupa;
_a_, caudal appendages; _b_, six pairs of natatory thoracic
legs; _c_, mouth, no doubt closed and functionless, as in
other pupæ; _g_, threads of attachment, with cement-ducts
in process of formation; _h_, antennæ.
7, Proteolepas bivincta, magnified about twenty-six times.
_m_, mouth, the summit being formed of the labrum and palpi
joined.
_^{1}c_, first segment of body; homologically the seventh or
last cephalic segment.
_^{2}t_ to _^{8}t_, second to eighth segment of body;
homologically, first to seventh thoracic segments.
_^{9}a_ to _^{11}a_, ninth to eleventh segment of body;
homologically, three segments of abdomen.
_d′_, three muscles attached on each side to the labrum, and
running to the gnathites.
_e e_, great ovarian sack.
_f_, true ovaria.
_g_, threads of attachment.
_h_, three terminal segments of the antennæ of the pupa.
_i_, vesicula seminalis.
_k_, papilla representing the penis.
Fig. 1, _Balanus tintinnabulum_: an enlarged longitudinal section
through the shell and sack, with the right-hand scutum and
tergum and right-hand half of shell and basis removed,
exhibiting the body of the animal not in section. The cirri
are exhibited only on one side.
A, A, orifice of shell, within which lies the operculum
formed by a pair of scuta (S), and pair of terga (T).
B, basis (homologically the anterior end of the shell).
K, carina of shell (or dorsal valve or compartment of shell).
K′, sheath of carina.
L, lateral compartment of shell. The carino-lateral
compartment is hidden by the scutum and tergum.
R, rostrum of shell (or ventral valve or compartment of
shell).
R′, sheath of rostrum.
O, O, opercular membrane, connecting the opercular valves
with the overhanging basal edge of the sheath.
S, scutum.
T, tergum.
_a_, adductor scutorum muscle, with the scutum on the near
side removed.
_b_, the whole space enclosed by a broken sinuous line, round
_a_ and _b_, shows the cut surface of attachment to the
near scutum, which has been removed.
_b′_, lower muscle, on the near side, running from near the
basal edge of the scutum to near the basal margin of the
labrum (_e_). Above this are three other muscles (all on
the near side), running to the membrane between the labrum
and adductor muscle.
_c_, prosoma, including the main portion of the stomach and
thickened ends of the vesiculæ seminales: homologically
this is formed by the development of the second thoracic
segment, which carries the first pair of cirri; and
possibly, in part, by the antecedent segment, _i. e._ the
first thoracic segment.
_c′_, thorax, extending from the prosoma to the posterior
end of the body: the letter (_c′_) stands on the segment
supporting the third cirrus; homologically, this segment is
the fourth thoracic.
_d′_, orifice of the acoustic sack, above which is the basal
articulation of the first cirrus.
_e_, labrum, forming the back (_i. e._ anterior end) of the
mouth, with the transverse palpi obscurely seen on the
summit.
_f_, sack in which the animal lies, with the tunic lining
it, continuous with that investing the prosoma (_c_), and
lining the under sides of both opercular valves, but cut
off round (_b_) and (_a_) by the removal of the near
scutum. The branchia on the further side, which occupies
the position represented at fig. 3, is covered by the body
of the animal.
_g g_, ovarian, inosculating cæca, branching from the simple
duct (of which the near one of the pair is represented),
leading to the ovaria (not represented) within the body.
_h_, rostral depressor muscle of the scutum: the lateral
depressor muscle of the scutum is hidden by the body and by
the membranes of the sack.
_i_, carinal depressor muscle of the tergum.
_z_, antennæ (three terminal segments) of the pupa; I
distinctly saw these in this species, but they are here
represented considerably too large, even on the supposition
that a young shell had been drawn, and magnified about
twelve times.
2, Testes of _Balanus perforatus_, greatly magnified.
3, Branchia of _Bal. tintinnabulum_, on the same scale as in
fig. 1, and in its natural position. This drawing was made
by lifting up the body in fig. 1; the organ being thus
completely exposed over its whole interior surface; _a_ is
the basal end of the spur of the tergum.
PLATE 26.
STRUCTURE OF THE MOUTH AND THORAX.
N.B.--_The same letters of reference are used for the parts of the
mouth throughout this plate. All the organs here represented have
been cleaned by boiling in potash, and consist exclusively of the
external membrane._
Fig. 1, Mouth of _Balanus perforatus_, viewed from vertically
above, with the first pair of cirri _x x_ in their proper
position, cut off close above their basal articulations.
_a_, outer maxilla; _a′_, ditto, cut closely off.
_b_, maxilla.
_c_, mandibles.
_d_, palpus; _d′_ ditto, cut closely off.
_e_, labrum, crest of.
_x x_, first pair of cirri, cut off.
2, Supra-œsophageal cavity of the mouth of _Balanus
improvisus_, torn open, with the palpi, mandibles, and
maxillæ removed, exhibiting the inner face of the labrum,
laid flat, and the inner faces of the outer maxillæ; scale
as in fig. 1.
_a_, outer maxillæ; _a′′_, inner and lower lobe of ditto.
_d′_, point of attachment to the torn-off palpus.
_e_, labrum, crest off; _e′_, central notch, with graduated
teeth on both sides.
_f_, triangular thickened portion of the inner fold of the
labrum; _f′_, rib or bar of membrane, thickened to support
the surrounding very thin membrane of the supra-œsophageal
cavity.
_g_, opening of the œsophagus.
_h_, forked bar of thickened membrane, for same purpose as
_f′_.
3, Mouth of _Coronula balænaris_, divided vertically in a
transverse line, with the outer and inner maxillæ and
mandible on the left side removed, with the labrum, the
other mandible and both palpi viewed on the inside, and
with the œsophagus adherent in its proper position.
_c_, Mandible, upper free segment; _c^1_, second or middle
segment; _c^2_, supposed third or basal segment, forming
the basal margin of the mouth, but not separated by any
articulation from the labrum.
_d d_, palpi; _d′_, aperture, through which the inside of
this palpus can be seen, caused by the mandible having been
cut off; the membrane on the under side of this aperture
is thickened, and affords the chief support to the palpus;
_d′′_, thin membrane, apparently part of the palpus,
connecting the attached basal end of the palpus to the
externo-lateral surface of the mandible.
_e′_, central notch in the crest of the labrum; _e′′ e′′_,
basal margin of the labrum, to which the membrane of the
body is joined.
_f_, thickened portion of membrane, part of the inner fold of
the labrum, corresponding with _f_ in fig. 2.
_g_, orifice of the œsophagus, with the front part of the
supra-œsophageal cavity cut away; _g′_, lower, expanded, or
bell-shaped, end of the œsophagus, in its natural condition.
_i_, _k_, concavities for the attachment of muscles running
to the mandibles.
_m_, cut off edge of membrane forming the side of the mouth,
which cut edge can be followed up above the aperture _d′_,
where the upper basal end of the palpus has been cut off,
and so onwards by _g_, to the mandible on the opposite side
of the mouth.
4, Mouth of _Coronula balænaris_, viewed from the outside, in
front, with the inner maxilla, mandible, and palpus on
the left side of the figure, and nearly the whole labrum,
and the greater part of the palpus on the right side, all
cut away. The maxilla, mandible, and basal portion of the
palpus on the right hand, are pulled out of their proper
positions; for the toothed edge of the mandible ought to
have stood higher, and nearly in a line between and behind
the outer maxillæ, and then the basal margin _e′′_ of the
lateral portion of the labrum would be raised, and curling
round, would stand nearly where the letter _b′_ is now
placed.
_a_ _a_, outer maxilla, upper _free_ lobe; _a′′_, lower
and inner _free_ lobe; _a^1_, supposed second or middle
segment; _a^2_, supposed third or basal segment; _a′′′_,
basal articulation of mouth, separating it from the ventral
surface of the thorax.
_b_, inner maxilla; _b′_, apodeme of do.
_c_, mandible, upper free segment; _c^1_, second segment;
_c^2_, third and basal segment, but not separated by an
articulation from the labrum.
_d_, palpus; _d′′_, membrane uniting the basal end of the
palpus, where attached to the labrum, to the side of the
mandible.
_e′′_, basal margin of the labrum, on one side, displaced and
pulled down.
_n_, orifice of the olfactory pouch.
5, _Balanus amaryllis_: mandible, upper free segment, seen
from the side that faces the labrum; _p_, an arched line,
where the thickened membrane of the upper free part
terminates, and is united by thin membrane to the near side
of the palpus; _q_, ligamentous bands for the attachment of
muscles.
6, _Balanus amaryllis_: palpus, on the same scale as the
mandible, fig. 5, seen from the outside, so that the
further or under face is the side which, towards the right
hand, is united to the near face of the mandible, fig. 5;
_r_, long bristles, springing from a protuberance near
the extremity; these bristles, in many species, form a
long single row, parallel to the basal margin; _s_, upper
row of shorter bristles; _t_, row of very short bristles,
arising from the further side and curling over the crest of
the labrum.
7, _Balanus amaryllis_: maxilla on twice the scale of the
mandible and palpus; _b_, upper free segment; _b′_,
apodeme; _b^1_, thickened portion of membrane, perhaps
answering to the second segment of the mandibles; _b^2_,
thin membrane, extending down to the basal edge of the
mouth, possibly answering to the basal segment of the
mandibles; or perhaps the thickened membrane forming the
protuberance _c^2_ in fig. 3, may be considered as the
basal segment of the maxillæ.
_u_, step-formed projection at lower angle of maxilla.
8, Thorax of _Coronula diadema_: outer membrane of the five
posterior segments laid almost flat, and viewed externally
as a transparent object.
^2 to ^6, the second to the sixth cirrus, cut off a little
above their basal articulations.
_a_, basal curved end of the probosciformed penis.
_b_, anus; _b′_, the membrane surrounding the anus, probably
consisting of a rudiment of the abdomen.
_c_, rectum, _seen through on the under side_.
_d_, basal articulation of the sixth cirrus; _d′_, do. of the
fifth cirrus; _d^2_, do. of the second cirrus.
_e e′_, posterior thoracic segment, carrying the sixth pair
of cirri; the dorso-lateral portion _e_ is separated, in
this one segment, from the corresponding portion _e′_ by a
narrow slip of thinner membrane, which may be a part of the
abdomen let in.
_e^2 e^2_, thoracic segment, supporting the second pair of
cirri.
_f_, _f^3_, folds or articulations between the adjoining
segments.
_g_, swollen, punctured portions of membrane, not differing
homologically from the rest of the segments.
_h_, posterior portion of the prosoma, or the specially
enlarged part of the thorax; it carries the first pair of
cirri; homologically _h_ forms part of the second segment
of the thorax of the archetype Crustacean.
PLATE 27.
NERVOUS SYSTEM AND SENSES.
Fig. 1, Nervous system of _Coronula diadema_, seen from the
ventral surface.
A, infra-œsophageal ganglion; two pairs of rather small
nerves, arising from the dorsal surface, cannot be here
shown.
B, supra-œsophageal ganglion, double.
C, ophthalmic ganglion, single.
D D, eyes, as believed to exist.
_a_, three pairs of nerves, running to the gnathites and
olfactory pouches;
_a′_, nerve rising from the collar, running towards the
mandibles.
_b_, single medial nerve, running to the great transverse
muscle, extending across the upper part of the stomach.
_c_, collar-nerve or chord, uniting the infra- and
supra-œsophageal ganglions.
_d_, great splanchnic nerve, here spread laterally out, but
properly running along the sides of the upper part of the
prosoma, and therefore under the collar-nerve;
_d′_, plexus by which this nerve is connected with the
supra-splanchnic nerve, _e_.
_e_, supra-splanchnic nerve, rising from the collar, almost
in contact with the supra-œsophageal ganglion;
_e′_, small nerve rising from the end of the supra-œsophageal
ganglion, and running to the adductor and surrounding
muscles.
_f_, _f_, pair of great nerves (_antennular_), distributed
over the sack and shell.
_g_, chord (in appearance single), uniting the
supra-œsophageal ganglion with the first (C), or ophthalmic
ganglion.
_h_, small medial nerve, running to near the adductor muscle.
_i_, nerve supposed to run from the ophthalmic ganglion to
the eye.
_k_, small nerve, rising from the main ganglion (A), between
_r^1_ and _r^2_, or the nerves running to the first and
second pairs of cirri.
_œ_, position of the œsophagus.
_r^1_ nerve entering the first cirrus.
_r^2_, _r^3_, _r^4_, _r^5_, _r^6_, nerves entering the five
succeeding pairs of cirri.
_s_, nerve entering the probosciformed penis.
2, nervous system of _Balanus tintinnabulum_; letters of
reference as in fig. 1.
3, acoustic sack of _Balanus tintinnabulum_.
4, " " _Coronula diadema_.
5, eye of _Balanus tintinnabulum_; _i_, nerve coming from the
ophthalmic ganglion; D, eye itself.
PLATE 28.
CEMENTING APPARATUS.
