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PALHONTOGRAPHICAL SOCIETY.

INSTITUTED MDCCCXLVII.

LONDON:

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MONOGRAPH

ON

ii HOSS iL REPTICTA

1 THI

LONDON CLAY.

PAE tr

SiO Opis. OPHIPDTA.

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t

-ROFESSOR OWEN, F-.R.S., F.L.S., F.G.S.. &c.

SISRARI . LONDON:
PRINTED FOR THE PALSONTOGRAPHICAS, SOCIETY

1S50.

Reprinted with the permission of the Palaeontographical Society, London

JOHNSON REPRINT CORPORATION JOHNSON REPRINT COMPANY LTD.
111 Fifth Avenue, New York, N. Y. 10003 Berkely Square House, London, W. 1

First reprinting, 1966, Johnson Reprint Corporation
Printed in the United States of America

MONOGRAPH

ON THE

FOSSIL REPTILIA OF THE LONDON CLAY.

SUPPLEMENT TO THE ORDER—CHELOMIA.

Familyj—PaLuDINOSAa.

PLATEMYS BOWERBANKII (?). Tab. XXIX, figs. 1, 2.

The evidence of species of Chelonia of the Fresh-water or Marsh-dwelling family,
Paludinosa, has hitherto been derived only from such parts of the skeleton of the
trunk as have been described, figured, and referred to the genera Platemys and mys,
in Part I of the present Monograph, pp. 62-76.

Since those pages were sent to press, Mr. Bowerbank has been so fortunate as to
obtain from the Kocene clay at Sheppy the portion of fossil skull, of which two views
are given of the natural size in T. XXIX, figs. 1, 2. If these figures, and especially
the side view, fig. 1, be compared with the corresponding view of the skulls, T. I,
fig. 1; T. III, fig. 1; T. XV, fig. 1, or T. XI, fig. 2, a marked difference will be dis-
cerned in the form and proportion of the orbit, which is smaller and more nearly
circular in fig. 1, T. XXIX.

But the bony chamber for the eyeball forms one of the characters by which the
skull is distinguished in the marine and fresh-water families of the order Chelonia.
The orbit, for example, is always much larger in proportion to the entire skull in the
marine species, and commonly of the oval form, which is preserved in the beautiful
fossil skull of the Chelone cuneiceps, T. XV, fig. 1; or with the upper and outer part
even more produced and angular than is there represented. In the families Hluviaha
(Trionyx) and Paludinosa (Emydians), the orbit is not merely much smaller in propor-
tion to the skull, it is circular, or nearly so, and not produced at the upper and outer
angle. By this character, we are led to refer the fossil skull under description to
the fresh-water division of the Chelonian order.

Our choice between the Fluviatile or Paludinose families of that division is guided
by the formation of the border of the orbit, and by the proportionate length and the

form of the face or muzzle in advance of it.
1

2 FOSSIL REPTILIA OF THE LONDON CLAY.

In the specics of Lmys (Podocnemys expansa) which I have selected for comparison,
as offering upon the whole the nearest approach, which any Chelonian skull at my
command gives, to the unique fossil in question, the malar bone (7, in Cuvier’s figure
of the skull of Limys expansa, pl. xi, fig. 9, of the ‘Ossemens Fossiles,’ tom. v, pt. ii,
1825; 26 im the figure of the fossil, fig. 1,T. X XIX), becomes much contracted
as it approaches the orbit, to which it contributes a small part of the posterior
border. In the Chelones the malar bone forms a larger proportion of the orbital
rim (see Cuvier, tom. cit., pl. xi, fig. 17), and contributes more to its under
than its back part, which is chiefly formed by the characteristically large postfrontal 12
(g in Cuvier’s figs.); and this character was manifested in the ancient Eocene
turtles as well as in the modern species, as may be seen by reference to the
bones numbered 26 and 12, in T. I, fig. 1; T. XV, fig. 1, of the present work.
The superior maxillary bone 21 (4 in Cuvier’s figs.) is longer in the Hmys, extends
further back in the orbit, and is deeper at its posterior termination, than in the
Chelones. In all these characters, derived from the bones entering into the formation
of the orbit, the fossil under comparison agrees with the Amys, and, indeed, departs
further from the Chelones than the Podocnemys eapansa does, by the much smaller
proportion in which the anteriorly contracted malar bone (26) contributes to the rim
of the orbit. In the Zrvonyces, the malar bone forms a larger proportion of the border
of the orbit than in the Podocnemys expansa, and a fortiori, than in the fossil in question.

