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.

A Monograph on the Sub-class Cirripedia (Volume 2 of 2) The Balanidæ, (or Sessile Cirripedes); the Verrucidæ, etc., etc. · 第 130 / 417 题