Fig. 1 _a_, Basal membrane, with the cementing apparatus, of
_Coronula balænaris_; a small portion of the parietal
membrane, _p p p_, which coats the folded shelly walls, is
left adherent to the basal membrane.
_b_, the circumferential slip (shaded more darkly than the
rest to catch the eye), separating the basal from the
parietal membrane.
_c′ c′_, slips of basal membrane, formed at each period of
growth, and overlapping each other.
_s s s s s s_, the six sutures in the walls, separating the
six compartments, of which--
A is the basal margin of the rostrum,
C C that of the lateral compartments,
D D that of the carino-lateral compartments, and
E that of the carina.
_r_, _r_, rays or spokes of membrane, prolonged from the
circumferential slip, and running under the trebly folded
wall of each compartment, but here cut off; a similar ray
should run under each line of suture (_s_). These rays, at
their extremities, expand transversely; and the shape and
length of the rays may be judged of from the basal outline
of the folded walls given in Pl. 16, fig. 5.
The two cement-ducts, proceeding from each cement-gland,
debouch opposite the middle folds of the lateral (C C)
and carino-lateral (D D) compartments. The layers of
cement have been removed. N.B. There is one considerable
error in this figure, the two main trunks, connecting
the cement-glands, and meeting at the centre, have been
represented as forming a straight line, but in fact they
form a very open angle, as is correctly shown in fig. 1 _c_.
1 _b_, Diagram, representing a vertical section through a
portion of the basal and parietal membranes, with the
thickness of the membrane enormously exaggerated.
_z z z_, layers of cement, which, if the section had been
made in the line of the cement-ducts, would have been seen
proceeding out of these ducts, as is represented at (_t_),
where the section is supposed to have taken the above
course.
_c′ c′_, the slips of basal membrane.
_b_, the circumferential slip; beneath this the coarsely
dotted layer represents the cement, lately excreted, and
before it has acquired its proper transparent structureless
character, elsewhere represented by fine dots.
_p_, membrane externally coating the walls of the shell.
_t_, cement-ducts opening beneath the basal membrane.
1 _c_, The central portion of fig. 1 _a_, considerably
magnified.
_c′ c′_, slips of the basal membrane; but the shell, when
these were formed, was so young that the walls had not
acquired their folded structure; in the centre the
prehensile antennæ of the pupa may be obscurely seen.
_f_, the main cement-trunk, connecting the cement-glands.
_h_, a cement-gland, from which two cement-ducts proceed.
_a a_, cement-ducts (cut off), leading to opposite the middle
fold of the carino-lateral compartment.
_b b_, cement-ducts (cut off), leading to opposite the middle
fold of the lateral compartment.
_a′ b′_, a pair of cement-ducts, with their orifices opening
on the under side of one of the slips of basal membrane
(_c′_), at a point which once was opposite the middle of
the carino-lateral and lateral compartments. The orifices
of the other ducts, towards the centre, may be seen forming
straight lines.
2, Cement-ducts and one cement-gland of _Chelonobia patula_,
represented without the basal membrane, to which they
adhere: _f_, _f_, main cement-trunk; _g_, enlarged portion;
_h_, gland; _a_, _b_, the two ducts proceeding from this
gland, and bifurcating several times before debouching
on the under side of the basal membrane. Several other
cement-ducts, proceeding from other glands, are represented
just as they appeared under the microscope.
3, Cement-ducts and glands of _Tubicinella trachealis_,
represented without the basal membrane to which they
adhered: (_f_) (_f_), main cement-trunk, connecting the
several glands; _h_, cement-gland; _a_, cement-duct, with a
singular loop (_a′_) having two projections or rudimentary
branches; _b_, spur or rudiment of a second cement-duct;
_c_, third cement-duct.
4 _a_, Chain of cement-glands of _Balanus tintinnabulum_,
_with all the ducts removed_, _excepting those proceeding
from the last formed gland_, which latter correspond in age
with the last-formed zone of the shelly basis; the whole of
the basis having been removed by acid.
_f_, main cement-trunk connecting the glands.
_gh_, last-formed cement-gland.
_k_, _t_, two cement-ducts, proceeding from a great common
duct; one of these bifurcates at (_t_), and one branch
joins at (_t′_) the corresponding branch from the
corresponding gland.
_i i i_, circumferential duct, into which the ducts _k_, _k_,
_t_, _t_, _t′_ all enter.
_i′ i′_, branches proceeding from the circumferential duct,
which branch and sub-branch till they form a sheet (_z z_)
of cement-tissue on the outside of the basis of the shell.
4 _b_, two cement-glands of _Bal. tintinnabulum_ (this figure,
to match with 4 _a_, ought to have stood upside down),
taken from near the centre of the basis, greatly enlarged;
(_f f_), main trunk; (_g_), enlarged portion of the trunk;
(_h_), gland; _k_, _t_, two cement-ducts proceeding from a
common point, one of them (_t_) bifurcates, and gives off
a rudimentary branch, _t′_; _m_, a spur, or rudimentary
duct. The gland, _h_, has been pushed on one side, it
ought to lie over the enlarged portion (_g_). There is a
considerable difference between these two glands and that
(_gh_) represented in fig. 4 _a_; the neck of the gland in
the latter being elongated into a great common duct, and
the spur or rudimentary duct (_m_) being absent.
PLATE 29.
CIRRI AND LARVÆ, FIRST STAGES.
Fig. 1, Sixth cirrus of _Balanus amphitrite_ (_var._ cirratus),
showing the muscles.
_a_, _b_, flexor and extensor muscles, moving the upper
segment (_i_ to _k_) of the pedicel.
_c_ and _d_, flexor and extensor muscles, apparently moving
the lower segments of both rami, as a whole; the lower
articulations in these rami being confluent.
_e_ and _f_, flexor and extensor muscles extending up both
rami (those only in the near ramus being figured) to their
tips.
_g_ _g_, flexors of the separate segments in both rami.
_h_, basal articulation of lower segment of the pedicel.
_i_, upper articulation of lower segment of the pedicel.
_k_, upper articulation of upper segment of pedicel.
2, Fourth cirrus of _Acasta sulcata_.
_i_, upper articulation of the lower } _These two letters
segment of pedicel. } apply to all the
_k_, upper articulation of the upper } figures 1 to 6._
segment of pedicel. }
_l_, curved teeth on the pedicel.
_m_, " " on the segments of the anterior ramus.
3, Third cirrus of _Chthamalus antennatus_.
4, Second cirrus of _Balanus perforatus_, viewed exteriorly and
laterally.
5, Third cirrus of _Xenobalanus globicipitis_.
6, Sixth " " "
7, Spermatozoa (copied from Mr. C. Spence Bate, in 'Annals and
Mag. of Natural History' (S. 2), vol. viii, Pl. VIII).
_a_, of _Verruca Strömia_, in an early condition.
_b_, of _Balanus balanoides_, more mature.
_c_, of _Balanus perforatus_, apparently mature.
8, Larva of _Scalpellum vulgare_, immediately after coming out
of the egg, seen on the ventral surface (letters of
reference given below).
9, Larva of _Balanus balanoides_, immediately after coming out
of the egg, seen on the ventral surface, copied from Mr. C.
Spence Bate's drawing in 'Annals and Magazine of Natural
History' (S. 2), vol. viii, Plate VI, fig. 1 (letters of
reference given below).
10, Larva of _Chthamalus stellatus_, after the first moult, but
during the first stage, seen on the ventral surface,
copied; with some alterations, from Mr. C. Spence Bate's
drawing in 'Annals and Magazine of Natural History' (2
Ser.), vol. viii, Plate VIII, fig. 13.
(_The following letters apply to above figs. 8, 9, 10._)
_a_, eye.
_b_, first pair of antennæ, not observed in fig. 9.
_b′_, the same antennæ, as yet encased (in fig. 8) within
small horns.
_c_, horns, including the second pair of antennæ.
_d_, mouth, probosciformed: in fig. 8, the specimen having
been acted on by potash, the œsophagus can be seen within,
with the orifice beneath the swelling, which latter perhaps
answers to the labrum.
_e_, first, uniramous natatory leg (homologically the second
thoracic limb).
_f_ _g_, second and third, biramous natatory legs
(homologically the third and fourth thoracic limbs.)
_h_, posterior point of carapace.
_i_, forked terminal projection of the body or abdomen.
_l_, second forked projection.
_m_, two sharp points, apparently representing a third forked
projection.
_n_, a rounded swelling, apparently lying between the
carapace and the supposed abdomen, and believed to form the
anus.
PLATE 30.
LARVÆ OF LEPAS: SECOND AND LAST STAGES OF DEVELOPMENT.
Fig. 1, Larva of Lepas in the second stage of development; _a_,
supposed antennæ (second pair); _m_, mouth; _c_, three
pairs of legs. (Copied from Burmeister's 'Naturgeschichte
der Rankenfüsser,' Tab. 1, fig. 3.)
2, _Lepas australis_: pupa or larva (2 _a_, of natural size) in
the last stage of development, with the young cirripede in
its natural position, obscurely seen within,--the specimen
having been treated with caustic potash, _and so rendered
transparent_,--viewed laterally and greatly enlarged.
Scarcely more than the outline of the shell or carapace is
represented. The darkly shaded part to the left represents
the extent of the sack of the pupa, or the cavity occupied
by the thorax, with its limbs.
_a_, both antennæ, adhering by their discs to a piece of wood.
_b_, dorsal surface of the shell or carapace; and immediately
underneath this letter is the point of reflection of the
membrane investing the thorax, so as to line the sack of
the pupa.
_b′_, is the posterior end of the animal, and of the orifice
on the ventral surface, through which the legs are
protruded.
_c_, six pairs of natatory legs.
_d_, pair of caudal appendages, seated on a minute abdomen.
_m_, mouth, from which the œsophagus can be seen running into
the stomach, the latter having two dark cæca; the stomach
sweeps round in the pupa to the abdomen, near _b′_; but in
the young included cirripede, only as far as the letter
(_b_), where the bases of the posterior pair of cirri and
the anus lie.
_n_, apodemes, supporting the eyes, produced deeply inwards
from the eye-bearing segment (N, in fig. 4) of the antennæ.
_s_, bottom of sack of the young cirripede immediately after
its metamorphosis; it extends as far as _s′_. (See _s_, in
fig. 3.)
_t_, gut-formed cement-gland (or incipient ovaria), seen on
the near side of the animal, whence a cement-duct, _t′_,
runs into the near antenna.
_u_, internal and anterior part of the pupa, filled with
pulpy, oily matter, together with the incipient muscles of
the peduncle; when stretched out it forms the peduncle of
the young cirripede. See _u′_, in fig. 3.
_x_, bases of the pedicels of the cirri of the young included
cirripede.
2 _a_, Pupa of _Lepas australis_ of natural average size,
within a half-inch wide.
3, Young cirripede (on a smaller scale than Fig. 2),
immediately after the exuviation of the bivalve-like
pupal carapace, the basal segments of the antennæ, the
eye-apodemes and eyes. The young cirripede has just assumed
its proper position at nearly right angles to what it held
whilst packed within the pupa,--this change of position
having been effected by the opening out or stretching of
the deep fold of membrane (see _n_, in fig. 2) formed
over the eye-apodemes and eyes, previous to the act of
exuviation.
_a_, the three terminal segments of the antennæ of the pupa,
still remaining cemented, in the same position as before,
to the same piece of wood: the basal, or eye-bearing
segment (N, in fig. 4), has been moulted with the pupal
carapace.
_c_, _d_, legs and caudal appendages of the pupa, not as yet
moulted, but quite functionless; the external membrane of
the thorax, and that lining the sack of the pupa, are,
likewise, as yet retained, but soon will be cast off.
_s_, bottom of the sack of the young cirripede, which can now
be easily distinguished.
_u′_, the peduncle.
_x_, _y_, _z_, primordial valves, composed of chitine: _x_,
being the scutum; _y_, the tergum; _z_, the carina.
3 _a_, small portion of one of the primordial, non-calcified
valves, much magnified.
4, ventral surface of pupa; on the same scale, and in the same
semi-transparent condition as in fig. 2.
_a_, antennæ.
_b_, limit of sack on the sides in the ventral or lower half
of the pupa.
_b′_, posterior end of the carapace.
_c_, _d_, bristles of the natatory legs and of the caudal
appendages, protruded through the orifice of the carapace.
_e_, orifice of the acoustic sacks.
_m_, mouth, with the œsophagus attached to it, obscurely seen
through the carapace; the stomach having been removed.
_n_, apodemes supporting the eyes.
N, the eye-bearing or basal segments of the antennæ.
_o_, the second or main segment of the antennæ.
_p_, the third or disc segment of the antennæ.
_v_, the fourth or terminal segment of the antennæ.
W, ventral surface, bordered by the edges of the carapace;
the letter stands near the extremity of the œsophagus.
5, First pair of natatory legs: _f_, lower segment of pedicel;
_g_, upper segment of pedicel; _h_, lower segment of ramus;
_i_, upper segment of ramus; _k_, outer ramus; _l_, inner
ramus of same leg; _r_, sternal surface between the first
and second pairs of legs, with singular thickened ridges
and folds.