The choice between the Fluviatile or Paludinose tribes of the fresh-water Chelonians,
in the determination of this fossil, is better guided by the form and proportions of the
skull anterior to the orbit. In the recent Zrionyces the muzzle is more acute, and in
inost of them more prolonged than in the Emydians, with which the fossil skull
agrees in the shortness of the muzzle; whilst it departs further than most recent
Emydians from the 7ronycide, in the broad truncated character of its anterior termi-
nation. There is also a very well-marked character of affinity to the Podocnemys
expansd, i the smooth and shallow canal which extends from the fore part of the
orbit forwards to the border of the external nostril across the upper part or nasal process
of the superior maxillary bone (21). This groove is very accurately represented in fig. 1,
T. XXIX, in the fossil; itis rather broader in proportion to its length in the Podocnemys
capansa ; Dut so far as it has depended upon the presence and arrangement of the
facial scutes, it is decisive against the fossil having appertained to any species of soft
turtle (Zrionyr), in which such epidermal parts were entirely wanting.

The marks of the supracranial scutes in the fossil are, as in some Emydians, too
feebly and obscurely traceable to permit of a satisfactory comparison of their arrange-
ment. The exterior surface of the prefrontal (16), frontal (11), postfrontal (12), and
parietal (7) bones is subreticulate. The substance of the bones is thick and coarsely
cancellous. The nasal bone is connate with the prefrontal, as in most modern Emydians ;
in the proportion of this compound bone the fossil resembles more the ordinary

CHELONIA. 3

Emydians (inys europea, e. g.) than it does the Podocnemys eapansa. The border of the
prefronto-nasal bone forming the upper part of the nostril is thick and rounded; as is
also the lateral border of the same cavity formed by the maxillary. The lower part
of this border of the maxillary shows the suture for the premaxillary, which must
have presented similar proportions to the premaxillary of the Podocnemys expansa and
other Emydians. The shape of the frontal (11), the proportion of the upper border of
the orbit which it forms, and the course of its sutures with the contiguous bones, are
clearly indicated in fig. 2, T. KXXIX. The straight line formed by the suture between
the frontal (11) and postfrontal (12) resembles that in the Podocnemys expansa ; it is bent
or curved in the Che‘ones. To what extent the postfrontal (12) was continued backwards,
whether so, as with the parietal, to roof over the temporal fossa, as in the Podocnemys
expansa,* or, in a less degree, leaving that fossa open superiorly, as in the Emydians
generally, is a question which will require for its determinauion a more perfect
specimen than the fossil under description. The thickness, however, of the fractured
posterior part of the postfrontal indicates that the bone had been broken not very
close to its natural posterior border, on the supposition that this was free, as in the
Emydians generally ; and the part of the suture of the postfrontal with the parietal
which has been preserved, extends obliquely outwards and backwards, as in Po-
docnemys expansa, not directly backwards, as in most of the Limydes with open temporal
fossee. (Compare Cuvier, loc. cit., fig. 10 with fig. 14, the suture between y and /.)
With respect to the parietal bones (7), these are too much mutilated to show more than
the position and extent of the coronal suture.