6, Abdomen with two caudal appendages: ^1, first abdominal
segment, attached to the posterior thoracic segment; ^2,
second abdominal segment; ^3, third or last abdominal
segment; _h_, lower segment; _i_, upper segment of caudal
appendage.
7, Transverse section of the pupa, close to the eye-apodemes;
these being made to stand more upright than in fig. 2. The
internal organs of the animal have all been removed.
_a a a′_, section of carapace; from _a_ to _a′_, on the lower
side, forms the ventral surface, with three longitudinal
furrows, here in some degree opened out.
_c_, crest of thick membrane on each side, forming the sides
of the lateral furrows.
N, the posterior margin of the eye-bearing segment (see fig.
4) of the antennæ; the edges of which are hardened and
thickened, and are produced inwards, forming
_n_, the apodemes, to which are attached the (_n′_) great
compound eyes.
8, Terminal portion of an antenna, greatly magnified, seen
from above.
_o_, part of the second or main segment; an oblique line
shows a line of separation of the upper thinner and lower
thicker membrane; _o′_, single spine borne at this segment.
_p_, disc-segment, bearing seven spines; _p′_, an irregular
border of the cement-tissue, believed to have debouched
through the spoke-like minute tubes seen on the disc.
_v_, terminal segment, projecting almost rectangularly
outwards; _v′_, most delicate tube or ribbon, believed to
consist of cement-tissue.
ERRATA.
Page
104, thirteen lines from bottom, _for_ "_Balanus elongatus_," _read_
"_Balanus galeatus_."
105, six lines from bottom, _for_ "_Balanus elongatus_," _read_
"_Balanus galeatus_."
174, Since the table on the distribution of the fossil species was
drawn up, I have examined some more specimens, sent me by Mr.
Wood, which show that _Balanus concavus_ and _Hameri_ are found
in the Red Crag; and that _Verruca Strömia_ is found in the
Coralline Crag; this will make the total in the Coralline Crag
ten, and in the Red Crag eight. I should, however, add, that
the identification of _Verruca Strömia_ in the two Crags is
a little doubtful, as the specimen was without the opercular
valves.
235, to _Fossil_ localities of _Bal. concavus_, add Red Crag
(Sutton) Mus. S. Wood.
255, six lines from the top, _Bal. crenatus_, I have now seen a
single Red Crag specimen .5 of inch in basal diameter.
278, to _Fossil_ localities of _Bal. Hameri_, add Red Crag (Sutton)
Mus. S. Wood.
293, nineteen lines from the top, _for_ "Cotantin," _read_
"Cotentin."
300, ten lines from bottom, _for_ "parieted" _read_ "parietal."
INDEX.
N.B. The names in _italics_ are synonyms or doubtful species.
Abdomen of the Balanidæ, 65.
of larvæ in first stage, 108.
in pupa of Alcippe, 548.
Abdominalia, order of, 21, 563.
Acasta, sub-genus, 302.
cyathus, 312.
fenestrata, 316.
glans, 314.
lævigata, 315.
_Montagui_, 308, 492.
purpurata, 318.
_spinulosa_, 321.
spongites, 308.
sporillus, 319.
sulcata, 310.
_tubulosa_, 320.
undulata, 313.
Acorn shell, 33.
Adductor scutorum, 53.
_Adna_, genus, 354.
_Anglica_, 360.
Affinities of the Balanidæ, 152.
Alæ, structure of, 37, 47.
Alcippe, genus, 529.
lampas (female), 530.
lampas (male), 555.
sexes of, 23.
Anelasma, false, resemblance to Xenobalanus, 445.
compared with Alcippe, 528.
Antennæ of larva in first stage, 105.
of pupa, 114.
of Alcippe, 549.
of Cryptophialus, 581.
of Proteolepas, 601.
Anus, 87.
none in Proteolepas, 596.
none in the male Alcippe and Cryptophialus, 546, 562, 585.
Apoda, order of, 22, 587.
Apodemes for the attachment of eyes of pupa, 120, 126.
Apparatus, cementing, 133.
Appendages, caudal, 65, 85, 479, 481, 491.
in Verruca, 510.
in Alcippe, 543.
filamentary, to sack, 64.
to limbs, 64, 83.
_Asemus_, genus, 321.
_porosus_, 329.
_Astrolepas_, genus, 382.
_lævis_, 396.
_rotundarius_, 392.
Attachment of Chelonobia to turtles, 390.
of Coronula to whale's skin, 411.
of Tubicinella, 436.
Baer, Von, on morphological differentiation, 19.
Balanidæ, family of, 33.
highest Cirripedes, 20.
Balaninæ, sub-family of, 175.
characters of, 152.
Balanus, genus, 177.
shell of, immediately after the metamorphosis, 41.
supposed male of, 271.
Ajax, 214.
allium, 281.
amaryllis, 279.
_amphimorphus_, 494.
amphitrite, 240.
_angulosus_, 256.
_arctica patelliformis_, 256.
_arcticus_, 262.
_balænaris_, 415.
balanoides, 267.
pupa of, 130.
monstrous specimens with imperforate penis, 102.
cementing apparatus of, 146.
_balanoides_, 240, 493.
bisulcatus, 293.
_Blainvillii_, 231.
_borealis_, 262.
calceolus, 218.
_candidus_, 277.
Capensis, 209.
_carbonarius_, 492.
cariosus, 273.
_cassis_, 218.
cepa, 283.
_chelytrypetes_, 394.
_clavatus_, 261, 267.
_circinnatus_, 492.
_communis_, 231, 492.
concavus, 235.
_cornubiensis_, 231.
corrugatus, 254.
_Coquimbensis_, 227.
_costatus_, 493.
_Cranchii_, 231.
_crassus_, 195, 493.
crenatus, 261.
_crispus_, 494.
_cylindraceus_, 206, 235, 494.
cymbiformis, 221.
_Cumingii_, 335.
declivis, 275.
decorus, 212.
_delphinus_, 494.
_dentiformis_, 493.
_diadema_, 417.
_discors_, 227.
dolosus, 295.
_d'Orbignii_, 195.
_duploconus_, 365.
eburneus, 248.
_elongatus_, 220, 261, 267.
_erisma_, 296.
_fasciatus_, 201.
_Finchii_, 492.
_fistulosus_, 267.
flosculus, 290.
galeatus, 104, 105, 220.
_geniculatus_, 256.
_gigas_, 210.
_glacialis_, 261.
glandula, 265.
_goissopomo_, 495.
Hameri, 277.
_Holgeri_, 492.
_humilis_, 495.
imperator, 288.
improvisus, 250.
inclusus, 299.
_intertextus_, 518.
lævis, 227.
_lævis_, 495.
_latiradiatus_, 493.
_miser_, 235, 494.
_Montagui_, 308.
navicula, 221.
nigrescens, 210.
nubilus, 253.
_ornatus_, 493.
_ostrearum_, 492.
_ovularis_, 267, 494.
patellaris, 259.
_patellaris_, 494.
_pectinarius_, 493.
_peregrinus_, 492.
perforatus, 231.
upfilled parietal tubes of, 44.
_perforatus_, 494.
_perplexus_, 296.
_picos_, 206.
_pictus_, 493.
_plicarius_, 493.
_plicatus_, 337.
pœcilus, 246.
_polythalamius_, 392.
porcatus, 256.
_porosus_, 493.
_proteus_, 492.
psittacus, 206.
_punctatus_, 267, 493.
_puncturatus_, 337.
_pustula_, 492.
_pustularis_, 235, 493.
_pyramidalis_, 494.
quadrivittatus, 284.
_radiatus_, 242, 494, 495.
_rhomboicus_, 494.
_roseus_, 279.
_rugosus_, 261.
_sagittata_, 493.
_Scoticus_, 256.
_semiplicatus_, 494.
_spongeosus_, 308.
spongicola, 225.
_spongites_, 308.
_squamosus_, 328, 494.
_stalactiferus_, 329.
_stellaris_, 494.
_striatus_, 494.
stultus, 216.
_sublævis_, 493.
_sulcatinus_, 293.
_sulcatus_, 256, 493.
terebratus, 285.
_tertiarius_, 494.
_tesselatus_, 256, 494.
tintinnabulum, 194.
varieties of, 201.
tintinnabulum, eyes of, 93.
nervous system of, 92.
cementing apparatus of, 147.
_tintinnabulum_, 195, 204, 206, 214, 493.
trigonus, 223.
_tulipa_, 195, 205, 277, 494.
tulipiformis, 204.
_Uddevallensis_, 277, 494.
unguiformis, 296.
varians, 298.
_verruca_, 518.
vestitus, 286.
vinaceus, 213.
_virgatus_, 494.
_vulgaris_, 267.
_zonarius_, 235, 494.
Ball, Dr., on the attachment of Chelonobia, 392.
Basis of shell, 49.
cancellated in Bal. lævis, 230.
perforated in Bal. terebratus, 285.
hardly distinguishable in Bal. flosculus, 291.
affected by adhering to corals, 301.
surrounded by a ledge in Chthamalus, 452, 466, 467.
Bate, Mr. C., on the spermatozoa of cirripedes, 99.
on the larva of cirripedes, 103.
on the excavation of Verruca, 514, 518.
Bisexuality of cirripedes, 23.
Boring powers of, in Verruca, 512.
in Alcippe, 549.
_Boscia_, genus, 354.
Bosquet, M., Monographie, &c. 526.
Branchiæ, 63.
in Alcippe, 537.
Brewster, Sir D., on lime and animal matter, 552.
Bristles on membranes of shell and operculum, 59.
Bronn, list of Balanidæ in his Index Palæont., 173.
Burmeister on the rank of cirripedes, 17.
branchiæ of Coronula, 64.
labrum of Coronula, 77.
reproductive system of cirripedes, 97, 100.
larvæ in the second stage, 109.
larvæ in the last stage, 114.
Cæca, ovarian, 100.
to stomach, 85.
Calcar tergi, 52.
Caligus, nerves of, 88.
Canal, alimentary, 85.
Carapace, homologies of, 131.
of pupa, 112.
aborted in Proteolepas, 595, 603.
Carina, posterior compartment of shell, 39.
Catophragmus, genus, 485.
connecting link between the Balanidæ and Lepadidæ, 41.
imbricatus, 490.
polymerus, 487.
Cement, tube of, proceeding from the pupal antennæ, 118.
Cementing apparatus in the pupa, 118, 122.
in the mature animal, 133.
in Platylepas, 426.
in Xenobalanus, 442.
in Proteolepas, 599.
organs of, in crustacea, 151.
_Cetopirus_, genus, 397.
Chamæsipho, genus, 470.
columna, 470.
scutelliformis, 472.
Changes during growth in the Balanidæ, 128, 189.
Characters, variability of, 155, 184, 197.
value of, in the Balanidæ, 154.
Chelonobia, genus, 382.
caretta, 394.
patula, 396.
cementing apparatus of, 145.
_Savignii_, 392.
testudinaria, 392.
_Chirona_, genus, 177.
Chitine, 7, 58.
Chthamalinæ, sub-family of, 446.
characters of, 152.
Chthamalus, genus, 447.
antennatus, 460.
cirratus, 461.
dentatus, 463.
fissus, 462.
_giganteus_, 477, 192.
_glaber_, 455.
Hembeli, 465.
intertextus, 467.
scabrosus, 468.
stellatus, 455.
_stellatus_, 492.
Ciliæ, not present in the articulata, 516.
Circulation, 87.
Cirri, their muscles and movements, 71.
their structure, 81.
their reparation, 158.
in Acasta, with variable hook-like teeth, 306, 311.
antenniformed in Chthamalus, 453, 460.
variability in numbers of segments in Tetraclita porosa, 332.
structure of, variable in Chamæsipho columna, 472.
with one ramus, converted into a triturating button in Alcippe, 542.
abdominal in Cryptophialus, 575.
none in Proteolepas, 587.
Cirripedia, their rank and affinities, 10 to 20.
Cirripedes, how their position is acquired, 127.
young, changes in, 128.
sessile, 33.
_Clisia_, genus, 496.
_striata_, 519.
_Clitia_, genus, 496.
_verruca_, 519.
_Clysia_, genus, 496.
Coldstream, Dr., on the shells of cirripedes, 33.
on a clicking noise, made by cirripedes, 64.
_Columellina_, genus, 424.
_bissexlobata_, 428.
Compartments of shell, number of, 39.
structure of, 43.
Concholepas Peruviana, inhabited by Cryptophialus, 567.
_Conia_, genus, 321.
_depressa_, 337.
_Lyonsii_, 343.
_porosa_, 329.
_radiata_, 343.
_rosea_, 335.
_Conopea_, genus, 177.
remarks on, 190.
_elongata_, 220.
_ovata_, 218.
Corals, effect of attachment to, in Balanus, 301.
Coronula, genus, 397.
balænaris, 415.
cementing apparatus of, 135.
barbara, 421.
_bifida_, 423.
_bissexlobata_, 428.
_Californiensis_, 428.
diadema, 417.
nervous system of, 88.
_patula_, 396.
reginæ, 419.
_sulcata_, 394.