A few words may be perhaps expected relative to the difference which the fossil
in question presents to the land-tortoises. In comparison with the skull of a Zestudo
indica of corresponding dimensions with the fossil, the larger proportional size of the
orbits distinguishes the skull of that terrestrial species almost as strongly as the same
character does the skull of the marine turtles. But in addition to this, the malar bone
forms a larger proportion of the back part of the orbit in the TZestudo, and the
prefronto-nasal part of the skull is more bent down ; the suture between the frontals
and prefrontals describes a curve convex forwards in the Testudo, whilst it deviates
very little from a straight line in the fossil, and that little is convex backwards. The
extent also of the upper surface of the postfrontals and parietals, so far as these are
preserved in the fossil, is greater than the whole of those bones in the land-tortoise
compared.

Having been led by the foregoing comparisons to refer the fragment of the fossil
skull (T. XXIX, figs. 1, 2) to the family Paludinosa, it is reasonable to conjecture that
it may have appertained to some one of the large Emydians, which we already know
to have left their carapaces in the Eocene clay of Sheppy. One commonly finds in

* See Cuvier, loc. cit., pl. xi, fig. 1 a.

A FOSSIL REPTILIA OF THE LONDON CLAY.

the recent skeletons of Emydians, that any particular character of the exterior surface
of the bones of the trunk is repeated on the upper surface at least of the bones of the
head. This comparison, in the present instance, indisposes me to regard the fossil in
question as having belonged to the Hmys levis, or to the mys bicarinata, or to the
Platemys Bullockit with the punctate plastron. I should be rather led to select the
Platemys Bowerbanku from the character in question, as exhibited by the carapace and
plastron described at p. 66. But, in provisionally registering the fossil skull in question
under the name of Platemys Bowerbanki, 1 should wish to be understood as by no
means vouching for the accuracy of the reference. The conjecture rests solely on the
character above referred to, which is far from being decisive; and its only value is,
that it happens to be the only one by which we can be guided at present in forming
any opinion at all as to the specific relations of the fossil in question.

PAE UT.

OrnDER—CROCODILIA.

CROCODILES, ALLIGATORS, GAVIALS.

OF the numerous and various kinds of Reptiles, the fossil remains of which have
been discovered in the tertiary and secondary strata of Great Britain, many are
found to have their nearest representatives, amongst the actual members of the
class, in the present order; and here more particularly in the long and narrow-snouted
genus called, through a corrupt latinization of its native name, Gavialis, which is now
represented by the Gavial or, more properly, Garrhial, of the river Ganges.

In the interpretation of the fossil remains of Reptiles, no skeleton has more
frequently to be referred to than that of the Gavial or Crocodile, or has thrown more
light on the nature of those singularly modified forms of the class which have long
since passed away.

It is accordingly requisite for the paleontologist who would describe the fossil
remains of reptiles, to make himself, in the first place, thoroughly conversant with the
osteology of the recent Crocodila. This knowledge can be gained only by assiduous
study of the skeletons themselves, with the aid of the best descriptions, or the guide
of a competent teacher. But to enable the reader to follow or comprehend the
description of the fossil Saurians, some elementary account of the Crocodilian skeleton
is at least necessary, accompanied with illustrations of the parts which, in the sequel,
will have to be frequently referred to under special or technical names.

In Tab. XI of the present part of this Monogragh is given a reduced or miniature
side view of the skeleton of a Gavial which was twenty-five feet in length—dimersions
which are rarely found to-be surpassed in the present day. Beneath it is a restoration
of the skeleton of the Teleosaur, or extinct Gavial of the Triassic or Oolitic period,
showing how closely the general type of conformation has been adhered to, the modi-
fications of the more ancient form of Crocodile evidently adapting it for moving with
greater speed and facility through the water, and indicating it to have been more
strictly aquatic, and probably marine.

The particular nature of these modifications will be explained when I come to
describe the Crocodiles of the secondary strata. I propose at present to give a pre-
liminary sketch of the osteology of the recent Crocodilia.