_testudinaria_, 392.
_tubicinella_, 431.
_Coronula_, genus, 382, 424, 430.
_Coronulinæ_, sub-family of, 153.
_Coronulites diadema_, 421.
Creusia, sub-genus, 375.
_Childreni_, 382.
_decorata_, 382.
_grandis_, 376, 381.
_gregaria_, 376, 378.
_madreporarum_, 367, 382.
_multistriata_, 382.
_radiata_, 382.
spinulosa, 376.
_striata_, 382.
_Strömia_, 518.
_verruca_, 518.
_Creusia rayonnante_, 362.
_Creusia_, genus, 354, 497.
Cross-impregnation of Cirripedes, 102, 197.
Crustacea, cementing organs of, 151.
Cryptophialus, sexes of, 23.
minutus (female), 566.
(male), 584.
Dana, Mr., on the classification of Crustacea, 11, 17.
on centralisation in Crustacea, 19.
on nerves of Caligus, 88.
on the antennæ of the larvæ of cirripedes, 106, 114.
on the conversion of mandibles into legs, 107.
on the abortion of the segments of body, 111.
on the distribution of Crustacea, 161, 167.
Daphnia, organs of hearing, 114.
eyes of, 121.
moulting of, 157.
_Daracia_, Genus, 354.
_monticulariæ_, 372.
_Linnæi_, 374.
Denticuli, on the parietal septa, 43.
Deposit, calcareous, inorganic in chamber of Alcippe, 552.
Depths at which cirripedes live, 163.
_Diadema_, genus, 397.
_bifida_, 492.
_vulgare_, 492.
Du Cane, Capt., on the larvæ of Crustacea, 108.
Edwards, Prof. Milne, on the classification of Crustacea, 11, 17.
on the lengthening and shortening of the limbs in Crustacea, 14, 73.
on the 'tige' and 'palpe' of the limbs, 83.
on the sclerodermic plates of the Carapace, 35.
on the Carapace, 131.
on general classification, 529, 565.
Elminius, genus, 345.
Kingii, 348.
cementing apparatus of, 146.
_Leachii_, 348.
modestus, 350.
plicatus, 351.
simplex, 353.
Epidermis, so called, 58.
Epithelium of stomach, 86.
Epizoons (male), 27.
_Euraphia_, genus, 447.
_Hembeli_, 465.
Excavation, powers of, in Verruca, 512.
in Alcippe, 549.
Exuviation of the pupal membranes, 123.
in the mature animal, 157.
in relation to the habits of cirripedes, 56.
anomalous in Isaura, 13.
Eye of larva in first stage, 104.
Eyes of pupa, 119.
of pupa, their exuviation, 126.
in pupa of Alcippe, 549.
the metamorphoses of, in Cirripedes, 120.
of Balanus, 93.
Female organs, 100.
Flabellum of limbs, 83.
Food of Cirripedes, 87.
Fouet of limbs, 83.
Fræna, ovigerous, homologous with branchiæ, 65.
Fræna, ovigerous, modified in Alcippe, 537.
Generation, organs of, 97.
Geography of Cirripedes, 159.
Geological history of Cirripedes, 172.
Glands for cement, 138.
Gnathites, structure and muscles of, 75, 77.
Goodsir, Mr., on the larvæ of Cirripedes, 103, 108.
on his male Balanus, 271.
Gray, Dr., on the homologies of sessile and pedunculated Cirripedes, 34.
on the Coronulinæ, 153.
on marks on the shell of a Balanus, produced by growing on wood, 185.
on the structure of Chthamalus, 448.
Growth of shell, 54.
changes during, 128, 189.
rate of, 156.
of shell in Tubicinella, 436.
Habits of Cirripedes, 159.
Hancock, Mr., on the action of the cirri, 14.
on the excavations of Verruca, 512, 516.
on Alcippe, 530.
on the excavations of Alcippe, 549.
Hanley, Mr. S., on Bal. Hameri, 277.
Hearing, organs of, 95.
Hectocotyle, 23.
Hepatic system, 86.
Hermaphrodite condition of Cirripedes, 23.
Hippolyte varians, larvæ of, 108.
History, geological, of Cirripedes, 172.
Homologies of Cirripedes, 102.
of the carapace and valves, 131.
of body in Proteolepas, 595.
Horner, Mr., on lime and animal matter, 552.
Horsford, Dr., on the setting of lime, 553.
Ibla, sexes of, 23.
Imbedment of Chelonobia in turtles, 390.
of Coronula in whale's skin, 411.
Impregnation, mutual, of Cirripedes, 102, 197, 271.
Injuries, reparation of, 158.
Interbreeding possible in Cirripedes, 102, 197.
Isaura, not moulting its carapace, 13.
Jaws, movements of, 76.
Joly, M., on the Isaura, 13.
on the larvæ of Caradina, 106.
Jones, Dr., on the liver in Daphnia, 86.
Jussieu, Adrien, on classification, 528.
Karsten on the testes and ovaria of Cirripedes, 98, 100.
on supposed hepatic organs, 86.
Kölliker on the spermatozoa of Cirripedes, 99.
Labrum, structure of, 75.
remarkable, in Alcippe, 540.
moveable, in Cryptophialus, 572.
Lamellæ, ovigerous, 101.
in Xenobalanus, 444.
Lamina, internal of wall of shell, cancellated, 213.
Larvæ, first stage, 103.
second stage, 109.
during their early stages, egg-like in Cryptophialus, 579.
last or pupal stage, 110.
Leach, Dr., on the Coronulinæ, 153.
Legs of pupa, 121.
Leidy, Dr., on the eyes of Balanus, 94.
Lepadidæ, family of, 526.
_Lepas angusta_, 231.
_angustata_, 231.
australis, pupa of, 110.
_balanoides_, 231, 240, 267.
_balanus_, 231, 256.
_borealis_, 251.
_caretta_, 394.
_cariosa_, 273, 329.
_cœrulescens_, 342.
_columna_, 470.
_costata_, 256.
_crispata_, 195.
_depressa_, 455.
_diadema_, 417.
fascicularis, cement, float of, 118.
_fistulosus_, 231.
_foliacea_, 261, 277.
_fungites_, 329.
_galeata_, 220.
_Hameri_, 277.
_minor_, 240.
_mitra_, 344.
_ore angustiore_, 231.
_radiata_, 240.
_patellaris_, 259.
_porcata_, 195.
_porosa_, 329.
_psittacus_, 206.
_punctatus_, 455.
_purpurascens_, 337.
_purpurea_, 241.
_Scotica_, 256.
_spinosa_, 195.
_spongiosa_, 308.
_spongites_, 308.
_stellata_, 455.
_striata_, 518.
_Strömia_, 518.
_testudinaria_, 392.
_tintinnabulum_, 194, 235.
_tulipa_, 204, 277.
_verruca_, 518.
Lesson on the Triton, 158.
Limbs of pupa, 121.
Limbus occludens in Pyrgoma, 54.
Lime, tendency to harden with animal matter, 552.
Limulus, nervous system of, 88, 90.
Linnæus on the Triton, 158.
Lithotrya, powers of boring, 535.
Liver, 86.
Lyell, Sir C., on _Bal. Uddevallensis_, 277.
Male organs, 97.
of Alcippe lampas, 555.
of Cryptophialus minutus, 584.
Males of Cirripedes, 23.
of Cirripedes, not of much classificatory importance, 566, 586.
Manatee, with Platylepas attached to the skin, 427.
Mandibles structure of, 76, 79.
reversed in Proteolepas, 589.
Maxillæ, structure of, 76, 80.
reversed in Proteolepas, 590.
_Megatrema_, genus, 354.
_Anglica_, 360.
_semicostata_, 374.
Membrane, covering the shell, 58.
opercular, 58.
_Messula_, genus, 177.
Metagenesis, 124.
Metamorphoses in the Thoracica, 102.
of Alcippe, 548.
remarkable, in Cryptophialus, 579.
Moulting of the pupal membrane, 123.
in the mature animal, 157.
Mouth, structure of, 74, 78.
of pupa, 121.
none in the male Alcippe and Cryptophialus, 562, 585.
suctorial in Proteolepas, 589.
Movements of the cirri, 71.
Muscles of attachment of the body to the operculum, 68.
of the sack, 62.
of the thorax, 69.
of the cirri, 71.
of the jaws, 76.
of sack, weak in Chelonobia, 389.
Nerves splanchnic, 91.
Nervous system, 88.
_Nobia_, genus, 354.
_grandis_, 365.
Nomenclature, rules of, 204.
_Ochthosia_, genus, 497.
_Stroemia_, 519.
Octomeris, genus, 482.
angulosa, 483.
_augubra_, 483.
brunnea, 484.
_Stutchburii_, 483.
Œsophagus, 85.
in the pupa, 121.
with teeth, in Cryptophialus, 577.
Operculum, 50.
remarkable, in Pyrgoma, 356.
remarkable, in Verruca, 497.
Orders of Cirripedes, 20.
Organs, acoustic, 95.
of pupa, 113.
of reproduction, 97.
for cementing, 133.
in the pupa, 122.
Orifice of shell, 38.
Otolithes absent in Crustacea, 97.
Ova, sizes of, 101.
Ovaria, 100.
incipient, in pupa, 122.
occupy a chamber in the shell in Coronula, 411.
in Proteolepas, 597.
Owen, Prof., on Metagenesis, 124.
on vegetative repetition, 20.
Pachylasma, genus, 475.
structure of alæ, 36.
giganteum, 172, 477.
aurantiacum, 480.
Palpi, structure of, 75.
absent in Alcippe, 540.
peculiar in Cryptophialus, 572.
in Proteolepas, 590.
Parasite, allied to Bopyrus, 102.
Parietes of shell, 43.
internal lamina of, cancellated, 213.
perforated in Acasta, 305.
with numerous rows of pores in Tetraclita, 323.
much folded in Coronula, 399.
Penis, probosciformed, 99.
imperforate, 102, 271.
its reparation, 159.
of wonderful length in Cryptophialus, 586.
Perforations in shell of Acasta, 305.
Phyllosoma, affinities to cirripedes, 18.
Platylepas, genus, 424.
bissexlobata, 428.
decorata, 429.
cementing apparatus of, 143.
_pulchra_, 428.
Pœcilasma Kæmpferi, 499.
Pollicipes compared with sessile cirripedes, 34, 41, 52.
_Polylepas_, genus, 397.
_Polytrema_, genus, 321.
Position of cirripedes, how acquired, 127.
Potash, caustic, action on shell, 56.
Primordial valves, 129.
Prosoma, part of thorax, 67.
Proteolepas bivincta, 589.
Pupæ of cirripedes, 110.
Pupa without natatory legs in Cryptophialus, 580.
Pyrgoma, genus, 354.
Anglicum, 360.
cancellatum, 362.
conjugatum, 364.
_corymbosa_, 374.
crenatum, 370.
dentatum, 369.
grande, 365.
_lobata_, 362
milleporæ, 367.
monticulariæ, 372.
_spongiarum_, 374.
_stellata_, 374.
Stokesii, 361.
_sulcatum_, 360, 492.
_undata_, 492
Radii, structure of, 37, 45.
remarkable, with outer lamina of great thickness in Chelonobia, 385.
compound in Coronula, 405.
in Xenobalanus, 441.
Rami of the cirri, their movements and homologies, 71, 83.
modified in Alcippe into triturating organs, 542.
Range, geographical, of cirripedes, 159.
Rathke, on the position of the anus in young Crustacea, 109.
on the cementing organs of Crustacea, 151.
Rectum, 87.
Reparation of injuries, 158.
Reproduction of cirripedes, 97.
Respiration of cirripedes, 64.
Rostrum, anterior compartment of shell, 39.
not symmetrical in Chelonobia caretta, 383.
compounded in Chelonobia, 386.
in Pachylasma, 478, 481.
not quite medial in Platylepas, 424.
Rules of nomenclature, 204.
Sack, 61.
muscles of, 62.
formation of, 124.
Sacks, acoustic, 95.
olfactory, 97
_Savignium_, genus, 354.
Scalpellum, sexes of, 23.
vulgare, larvæ of, 103
Schödler, on organs of hearing in Daphnia, 114.
Schumacher, date of memoir, 321.
Scuta, their structure, 51.
Scutum, with horny articular ridge in Chelonobia, 388.
remarkable from great adductor ridge and occludent ledge in Pyrgoma, 356.
fixed in Verruca, 502.
Sections, transverse, of shells of Balanidæ, 39.
Segments of body in cirripedes, 111, 125.
in Alcippe, 537.
in Cryptophialus, 571.
Senses of cirripedes, 94.
Septa, longitudinal, of the parietes, 43.
Sexes of cirripedes, 23.
Sheath of shell, 38, 48.
peculiar in Tubicinella, 432.
remarkable in Chelonobia, 385.
Shell of a sessile cirripede compared with the shell of the Lepadidæ,
34, 41, 52.
of sessile cirripede compared with the carapace of Crustacea, 35.
elements of, 37.
sheath of, 38, 48.
orifice of, 38.
compartments of, 39, 43.
immediately after the metamorphosis, 41.
walls of, or parietes, 44.
radii of, 37, 45.
alæ of, 37, 47.
basis of, 49,
operculum of, 50.
growth of, 54.
minute structure of, 57.
membranes of, 58.
sack of, 61.
with tubular perforations in Chamæsipho, 473.
almost rudimentary in Xenobalanus, 441.
asymmetrical in Verruca, 499.
minute structure of, in Verruca, 506.
Shells of mollusca excavated by Verruca, 512.
excavated by Alcippe, 549.
Siebold, Von, on Syngamus trachealis, 23.
on the abdomen of cirripedes, 65.
on the cæca of the stomach in Crustacea, 86.
on the vision of cirripedes, 94.
on the acoustic organs of Crustacea, 97.
on the spermatozoa of cirripedes, 99.
on the eyes of cirripedes, 121.
_Siphonicella_, genus, 438.
Size of cirripedes in relation to temperature, 162.
Sowerby, Mr. G. B., on the species of Balani having cup-formed bases, in
the southern hemisphere, 192.
Sowerby, Mr. G. B., jun., on an inorganic deposit in connexion with a
Pholas, 553.
Species, variability of, 155, 184, 197.
Spermatozoa, 98.
Spur of tergum, 52.
St. Ange, M. M., on the mouth of cirripedes, 19, 76.
on the reproductive system of cirripedes, 97, 100.
on the nervous system, 92.
Stomach, 85.
none in Proteolepas, 596.
none in the male Alcippe and Cryptophialus, 562, 585.
Stomopoda, affinities to cirripedes, 19.
Straus, on the affinities of cirripedes, 9.
on the moulting of Daphnia, 157.
Structure, microscopical, of shell, 57.
peculiar in Verruca, 506.
Stutchbury, Mr., on the shell of Chelonobia, 386.
Syngamus trachealis, 23.
System, muscular, of the body and cirri, 62, 68, 71, 76.
alimentary, 85.
circulatory, 87.
nervous, 88.
reproductive, 97.
Temperature, effects of, on cirripedes, 161.
Terga, their structure, 51.
Tergum, occludent ledge and shape of, in Pyrgoma, 356.
fixed in Verruca, 503.
Testæ valvæ, 39, 43.
Testes, 97.
development of, in Proteolepas, 598.
Tetraclita, Genus, 321.
cœrulescens, 342.
costata, 339.
porosa, 328.
upfilled parietal tubes of, 44.
purpurascens, 337.
radiata, 343.
rosea, 335.
serrata, 333.
_squamulosa_, 329.
vitiata, 340.
Thoracica, order of, 21, 30.
Thorax of the Balanidæ, 66.
of pupa, 85.
rudimentary, in the male Alcippe, 562.
Tige of limbs, 83.
Triton of Linnæus, 158.
Tubicinella, genus, 430.
cementing apparatus of, 143.
trachealis, 431.
_balænarum_, 431.
_Lamarckii_, 431.
_major_, 431.
_maxima_, 172, 438, 492.
Turtles, with their carapaces perforated by Chelonobia, 391.
with Platylepas attached, 427.
Unisexuality of cirripedes, 23.
Vagina or sheath, 38, 48.
Valves, opercular, 50.
of shell, 39, 43.
primordial, 129.
homologies of, 131.
Van de Hoeven on Limulus, 88, 90.
Variation of characters, 155, 184, 197.
in Tetraclita, 327.
in Chthamalus, 459.
Verruca, genus, 496.
_cancri Americani_, 396.
lævigata, 520.
nexa, 522.
prisca, 525.
Spengleri, 521.
Strömia, 518.
_Verruca testudinaria_, 392.
Verrucidæ, family of, 495.
Vesicle, acoustic, 95.
Vesiculæ seminales, 98.
Vesicula seminalis, compound in Proteolepas, 598.
Vision of cirripedes, 94.
Wagner, R., on the reproductive system of cirripedes, 97, 99.
Walls of shell, 43.
Water, brackish, effects of, on cirripedes, 163.
Whales' skin, growth of, into the shells of Coronula, 413.
perforated by Tubicinella, 436.
Xenobalanus, genus, 438.
globicipitis, 440.
Zenker on the eyes of Daphnidæ, 121.
Zoosperms, 98.
PLATES.
[Illustration: _Plate I._
_George Sowerby._
BALANUS TINTINNABULUM.]
[Illustration: _Plate II._
_George Sowerby._
BALANUS.]
[Illustration: _Plate III._
_George Sowerby._
BALANUS.]
[Illustration: _Plate IV._
_George Sowerby._
BALANUS.]
[Illustration: _Plate V._
_George Sowerby._
BALANUS.]
[Illustration: _Plate VI._
_George Sowerby._
BALANUS.]
[Illustration: _Plate VII._
_George Sowerby._
BALANUS.]
[Illustration: _Plate VIII._
_George Sowerby._
BALANUS.]
[Illustration: _Plate IX._
_George Sowerby._
ACASTA.]
[Illustration: _Plate X._
_George Sowerby._
TETRACLITA.]
[Illustration: _Plate XI._
_George Sowerby._
TETRACLITA: ELMINIUS.]
[Illustration: _Plate XII._
_George Sowerby._
ELMINIUS: PYRGOMA.]
[Illustration: _Plate XIII._
_George Sowerby._
PYRGOMA: CREUSIA.]
[Illustration: _Plate XIV._
_George Sowerby._
CREUSIA: CHELONOBIA.]
[Illustration: _Plate XV._
_George Sowerby._
CHELONOBIA: CORONULA.]
[Illustration: _Plate XVI._
_George Sowerby._
CORONULA.]
[Illustration: _Plate XVII._
_George Sowerby._
PLATYLEPAS: TUBICINELLA: XENOBALANUS.]
[Illustration: _Plate XVIII._
_George Sowerby._
CHTHAMALUS.]
[Illustration: _Plate XIX._
_George Sowerby._
CHTHAMALUS: CHAMÆSIPHO: PACHYLASMA.]
[Illustration: _Plate XX._
_George Sowerby._
PACHYLASMA: OCTOMERIS: CATOPHRAGMUS.]
[Illustration: _Plate XXI._
_George Sowerby._
VERRUCA.]
[Illustration: _Plate XXII._
_George Sowerby._
ALCIPPE LAMPAS.]
[Illustration: ALCIPPE. _Plate XXIII._
_George Sowerby._
CRYPTOPHIALUS.]
[Illustration: CRYPTOPHIALUS. _Plate XXIV._
_George Sowerby._
PROTEOLEPAS.]
[Illustration: PROTEOLEPAS. _Plate XXV._
_George Sowerby._
BALANUS.]
[Illustration: _Plate XXVI._
_George Sowerby._
MOUTH: THORAX.]
[Illustration: _Plate XXVII._
_George Sowerby._
NERVOUS SYSTEM.]
[Illustration: _Plate XXVIII._
_George Sowerby._
CEMENTING APPARATUS.]
[Illustration: CIRRI. _Plate XXIX._
_George Sowerby._
LARVÆ, FIRST STAGES.]
[Illustration: _Plate XXX._
_George Sowerby._
LARVÆ, LAST STAGES.]
Detailed Transcriber's Notes
Words which were in italics in the original book are surrounded by
underlines (_italic_). Words which were originally printed in small
caps are in ALL CAPS. Obvious misprints have been fixed. Archaic and
unusual words, spellings and styling have been maintained when they
are used consistently. Spelling has been standardized within the book
where one spelling was predominant, even if it is not in common usage.
Inconsistent spelling and hyphenation has been maintained when there
was not a predominant spelling. Details of the changes follow.
No attempt has been made to correct the capitalization or standardize
the italicization of biological names--they are left as in the original.
All of Darwin's errata have been applied to the text, except the one on
page 255. It was not applied because the wording is not obvious.
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.
Details of the Changes
Page vii, footnote 1: du terrain Crétacé du D. de Limbourg
In the original book: du terrain Crétacée du D. de Limbourg
Page 4: the shell, without the operculum being removed,
In the original book: the shell, without the operculum be removed,
Page 5, footnote 3: by some authors as "intersticia."
In the original book: by some authors as "intersticia".
Page 7: carrying two multiarticulated _rami_
In the original book: carrying two multi-articulated _rami_
Page 8: interjection (!)
In the original book: interjection [!]
Page 9: _biramous, multiarticulated limbs_
In the original book: _biramous, multi-articulated limbs_
Page 11: knowledge of the class
In the original book: knowlege of the class
Page 12: first and second pairs of antennæ
In the original book: first and second pairs of attennæ
Page 20: what has happened to one organ
In the original book: what has happenend to one organ
Page 30: ORDER I.--THORACICA.
In the original book: I.--ORDER THORACICA.
Page 33. The table of contents has been completely restructured, to
make the entries match (or compromise with) the titles in the text.
In this ebook: Structure of the alæ
Structure of the sheath
In the original book: Structure of the alæ and sheath
In this ebook:
Structure of the opercular valves (scuta and terga)
In the original book:
Structure of the scuta and terga
In this ebook:
Growth of whole shell and microscopical structure
In the original book:
Growth of whole shell and minute structure of valves
In this ebook: Muscles of sack
In the original book: Sack, muscles of
In this ebook: Movements and muscles of the cirri
In the original book: Cirri, muscles of
In this ebook: Olfactory sacks
In the original book: Olfactory organs
In this ebook: Larva, first stage
In the original book: Larva, first stages
In this ebook: Larva, second stage
Originally this TOC entry was not present in the book.
In this ebook: Affinities, classification, variation
In the original book: Classification and variation
Page 34. Inserted missing title (which appeared in the TOC):
_Structure of Shell._
Page 37: in all cases (as is obvious in Pachylasma)
In the original book: in all cases (as is obvious in _Pachylasma_)
Page 40: the upper lateral valves alternate
In the original book: the upper Lateral valves alternate
Page 45: owing to its midrib, is generally thrust
In the original book: owing to its mid-rib, is generally thrust
Page 45. Inserted missing title (which appeared in the TOC):
_Structure of the Radii._
and left the original paragraph title as it was:
_Radius._
Page 47: and of the recipient furrow in Octomeris (Pl.
In the original book: and of the recipient furrow in _Octomeris_ (Pl.
Page 47: _Structure of the Alæ._
In the original book: _Alæ._
Page 48: _Structure of the Sheath._
In the original book: _Sheath._
Page 49: _Structure of the Basis._
In the original book: _Basis._
Page 51:
_Structure of the Opercular Valves (Scuta and Terga)._
In the original book:
_Opercular Valves._
Page 61: _Muscles of Sack._
In the original book: _Sack; muscles of, &c._
Page 62: can only act as depressores;
In the original book: can only act as depressors;
Page 65:
Balanidæ are the _ovigerous fræna_ of the Lepadidæ
In the original book:
_Balanidæ_ are the _ovigerous fræna_ of the _Lepadidæ_
Page 65. Inserted title from the TOC:
_Thorax and Body._
and demoted the existing title:
_Parts of the body included within the shell or carapace._
to the head of the next paragraph.
Page 67: p. 440, (foot-note), consider the articulated
In the original book: p. 440, foot-note), consider the articulated
Page 68: _Muscular System._
_Attachment of the Body to the Shell._
In the original book:
_Attachment of the Body to the Shell. Muscular System._
Page 76: there is no trace of any labrum
In the original book: there is no trace of any labium
Page 77: (Pl. 26, figs. 3, 4, _c^1_;
In the original book: (Pl. 26, figs. 3, 4, _c_ 1;
Page 78: pairs of gnathites, which latter have only
In the original book: pairs of gnathites; which latter have only
Page 79: thickened membrane (fig. 3, _c^1_)
In the original book: thickened membrane (fig. 3, _c_ 1)
Page 79: thickened membrane (_c^2_), the basal
In the original book: thickened membrane (_c_ 2), the basal
Page 80: piece of membrane (_c^1_) represents
In the original book: piece of membrane (_c_ 1) represents
Page 81: middle segment (fig. 4, _a^1_) of each maxilla
In the original book: middle segment (fig. 4, _a_ 1) of each maxilla
Page 81: this latter (_a^2_) is separated from
In the original book: this latter (_a_ 2) is separated from
Page 81: There are always six pairs; each biramous
In the original book: There are always six pair; each biramous
Page 83: they carry as many as ten pairs in a
In the original book: they carry as many as ten pair in a
Page 84: in the sub-genus Acasta, in which, differently
In the original book: in the sub-genns Acasta, in which, differently
Page 85: developed into strong, downwardly curved teeth
In the original book: developed into strong, downwardly-curved teeth
Page 89: The nerves (Pl. 27, fig. 1, _r^5_, _r^6_)
In the original book: The nerves (Pl. 27, fig. 1, _r_ 5, _r_ 6)
Page 89: outer larger pair (_r^1_) entering
In the original book: outer larger pair (_r_ 1) entering
Page 93. Inserted missing title (which appeared in the TOC):
_Eyes and Vision._
Page 97: _Male Organs of Generation._
In the original book: _Reproductive System._
_Male Organs._
Page 98: _B. Perforatus_ and in the Chthamalus, that
In the original book: _B. Perforatus_ and in the _Chthamalus_, that
Page 99: including the prosoma: in Pachylasma and in
In the original book: including the prosoma: in _Pachylasma_ and in
Page 100: _Female Organs of Generation._
In the original book: _Female Organs._
was a paragraph title.