A glance at a natural or well-articulated skeleton of one of these reptiles, such as

6 FOSSIL REPTILIaA OF THE LONDON CLAY.

is figured in T. XI, will show that it consists mainly of a series of segments, more or
less alike. From the back of the head to the end of the tail, the chief part of each
segment consists of a cylindrical portion or ‘body,’ differing only in its proportions,
and diminishing as it recedes from the trunk. Every segment sends a plate of bone
upwards from its upper or dorsal surface, which plate or ‘ spine’ is supported by an
arch of bone, except in the diminishing segments at the end of the tail.

Other plates of bone, of more variable forms and dimensions, project from each side
of the segments of the trunk and basal part of the tail. In a less proportion, but still
in a great number of the segments, an arch of bone is formed below, or on the ventral
side of the cylindrical body ; but this lower arch is more variable in its proportions
and mode of composition than the upper arch: it is open or incomplete in the neck.
Under all these variations, however, there is plainly manifested a fundamental unity
of plan in the composition of the different segments, which have accordingly received
the common appellation of ‘ vertebra.’

For the convenience of description, the vertebree are divided, though somewhat
arbitrarily, mto groups bearing special or specific names. Those next the head, with
the inferior arch incomplete below, are called ‘ cervical vertebra;’ they are usually nine
in. number: those that follow with the inferior arch closed below, or which have the
laterally projecting parts slender and freely moveable, are called ‘dorsal vertebre ;’
the other vertebra of the trunk that have no lateral moveable appendages, are called
‘lumbar vertebra ;’ the last vertebrae of the trunk, always two in number in the
Crocodilia, the inferior arches of which coalesce to support and be supported by the
hind limbs, are the ‘ sacral vertebra ;’ the segments of the tail are the ‘ coccygeal,’
or ‘caudal vertebree, whether they possess or not an inferior arch, or whatever other
modifications they may offer.

These names, ‘cervical,’ ‘dorsal,’ ‘lumbar,’ ‘sacral, ‘coccygeal, were ori-
ginally applied to corresponding segments or vertebre in the human skeleton, from the
study of which the nomenclature of osteology takes its date: 1t may well be supposed,
therefore, that a classification and designation of vertebrse based upon knowledge
limited to their characters in a single example of the vertebrated series, and that
example one in which the common type has been most departed from, to adapt it to
the peculiar attitude and powers of the human species, would fall far short of what
is required to express the general ideas derived from a comparison of all the leading
modifications of the vertebrate skeleton ; and accordingly the anatomist who passes
from a previous acquaintance with human osteology only, to the study of those of the
lower Vertebrata, finds that he has to rectify, in the first place, the erroneous notions
which anthropotomy has taught him of the nature of the primary segment of his own
and other vertebrated skeletons, and to acquire true ideas, with the concomitant
nomenclature, of the essential constituents or anatomical elements of such segment.

In human anatomy, for example, the costal elements are only recognised when they

CROCODILIA. fl

retain throughout life that distinctness, or moveable union with the rest of their
segment, which they manifest at their first appearance; and they are then classified as
distinct bones from the rest of their segment, to which the term ‘vertebra’ is
restricted, and which is equally regarded as a single bone; as, e. g., in the dorsal
region of the skeleton. In the cervical region the whole segment is called ‘ vertebra,’
and is recognised as the equivalent bone to a dorsal vertebra, although it includes the
costal elements, because these have coalesced with the rest of their segment, which
anchylosis is misinterpreted as a mere modification of a transverse process; and the
‘cervical vertebra’ is distinguished by having that process ‘perforated,’ and not
entire as in the other vertebre.

But, in the Crocodile, the embryonic condition of the cervical ribs in Man is
retained throughout life; and, therefore, if we were to be guided by the characters laid
down by the recognised authorities in anthropotomy for the classification of its vertebre,
we should seek in vain for any vertebrze with “ transverse processes perforated for the
transmission of vertebral arteries,” whilst we should find all the vertebrze from the
head to the loins, ‘‘ with articular surfaces, either on their sides or their transverse
processes, where they join with ribs,” and should accordingly have to reckon these as
“ dorsal vertebre.”’