Page 100:
distinctly in Balanus, Tetraclita, and Coronula, the
In the original book:
distinctly in _Balanus_, _Tetraclita_, and _Coronula_, the
Page 103: _Larva, First Stage._
In the original book this was a paragraph title.
Page 103: the larva (Pl. 29, fig. 9)
In the original book: the larva (Pl. 29, (fig. 9)
Page 104: to the eye, we see, in _Scalpellum vulgare_,
In the original book: to the eye, we see, in _Scalpellam vulgare_,
Page 104. Applied Darwin's errata:
In _Balanus galeatus_, in the immature
In the original book:
In _Balanus elongatus_, in the immature
Page 105. Applied Darwin's errata:
Lepadidæ; but in _Balanus galeatus_
In the original book:
Lepadidæ; but in _Balanus elongatus_
Page 106: the large second pair remains thus
In the original book: the large second pair remain thus
Page 110: _Larva in the Last or Pupal Stage._
In the original book: _Larva in the third or Pupal Stage._
Page 120: though enclosed fairly within the carapace
In the original book: though inclosed fairly within the carapace
Page 121: these latter may possibly be multiplied into
In the original book: these latter may possible be multiplied into
Page 126. Inserted missing title (which appeared in the TOC):
_Act of Metamorphosis._
Page 132: separation of the three cephalic segments.
In the original book: separation of the three caphalic segments.
Page 152: _Affinities, Classification, Variation._
In the original book: _Affinities, Value of Characters, Variation._
Page 163: in Southern Patagonia, and near Guayaquil.
In the original book: in Southern Patagonia, and near Guyaquil.
Page 168. The first two appearances of the text: (1 to 5).
In the original book: (1 to 5.)
Page 169: (3.) SECOND PROVINCE … Balanus … pœcilus*
In the original book: poecilus*
Page 170: (3.) SECOND PROVINCE …
Tetraclita porosa north and south (4 and 5 and W. Indies).
In the original book:
Tetraclita porosa north and south 4 and 5 and W. Indies).
Page 170: (4.) THIRD PROVINCE … Tetraclita … cœrulescens
In the original book: coerulescens
Page 172: species of Balanus abounds in individuals
In the original book: species of Balanns abounds in individuals
Page 172: specimens, said to have come from the eocene
In the original book: specimens, said to have come form the eocene
Page 174. Applied Darwin's errata to the table. Added concavus
and Hammeri to C (Red crag) and Verruca to D (Corraline crag) and
corrected totals.
Page 184: inwardly projecting filamentary appendages.
In the original book: inwardly projecting filimentary appendages.
Page 195: BALANUS TULIPA. _Bruguière._ Encyclop. Meth.,
In the original book: BALANUS TULIPA. _Bruguiere._ Encyclop. Meth.,
Page 196: on ships' bottoms arriving from West Africa
In the original book: on ships'-bottoms arriving from West Africa
Page 198: Archipelago, of a uniform grayish-blue.
In the original book: Archipelago, of a uniform greyish-blue.
Page 200: of America, from Monterey, in lat. 37° N.
In the original book: of America, from Monterez, in lat. 37° N.
Page 206: Zoolog. Journal,
In the original book: Zoolog. Zournal,
Page 211: longitudinally ribbed: colour ashy-gray tinged
In the original book: longitudinally ribbed: colour ashy-grey tinged
Page 211: along a line running from the apex
In the original book: along a line runing from the apex
Page 213: septa are united together, making a network
In the original book: septa are united together, making a net-work
Page 216: Parietes and base porose: shell white, or
In the original book: Parietea and base porose: shell white, or
Page 223: California; Sydney; New Zealand.
In the original book: California; Sidney; New Zealand.
Page 233: _B. nubilus_ and _cariosus_
In the original book: _B. nubibus_ and _cariosus_
Page 234: adhering to the floating _Lepas Hillii_.
In the original book: adhering to the floating _Lepas Hilii_.
Page 235: _Bronn._ Italiens Tertiär-Gebilde (1831)
In the original book: _Bronn._ Italiens Tertiar-Gebilde (1831)
Page 235: _var._ c. _Lamarck._ Animaux sans Vertèbres
In the original book: _var._ c. _Lamarck._ Animaux sans Vertebres
Page 235. Applied Darwin's errata. Added fossil locality:
Red Crag (Sutton) Mus. S. Wood.
Pages 240-241. Italicized the abbreviation "var." in each of the
varieties.
Page 240: lower part uniform blueish-gray,
In the original book: lower part uniform blueish-grey,
Page 241: _Var._ (9) (_an. spec._?) cirratus:
In the original book: Var. (9) (an. spec.?) cirratus:
(See also note above on abbreviation "var.")
Page 244: part of the shell was uniformly blueish-gray.
In the original book: part of the shell was uniformly blueish-grey.
Page 246: 20. BALANUS PŒCILUS.
In the original book: 21. BALANUS PŒCILUS.
Page 248: 21. BALANUS EBURNEUS.
In the original book: 22. BALANUS EBURNEUS.
Page 248: _General Appearance._--Shell conical, or almost
In the original book: _General Appearance_--Shell conical, or almost
Page 250: Southern Patagonia, Guayaquil, West Colombia;
In the original book: Southern Patagonia, Guyaquil, West Colombia;
Page 255. Did not apply Darwin's errata because desired wording
is not obvious. Darwin's errata: six lines from the top, _Bal.
crenatus_, I have now seen a single Red Crag specimen .5 of inch in
basal diameter.
Page 263: their sutural edges are rather thick and
In the original book: their suteral edges are rather thick and
Page 264: vessels are thickly encrusted with this
In the original book: vessels are thickly incrusted with this
Page 264: enormous range and capability of resisting
In the original book: enormous range and cabability of resisting
Page 265: sent me by Krantz from the miocene formation
In the original book: sent me by Krantz from the miocence formation
Page 271: _Range, Habitats, &c._--This species is
In the original book: _Range, Habits, &c._--This species is
Page 278. Applied Darwin's errata. Added fossil locality:
Red Crag (Sutton) Mus. S. Wood.
Page 279: brought from Barbados, in the West Indies,
In the original book: brought from Barbadoes, in the West Indies,
Page 282: coral-inhabiting sub-genus Creusia
In the original book: coral-inhabiting genus Creusia
Page 288: or more strictly "imperial purple;"
In the original book: or more strictly "imperial purple";
Page 292: prominent ridges than even _B. flosculus_
In the original book: prominent ridges than even _var. flosculus_
Page 293. Applied Darwin's errata: Rauville, dans le Cotentin,
In the original book: Rauville, dans le Cotantin,
Page 294: the crests for the tergal depressor muscles
In the original book: the crests for the tergal depressores muscles
Page 297: In both varieties, the parietes
In the original book: In both varietes, the parietes
Page 297: or deeply folded so as to be strongly ribbed
In the original book: or deeply folded so as be strongly ribbed
Page 300. Applied Darwin's errata:
internally, without parietal pores.
In the original book:
internally, without parieted pores.
Page 309: for the depressor muscles are feebly
In the original book: for the depressores muscles are feebly
Page 310: Sydney, Port Fairy,
In the original book: Sidney, Port Fairy,
Page 311, footnote 104: Sydney, which I fully
In the original book: Sidney, which I fully
Page 314: for the depressor muscles are very feebly
In the original book: for the depressores muscles are very feebly
Page 331: The width of the valve and of the spur,
In the original book: Te width of the valve and of the spur,
Page 335: Die Südafrikanischen Mollusken
In the original book: Die Sudafrikanischen Mollusken
Page 337: judging from some dried specimens,
In the original book: judging from some dryed specimens,
Page 338: rostral or lateral depressor muscles
In the original book: rostral or lateral depressores muscles
Page 347: form of a network, or of separate tubes
In the original book: form of a net-work, or of separate tubes
Page 353, footnote 113: I have seen two specimens of
In the original book: I have seen too specimens of
Page 357: forms a mere border to the occludent ledge;
In the original book: forms a mere border to the occludent lodge;
Page 357: between this genus and Creusia, it is
In the original book: between this genus and _Creusia_, it is
Page 369: almost obliterated by the encrusting coral.
In the original book: almost obliterated by the incrusting coral.
Page 376: Under these circumstances I have thought it
In the original book: Under these circustances I have thought it
Page 376: 1. CREUSIA SPINULOSA. Pl. 13,
In the original book: CREUSIA SPINULOSA. Pl. 13,
Page 376:
CREUSIA SPINULEUSE. _De Blainville._ Dict. Sc. Nat.,
In the original book:
CREUSIE SPINULEUSE. _De Blainville._ Dict. Sc. Nat.,
(de Blainville spells it CREUSIA.)
Page 383: In Chelonobia, the parietes are remarkably
In the original book: In Chelonobia, the parieties are remarkably
Page 405: (see a black dot (_d´_) in section, fig. 7)
In the original book: (see a black dot (_d´_) in section 7)
Page 416: alæ are only 1/5th or 1/6th of the thickness
In the original book: alæ are only 1-5th or 1-6th of the thickness
Page 423: Eine tiefe Furche oder Spalte
In the original book: Eine tiefe Furcle oder Spalte
Page 423: von oben herab bis zur Hälfte
In the original book: von oben herab bis zur Halfte
Page 424:
9. _Genus_--PLATYLEPAS. Pl. 17, fig. 1 _a_-2 _b_.
In the original book:
9. _Genus_--PLATYLEPAS. Pl. 17, fig. 1 and 2.
Page 424: whole rostrum is pushed a little on one side
In the original book: whole rostum is pushed a little on one side
Page 431: In numerous other respects Tubicinella
In the original book: In numerous other respects Tubinicella
Page 431: 1. TUBICINELLA TRACHEALIS.
In the original book: 4. TUBICINELLA TRACHEALIS.
Page 431, footnote 127: specific name either of _major_ or _minus_
In the original book: specific name either of _minor_ or _majus_
Page 432: reverse of that usual with sessile cirripedes
In the original book: reverse of that usual with sessile-cirripedes
Page 440: 1. XENOBALANUS GLOBICIPITIS.
In the original book: XENOBALANUS GLOBICIPITIS.
Pages 455-6. Note that under species _Chthamalus stellatus_, both
varieties (_a_) and (_c_) have the name: communis. The original book
reads that way.
Page 465: 6. CHTHAMALUS HEMBELI. Pl. 18,
In the original book: 6. CHTHAMALUS HEMBELI. Tab. 18,
Page 467: causing the shell to be ashy gray.
In the original book: causing the shell to be ashy grey.
Page 471: the projecting, extremely rugged, overlapping,
In the original book: the projecting, extremely rugged, over-lapping,
Page 475: and these occur in Catophragmus
In the original book: and these occur Catophragmus
Page 487: moderately or slightly depressed; colour gray.
In the original book: moderately or slightly depressed; colour grey.
Page 488: as seen, when corroded, in fig. 4 _d_,
In the original book: as seen, when corrroded, in fig. 4 _d_,
Page 492: _Diadema bifidum_ … 'Italiens Tertiär-Gebilde'
In the original book: 'Italiens Tertiar-Gebilde'
Page 492:
_Balanus ostrearum_ … Organic Remains of the Cretaceous Group,'
In the original book:
Organic Remains of the Cretacean Group,'
Page 493: _B. costatus_ … is a synonym of _B. porcatus_
In the original book: is a synonym of _B. porcatns_
Page 493:
_B. ornatus_, Münster, 'Beiträge zur Petrifact.,'
In the original book:
_B. ornatus_, Münster, 'Beitrage zur Petrifact.,'
Page 493:
_B. pectinarius_, Bronn ('Italiens Tertiär-Gebilde,'
In the original book:
_B. pectinarius_, Bronn ('Italiens Tertiar-Gebilde,'
Page 493: _B. porosus_ … this species in Münster's 'Beiträge.'
In the original book: this species in Münster's 'Beitrage.'
Page 494: _B. pustularis_ … in Münster's 'Beiträge,'
In the original book: in Münster's 'Beitrage,'
Page 494: _B. cylindraceus_ … 'Italiens Tertiär-Gebilde,'
In the original book: 'Italiens Tertiar-Gebilde,'
Page 494: _B. Uddevallensis_, Linnæus,
In the original book: _B. Uddewallensis_, Linnæus,
Page 496: members of the Chthamalinæ, though abnormal
In the original book: members of the _Chthamalinæ_, though abnormal
Page 503: under an edge, _z_ (much foreshortened in T′
In the original book: under an edge, _z_ (much fore-shortened in T′
Page 504: umbo of growth matches the umbo of the four
In the original book: umbo of growth matches the umbo: of the four
Page 517: for themselves, could be pried vertically up
In the original book: for themselves, could be prised vertically up
Page 518:
BALANUS VERRUCA. _Bruguière._ Encyclop. Meth., 1789;
In the original book:
BALANUS VERUCA. _Bruguière._ Encyclop. Meth., 1789;
Page 526: The _moveable tergum_ has its upper articular
In the original book: The _moveable_ tergum has its upper articular
Page 529, footnote 141: Crustacés fossiles du terrain Crétacé
In the original book: Crustacés fossiles du terrain Crétacée
Page 529, footnote 142: 'Archives du Muséum'
exemples
comment à des
reconnaître
In the original book: 'Archives du Museum'
examples
comment a des
reconnaitre
Page 538: or the whole anterior part of the animal
In the original book: or the whole anterior part of theanimal
Page 552: although constantly agitated.
In the original book: although constanty agitated.
Page 553: a considerable residuum is left
In the original book: a considerable residium is left
Page 562: down the sides of the sack of the female till
In the original book: down the sides of the sack of female till
Page 562: mere flap, and only two pairs of cirri exist
In the original book: mere flap, and only two pair of cirri exist
Page 562: with four pairs of minute, modified cirri,
In the original book: with four pair of minute, modified cirri,
Page 563: ORDER II.--ABDOMINALIA.
In the original book: ORDER II.--CIRRIPEDIA ABDOMINALIA.
Page 564: close to Alcippe amongst the Lepadidæ
In the original book: close to Alcippe amonst the Lepadidæ
Page 581: dorsal surface, and the overlapping sides
In the original book: dorsal surface, and the over-lapping sides
Page 582, footnote 152: I may here add those of Cryptophialus,
In the original book: I may here add those of Crystophialus,
Page 583: to enclose and protect the antennæ
In the original book: to inclose and protect the antennæ
Page 587: compound structure of the vesicula seminalis
In the original book: compound structure of the visicula seminalis
Page 588: so peculiar, the hermaphrodite condition,
In the original book: so peculiar, the hermaphodite condition,
Page 596: the two threads enclosing the cement-ducts
In the original book: the two threads inclosing the cement-ducts
Page 601: common membrane, lined of course by corium;
In the original book: common membrane, lined of course by coriumi;
Page 604: it does not enclose the thorax or mouth
In the original book: it does not inclose the thorax or mouth
Page 606: Sub-Familia 1. BALANINÆ.
In the original book: Sub-familia 1. BALANINÆ.
Page 607: 2. _Sub-Genus_--ACASTA
In the original book: 2. _Sub-genus_--ACASTA
Page 607: 6. _Sub-Genus_--CREUSIA
In the original book: 6. _Sub-genus_--CREUSIA
Page 607, 9.: quæ basin membranaceam extrorsùs
In the original book: quæ basin membranacean extrorsùs
Page 608: Sub-Familia 2. CHTHAMALINÆ.
In the original book: Sub-familia 2. CHTHAMALINÆ.
Page 611: ORDO I. THORACICA.
In the original book: ORDER THORACICA.
Page 611: Sub-Fam. BALANINÆ.
In the original book: Sub-fam. BALANINÆ.
Page 613, 12.: Tergo lato, pæne æquilaterali.
In the original book: Tergo lato, pœne æquilaterali.
Page 614, 17.: Tab. 4, fig. 3,
In the original book: Tab. 4, fig. 4,
Page 614, 18.: Americâ septent. et meridionali.
In the original book: Americâ septent, et meridionali.
Page 614, 22.: superioribus lævibus, leniter arcuatis,
In the original book: superioribus lævibusleniter arcuatis,
Page 615, 27.: et Indiâ occident. et Africâ meridionali.
In the original book: et Indiâ occident, et Africâ meridionali.
Page 617, 42.: basi poris magnis perforatâ:
In the original book: basi poris maguis perforatâ:
Page 618: _Sub-Genus_--ACASTA
In the original book: _Subgenus_--ACASTA
Page 618, 1.: scuti cristâ articulari
In the original book: scuti cris â articulari
Page 619, 4.: longitudinaliter costatâ, aut corrosâ
In the original book: longitudinaliter costatá, aut corrosâ
Page 622, 2.: et Australiâ septent.; testudinibus affixa.
In the original book: et Australiâ septent; testudinibus affixa.
Page 622: 3. _Chelonobia patula_, Ranzani. (Darwin,
In the original book: 3. _Chelonobia patula_, Ranzani. (Drawin,
Page 624: 1. _Xenobalanus globicipitis_, Steenstrup.
In the original book: 1. _Xenobalanis globicipitis_, Steenstrup.
Page 624: Sub-Fam. CHTHAMALINÆ.
In the original book, this title was missing.
Page 634: 11. _Genus_--POLLICIPES.
In the original book: 11.--_Genus_--POLLICIPES.
Page 637:
18. _Pollicipes validus_ … _Foss._-- … Scaniâ et Maestricht.
In the original book:
Scaniâ et Mäestricht.
Page 638: [_Sectio_ ††.]
In the original book: [†† _Scuta, aut longitudinaliter aut
transversè (i. e. secundum lineas incrementi) costata._]
Page 640: ORDO II. ABDOMINALIA.
In the original book: ORDER II. ABDOMINALIA.
Page 640: ORDO III. APODA.
In the original book: ORDER III. APODA.
Page 641: PLATE 1.
BALANUS TINTINNABULUM.
In the original book: PLATE 1.
_Genus_--BALANUS.
Page 642, Plate 3, 2 _c_ and 2 _d_:
scutum, internal and external views.
In the original book:
scutum and tergum, internal views.
Page 643, Plate 5, 2 _d_: Stutchburii.
In the original book: Stutchburi.
Page 643, Plate 5, 2 _i_: Stutchburii.
In the original book: Stutchburi.
Page 643, Plate 5, 2 _m_, 2 _n_, and 2 _o_: Stutchburii.
In the original book: Stutchburi.
Page 644, Plate 9: _Sub-Genus_--ACASTA.
In the original book: _Sub-genus_--ACASTA.
Page 647, Plate 14: _Genera_--CREUSIA AND CHELONOBIA.
In the original book: _Genera_--CREUSIA, CHELONOBIA.
Page 650, Plate 16: Of the letters in italics, …
In the original book, this sentence was not aligned differently from
the surrounding items.
Page 650, Plate 16: open beneath, and filled up by the epidermis
In the original book: open beneath, and filled up by the epiderdermis
Page 651, Plate 17, 2 _a_: A, rostrum;
In the original book: A, rostum;
Page 653, Plate 19, 4 _b_, 4 _c_, 4 _d_: Chamæsipho scutelliformis,
In the original book: Chamæsipho columna,
Page 657, Plate 22, 8: appear like a bilobed lower lip;
In the original book: appear like a bi-lobed lower lip;
Page 658, Plate 23. Title inserted to match plate:
_Genera_--ALCIPPE AND CRYPTOPHIALUS.
Page 660, Plate 24. Title inserted to match plate:
_Genera_--CRYPTOPHIALUS AND PROTEOLEPAS.
Page 662, Plate 25. Title inserted to match plate:
_Genera_--PROTEOLEPAS AND BALANUS.
Page 664, Plate 26: STRUCTURE OF THE MOUTH AND THORAX.
In the original book: STRUCTURE OF THE MOUTH, &C.
Pages 664-6, Plate 26. The original book enclosed all the letters
of reference in parentheses for this Plate and Plate 29 only. The
parentheses have been removed for consistency with the descriptions
of all the other plates, without noting the individual changes.
Page 669, Plate 28, 4 _a_: _k_, _t_, … the corresponding branch
In the original book: the corrresponding branch
Page 669, Plate 28, 4 _b_: that (_gh_) represented in fig. 4 _a_;
In the original book: that (_g h_) represented in fig. 4 _a_;
Page 669, Plate 29: CIRRI AND LARVÆ, FIRST STAGES.
In the original book: CIRRI: LARVA, FIRST STAGE.
Pages 669-71, Plate 29. The original book enclosed all the letters
of reference in parentheses for this Plate and Plate 26 only. The
parentheses have been removed for consistency with the descriptions
of all the other plates, without noting the individual changes.
Page 670, Plate 29: _letters apply to all the figures, 1 to 6._
In the original book: _letters apply to all the figures, 1 to 5._
Page 674. In Darwin's errata on Page 174:
total in the Coralline Crag ten,
In the original book:
total in the Coralline Crag nine,
Page 675: _Adna_ … _Anglica_, 360.
In the original book: _anglica_, 360.
Page 675:
Baer, Von, on morphological differentiation, 19.
In the original book:
Baer, Von, on morphological differentation, 19.
Page 676: _d'Orbignii_, 195.
In the original book: _D'Orbignii_, 195.
Page 676. The index has been changed due to the errata on pages 104
and 105 changing the references from Balanus elongatus to Balanus
galeatus.
Page 676: (Balanus) _intertextus_, 518.
In the original book: intertextus, 518.
Page 677: (Balanus tintinnabulum) varieties of, 201.
In the original book, the "varieties" listing was at the same level
of indentation as the line before it.
Page 677: (Balanus) tulipiformis, 204.
In the original book: _tulipiformis_, 204.
Page 677: Chamæsipho, genus, 470.
In the original book: Chamœsipho, genus, 470.
Page 678: _Conopea_ … _ovata_, 218.
In the original book: _ovala_, 218.
Page 678: _Creusia rayonnante_, 362.
In the original book: _Creusie rayonnante_, 362.
Page 679: _Daracia_ … _monticulariæ_, 372.
In the original book: _monitculariœ_, 372.
Page 680: _Lepas_ … _fistulosus_, 231.
In the original book: _fistulosa_, 231.
Page 681: Male organs … of Cryptophialus minutus, 584.
In the original book: of Crytophialus minutus, 584.
Page 681. Index entries for _Ochthosia_ and Octomeris were put in
alphabetical order.
Page 681: Otolithes absent in Crustacea, 97.
In the original book: Otolites absent in Crustacea, 97.
Page 681: Penis, probosciformed, of wonderful length
In the original book: Penis, pubosciformed, of wonderful length
Page 683: Stomopoda, affinities to cirripedes, 19.
In the original book: Stomapoda, affinities to cirripedes, 19.
Page 683. Index entry for Tetraclita was put in proper alphabetical
order.
Page 684. Index entry for Verrucidæ was put in proper alphabetical
order.
Pages 685-714. The titles of the plates have been changed. In the
original book, the word plate was abbreviated, _i. e._ "Pl. II."
whereas in this ebook it is "Plate II."
*** END OF THE PROJECT GUTENBERG EBOOK A MONOGRAPH ON CIRRIPEDIA, VOL 2 ***
Updated editions will replace the previous one--the old editions will
be renamed.
Creating the works from print editions not protected by U.S. copyright
law means that no one owns a United States copyright in these works,
so the Foundation (and you!) can copy and distribute it in the
United States without permission and without paying copyright
royalties. Special rules, set forth in the General Terms of Use part
of this license, apply to copying and distributing Project
Gutenberg-tm electronic works to protect the PROJECT GUTENBERG-tm
concept and trademark. Project Gutenberg is a registered trademark,
and may not be used if you charge for an eBook, except by following
the terms of the trademark license, including paying royalties for use
of the Project Gutenberg trademark. If you do not charge anything for
copies of this eBook, complying with the trademark license is very
easy. You may use this eBook for nearly any purpose such as creation
of derivative works, reports, performances and research. Project
Gutenberg eBooks may be modified and printed and given away--you may
do practically ANYTHING in the United States with eBooks not protected
by U.S. copyright law. Redistribution is subject to the trademark
license, especially commercial redistribution.
START: FULL LICENSE
THE FULL PROJECT GUTENBERG LICENSE
PLEASE READ THIS BEFORE YOU DISTRIBUTE OR USE THIS WORK
To protect the Project Gutenberg-tm mission of promoting the free
distribution of electronic works, by using or distributing this work
(or any other work associated in any way with the phrase "Project
Gutenberg"), you agree to comply with all the terms of the Full
Project Gutenberg-tm License available with this file or online at
www.gutenberg.org/license.
Section 1. General Terms of Use and Redistributing Project
Gutenberg-tm electronic works
1.A. By reading or using any part of this Project Gutenberg-tm
electronic work, you indicate that you have read, understand, agree to
and accept all the terms of this license and intellectual property
(trademark/copyright) agreement. If you do not agree to abide by all
the terms of this agreement, you must cease using and return or
destroy all copies of Project Gutenberg-tm electronic works in your
possession. If you paid a fee for obtaining a copy of or access to a
Project Gutenberg-tm electronic work and you do not agree to be bound
by the terms of this agreement, you may obtain a refund from the
person or entity to whom you paid the fee as set forth in paragraph
1.E.8.
1.B. "Project Gutenberg" is a registered trademark. It may only be
used on or associated in any way with an electronic work by people who
agree to be bound by the terms of this agreement. There are a few
things that you can do with most Project Gutenberg-tm electronic works
even without complying with the full terms of this agreement. See
paragraph 1.C below. There are a lot of things you can do with Project
Gutenberg-tm electronic works if you follow the terms of this
agreement and help preserve free future access to Project Gutenberg-tm
electronic works. See paragraph 1.E below.