These and many similar instances which might be adduced, have compelled me to
premise a few brief explanations of the principles and nomenclature by which I shall
describe the fossil remains of the Hepéilia, and illustrate their nature by reference to the
skeletons of their existing representatives, in the present and succeeding Monographs.

The primary segment of the skeleton of all Vertebrata isa natural group of bones,
which may be severally recognised and defined under all the modifications to which
such segment may have been subjected in subservient adaptation to the habits and
exigencies of a particular species.

A view of such a segment, as it exists in the thorax of the crocodile, the tortoise,
and the bird, is given at p. 5, Part I, of the present Monograph.

The part marked c is the ‘centrum,’ or body of the vertebral segment; it is
always developed originally as a separate element, and retains its character of
individuality in the tortoise and crocodile. The bony arch above the centrum was
formed originally by two distinct side-plates,—the ‘neurapophyses,’ x, which coalesce
with one another at their summits and thence develope a median plate or process of
bone called the ‘neural spine’ zs. Other bony processes which shoot out from the
neurapophyses are more variable, and will be afterwards noticed. The arch so formed
coalesces with the centrum in the bird, and constitutes an apparently single bone, to
which, in anthropotomy, the name ‘ vertebra’ would be restricted. But it would be
as reasonable to confine it to the central element (c) in the tortoise and crocodile ; for
the parts of the inferior arch are not less essentially parts of the same natural segment,
than the neurapophyses which have formed the upper arch. The next pair of elements,

8 FOSSIL REPTILIA OF THE LONDON CLAY.

then, which we have to notice, is marked in figs. 4, 5, and 6, p/, signifying ‘ pleura-
pophysis,’ the name of these elements. In the segments figured they retain their
primitive distinctness, and acquire unusual length, in order to aid in encompassing the
dilated canal or cavity for the heart and lungs: so modified, these elements are
commonly called ‘ribs,’ or ‘ vertebral ribs.’

The elements more constantly employed to protect the vascular or ‘ heemal’ axis, in
other words, to form the inferior or hzmal canal, are those marked / in figs. 4 and 6:
they are the ‘ heamapophyses,’ which are usually articulated, like the neurapophyses,
with the centrum, but are displaced by the great centres of the vascular system in the
thorax, where they have got the special name of ‘ sternal ribs,’ and also that of ‘ costal
cartilages,’ or ‘cartilages of the ribs,’ when they do not become ossified. The hzmal
arch in the thorax is usually completed by a median element (As), called a ‘ heemal
spine, but which itself becomes vastly expanded in the bird (fig. 4) ; it 1s, nevertheless,
the part in the hemal arch which repeats below, or answers to the part (ws) in the
upper arch. In the segments of the trunk and tail, the element (zs) retains its normal
size and form as a ‘neural spine; but where the central axis of the nervous system
becomes unusually developed, as in the head, e. g., analogously to the development of
the vascular centres in the chest, the neural canal is correspondingly expanded, and the
cavity acquires a special name, and is called ‘ cranium,’ just as the analogously expanded
heemal canal is called ‘ thorax.’ Into the formation of the wall of the cranium other
vertebral elements enter besides the neurapophyses, those e. g. which are numbered 8
and 12 in fig. 9, p. 17, of the present Part; the neural spine (7 and 11 in the same figure)
retains its primitive distinctness, is expanded horizontally, and, like the ‘sternum’ in the
thorax of the bird (As, fig. 4, p. 5, Part I), it receives a special name (7), e. g. of ‘ parietal’,
and (9) of ‘ frontal’ in fig. 9. The elements a a (figs. 4 and 6, Part I) form a symmetrical
pair of bones or cartilages, attached at one end to the hemal arch, and projecting out-
wards and backwards. These are the ‘ prosartemata,’ or appendages ; they are, of all the
elements of the vertebral segment, those that are least constant in regard to their
presence, and, when present, are subject to the greatest amount of development and
metamorphosis: they become, e. g., the pterygoid appendages in the nasal segment of the
fish’s skull, the opercular bones in the frontal segment of the fish, the branchiostegal
rays in the parietal segment, the pectoral fins in the occipital segment, and they are deve-
loped into the fore limbs and hind limbs, the arms, wings, and legs of other Vertebrata.*