1.C. The Project Gutenberg Literary Archive Foundation ("the
Foundation" or PGLAF), owns a compilation copyright in the collection
of Project Gutenberg-tm electronic works. Nearly all the individual
works in the collection are in the public domain in the United
States. If an individual work is unprotected by copyright law in the
United States and you are located in the United States, we do not
claim a right to prevent you from copying, distributing, performing,
displaying or creating derivative works based on the work as long as
all references to Project Gutenberg are removed. Of course, we hope
that you will support the Project Gutenberg-tm mission of promoting
free access to electronic works by freely sharing Project Gutenberg-tm
works in compliance with the terms of this agreement for keeping the
Project Gutenberg-tm name associated with the work. You can easily
comply with the terms of this agreement by keeping this work in the
same format with its attached full Project Gutenberg-tm License when
you share it without charge with others.
1.D. The copyright laws of the place where you are located also govern
what you can do with this work. Copyright laws in most countries are
in a constant state of change. If you are outside the United States,
check the laws of your country in addition to the terms of this
agreement before downloading, copying, displaying, performing,
distributing or creating derivative works based on this work or any
other Project Gutenberg-tm work. The Foundation makes no
representations concerning the copyright status of any work in any
country other than the United States.
1.E. Unless you have removed all references to Project Gutenberg:
1.E.1. The following sentence, with active links to, or other
immediate access to, the full Project Gutenberg-tm License must appear
prominently whenever any copy of a Project Gutenberg-tm work (any work
on which the phrase "Project Gutenberg" appears, or with which the
phrase "Project Gutenberg" is associated) is accessed, displayed,
performed, viewed, copied or distributed:
This eBook is for the use of anyone anywhere in the United States and
most other parts of the world at no cost and with almost no
restrictions whatsoever. You may copy it, give it away or re-use it
under the terms of the Project Gutenberg License included with this
eBook or online at www.gutenberg.org. If you are not located in the
United States, you will have to check the laws of the country where
you are located before using this eBook.
1.E.2. If an individual Project Gutenberg-tm electronic work is
derived from texts not protected by U.S. copyright law (does not
contain a notice indicating that it is posted with permission of the
copyright holder), the work can be copied and distributed to anyone in
the United States without paying any fees or charges. If you are
redistributing or providing access to a work with the phrase "Project
Gutenberg" associated with or appearing on the work, you must comply
either with the requirements of paragraphs 1.E.1 through 1.E.7 or
obtain permission for the use of the work and the Project Gutenberg-tm
trademark as set forth in paragraphs 1.E.8 or 1.E.9.
1.E.3. If an individual Project Gutenberg-tm electronic work is posted
with the permission of the copyright holder, your use and distribution
must comply with both paragraphs 1.E.1 through 1.E.7 and any
additional terms imposed by the copyright holder. Additional terms
will be linked to the Project Gutenberg-tm License for all works
posted with the permission of the copyright holder found at the
beginning of this work.
1.E.4. Do not unlink or detach or remove the full Project Gutenberg-tm
License terms from this work, or any files containing a part of this
work or any other work associated with Project Gutenberg-tm.
1.E.5. Do not copy, display, perform, distribute or redistribute this
electronic work, or any part of this electronic work, without
prominently displaying the sentence set forth in paragraph 1.E.1 with
active links or immediate access to the full terms of the Project
Gutenberg-tm License.
1.E.6. You may convert to and distribute this work in any binary,
compressed, marked up, nonproprietary or proprietary form, including
any word processing or hypertext form. However, if you provide access
to or distribute copies of a Project Gutenberg-tm work in a format
other than "Plain Vanilla ASCII" or other format used in the official
version posted on the official Project Gutenberg-tm website
(www.gutenberg.org), you must, at no additional cost, fee or expense
to the user, provide a copy, a means of exporting a copy, or a means
of obtaining a copy upon request, of the work in its original "Plain
Vanilla ASCII" or other form. Any alternate format must include the
full Project Gutenberg-tm License as specified in paragraph 1.E.1.
1.E.7. Do not charge a fee for access to, viewing, displaying,
performing, copying or distributing any Project Gutenberg-tm works
unless you comply with paragraph 1.E.8 or 1.E.9.
1.E.8. You may charge a reasonable fee for copies of or providing
access to or distributing Project Gutenberg-tm electronic works
provided that:
* You pay a royalty fee of 20% of the gross profits you derive from
the use of Project Gutenberg-tm works calculated using the method
you already use to calculate your applicable taxes. The fee is owed
to the owner of the Project Gutenberg-tm trademark, but he has
agreed to donate royalties under this paragraph to the Project
Gutenberg Literary Archive Foundation. Royalty payments must be paid
within 60 days following each date on which you prepare (or are
legally required to prepare) your periodic tax returns. Royalty
payments should be clearly marked as such and sent to the Project
Gutenberg Literary Archive Foundation at the address specified in
Section 4, "Information about donations to the Project Gutenberg
Literary Archive Foundation."
* You provide a full refund of any money paid by a user who notifies
you in writing (or by e-mail) within 30 days of receipt that s/he
does not agree to the terms of the full Project Gutenberg-tm
License. You must require such a user to return or destroy all
copies of the works possessed in a physical medium and discontinue
all use of and all access to other copies of Project Gutenberg-tm
works.
* You provide, in accordance with paragraph 1.F.3, a full refund of
any money paid for a work or a replacement copy, if a defect in the
electronic work is discovered and reported to you within 90 days of
receipt of the work.
* You comply with all other terms of this agreement for free
distribution of Project Gutenberg-tm works.
1.E.9. If you wish to charge a fee or distribute a Project
Gutenberg-tm electronic work or group of works on different terms than
are set forth in this agreement, you must obtain permission in writing
from the Project Gutenberg Literary Archive Foundation, the manager of
the Project Gutenberg-tm trademark. Contact the Foundation as set
forth in Section 3 below.
1.F.
1.F.1. Project Gutenberg volunteers and employees expend considerable
effort to identify, do copyright research on, transcribe and proofread
works not protected by U.S. copyright law in creating the Project
Gutenberg-tm collection. Despite these efforts, Project Gutenberg-tm
electronic works, and the medium on which they may be stored, may
contain "Defects," such as, but not limited to, incomplete, inaccurate
or corrupt data, transcription errors, a copyright or other
intellectual property infringement, a defective or damaged disk or
other medium, a computer virus, or computer codes that damage or
cannot be read by your equipment.
1.F.2. LIMITED WARRANTY, DISCLAIMER OF DAMAGES - Except for the "Right
of Replacement or Refund" described in paragraph 1.F.3, the Project
Gutenberg Literary Archive Foundation, the owner of the Project
Gutenberg-tm trademark, and any other party distributing a Project
Gutenberg-tm electronic work under this agreement, disclaim all
liability to you for damages, costs and expenses, including legal
fees. YOU AGREE THAT YOU HAVE NO REMEDIES FOR NEGLIGENCE, STRICT
LIABILITY, BREACH OF WARRANTY OR BREACH OF CONTRACT EXCEPT THOSE
PROVIDED IN PARAGRAPH 1.F.3. YOU AGREE THAT THE FOUNDATION, THE
TRADEMARK OWNER, AND ANY DISTRIBUTOR UNDER THIS AGREEMENT WILL NOT BE
LIABLE TO YOU FOR ACTUAL, DIRECT, INDIRECT, CONSEQUENTIAL, PUNITIVE OR
INCIDENTAL DAMAGES EVEN IF YOU GIVE NOTICE OF THE POSSIBILITY OF SUCH
DAMAGE.
1.F.3. LIMITED RIGHT OF REPLACEMENT OR REFUND - If you discover a
defect in this electronic work within 90 days of receiving it, you can
receive a refund of the money (if any) you paid for it by sending a
written explanation to the person you received the work from. If you
received the work on a physical medium, you must return the medium
with your written explanation. The person or entity that provided you
with the defective work may elect to provide a replacement copy in
lieu of a refund. If you received the work electronically, the person
or entity providing it to you may choose to give you a second
opportunity to receive the work electronically in lieu of a refund. If
the second copy is also defective, you may demand a refund in writing
without further opportunities to fix the problem.
1.F.4. Except for the limited right of replacement or refund set forth
in paragraph 1.F.3, this work is provided to you 'AS-IS', WITH NO
OTHER WARRANTIES OF ANY KIND, EXPRESS OR IMPLIED, INCLUDING BUT NOT
LIMITED TO WARRANTIES OF MERCHANTABILITY OR FITNESS FOR ANY PURPOSE.
1.F.5. Some states do not allow disclaimers of certain implied
warranties or the exclusion or limitation of certain types of
damages. If any disclaimer or limitation set forth in this agreement
violates the law of the state applicable to this agreement, the
agreement shall be interpreted to make the maximum disclaimer or
limitation permitted by the applicable state law. The invalidity or
unenforceability of any provision of this agreement shall not void the
remaining provisions.
1.F.6. INDEMNITY - You agree to indemnify and hold the Foundation, the
trademark owner, any agent or employee of the Foundation, anyone
providing copies of Project Gutenberg-tm electronic works in
accordance with this agreement, and any volunteers associated with the
production, promotion and distribution of Project Gutenberg-tm
electronic works, harmless from all liability, costs and expenses,
including legal fees, that arise directly or indirectly from any of
the following which you do or cause to occur: (a) distribution of this
or any Project Gutenberg-tm work, (b) alteration, modification, or
additions or deletions to any Project Gutenberg-tm work, and (c) any
Defect you cause.
Section 2. Information about the Mission of Project Gutenberg-tm
Project Gutenberg-tm is synonymous with the free distribution of
electronic works in formats readable by the widest variety of
computers including obsolete, old, middle-aged and new computers. It
exists because of the efforts of hundreds of volunteers and donations
from people in all walks of life.
Volunteers and financial support to provide volunteers with the
assistance they need are critical to reaching Project Gutenberg-tm's
goals and ensuring that the Project Gutenberg-tm collection will
remain freely available for generations to come. In 2001, the Project
Gutenberg Literary Archive Foundation was created to provide a secure
and permanent future for Project Gutenberg-tm and future
generations. To learn more about the Project Gutenberg Literary
Archive Foundation and how your efforts and donations can help, see
Sections 3 and 4 and the Foundation information page at
www.gutenberg.org
Section 3. Information about the Project Gutenberg Literary
Archive Foundation
The Project Gutenberg Literary Archive Foundation is a non-profit
501(c)(3) educational corporation organized under the laws of the
state of Mississippi and granted tax exempt status by the Internal
Revenue Service. The Foundation's EIN or federal tax identification
number is 64-6221541. Contributions to the Project Gutenberg Literary
Archive Foundation are tax deductible to the full extent permitted by
U.S. federal laws and your state's laws.
The Foundation's business office is located at 809 North 1500 West,
Salt Lake City, UT 84116, (801) 596-1887. Email contact links and up
to date contact information can be found at the Foundation's website
and official page at www.gutenberg.org/contact
Section 4. Information about Donations to the Project Gutenberg
Literary Archive Foundation
Project Gutenberg-tm depends upon and cannot survive without
widespread public support and donations to carry out its mission of
increasing the number of public domain and licensed works that can be
freely distributed in machine-readable form accessible by the widest
array of equipment including outdated equipment. Many small donations
($1 to $5,000) are particularly important to maintaining tax exempt
status with the IRS.
The Foundation is committed to complying with the laws regulating
charities and charitable donations in all 50 states of the United
States. Compliance requirements are not uniform and it takes a
considerable effort, much paperwork and many fees to meet and keep up
with these requirements. We do not solicit donations in locations
where we have not received written confirmation of compliance. To SEND
DONATIONS or determine the status of compliance for any particular
state visit www.gutenberg.org/donate
While we cannot and do not solicit contributions from states where we
have not met the solicitation requirements, we know of no prohibition
against accepting unsolicited donations from donors in such states who
approach us with offers to donate.
International donations are gratefully accepted, but we cannot make
any statements concerning tax treatment of donations received from
outside the United States. U.S. laws alone swamp our small staff.
Please check the Project Gutenberg web pages for current donation
methods and addresses. Donations are accepted in a number of other
ways including checks, online payments and credit card donations. To
donate, please visit: www.gutenberg.org/donate
Section 5. General Information About Project Gutenberg-tm electronic works
Professor Michael S. Hart was the originator of the Project
Gutenberg-tm concept of a library of electronic works that could be
freely shared with anyone. For forty years, he produced and
distributed Project Gutenberg-tm eBooks with only a loose network of
volunteer support.
Project Gutenberg-tm eBooks are often created from several printed
editions, all of which are confirmed as not protected by copyright in
the U.S. unless a copyright notice is included. Thus, we do not
necessarily keep eBooks in compliance with any particular paper
edition.
Most people start at our website which has the main PG search
facility: www.gutenberg.org
This website includes information about Project Gutenberg-tm,
including how to make donations to the Project Gutenberg Literary
Archive Foundation, how to help produce our new eBooks, and how to
subscribe to our email newsletter to hear about new eBooks.