As the nervous and vascular centres become reduced in size, the bony canals or
arches protecting them are simplified and contracted, and the vertebra assumes a
symmetrical character. In the Crocodile, the hemal arch, in the tail,e. g., is formed
by the hzemapophyses, which ascend and articulate directly with the centrum ;
the pleurapophyses are shortened, directed outwards, and become anchylosed to

* The facts and arguments in support of this conclusion, are detailed in my works ‘On the Nature of
Limbs,’ and ‘On the Archetype of the Vertebrate Skeleton,’ 8vo (Van Voorst).

CROCODILIA. 9

form ‘transverse processes ;) but such a vertebra, when analysed as it is developed,

resolves itself very nearly into the ideal type Fig. 7.

given in the subjomed diagrammatic cut feeoint spine:
: 3 5 cygapophysis.~., Hh

(fig. 7); wis the neural axis, called ‘myelon, or ar Wir aes

\ .
‘ \ __-ereoneurapophysis.

‘spinal marrow ;’ /is the hema! axis, the chief
trunk of which is called ‘aorta,’ and ‘ caudal ston
artery. The names of the vertebral elements SO a @ ESS pleurapophysis.

. 5 e ° ca ZED
which, being usually developed from distinct _ parapophysis.““ ©
h Pe,
Bruins hamapophysis.

centres, are called ‘ autogenous,’ are printed in

Roman type ; the Jfalics denote the ‘ exogenous’ sygapophysis.~
parts, more properly called ‘ processes,’ which ~~ hvemal spine.
shoot out from the preceding elements. Ideal typical vertebra.

On comparing this form of the primary segment with that figured in Cut 4,
p. 5, Part I, it will be seen that they differ by altered proportions with some change of
position of certain elements ; but every mcdification resulting in the various forms of the
parts of the skeleton figured in T. XI, has its seat in one or other of the segmental
or ‘ vertebral’ elements above defined; and the same principle I believe that I have
established with regard to the internal skeleton in all vertebrate animals.

With this preliminary explanation, the nature and relations to the typical vertebra of
the parts of the Crocodilian vertebre, figured in T. IV, V, IX, will be, itis hoped, readily
appreciated. In T. IX, in which are figured, through opportunities kindly afforded
by the Marchioness of Hastings, and Searles Wood, Hsq., F.G.S., some of the most
perfectly-preserved fossil reptilian vertebree which have hitherto been discovered, the
elements and processes are indicated by the initial letter of their names. Figs. 1 and 2
give a side view and a back view of a cervical vertebra, apparently the fourth, of the
Crocodilus Hastingsie, from the Eocene deposits at Hordwell; cis the centrum, x the
neural canal formed by the neurapophyses, which have coalesced superiorly with each
other, and with the neural spine (vs). Inferiorly they articulate by a suture (which is
shown by the wavy line on each side of the process d in fig. 1) with the centrum; p/ is
the pleurapophysis, which articulates by two parts, the lower one called the ‘ head’
to the process from the centrum, the upper one called the ‘tubercle’ to the process
from the neurapophysis ; beyond the union of the head and tubercle, the pleurapophysis
projects freely outwards and downwards, but instead of being elongated in that
direction, it becomes expanded in the direction of the axis of the body, i. e. forwards
and backwards, and so acquires a shape which has given rise to the name ‘hatchet
bone’ or ‘ hatchet-shaped process,’ * applied to this element in the Pleszosaurus.

* «To compensate for the weakness that would have attended this great elongation of the neck, the
Plesiosaurus had an addition of a series of hatchet-shaped processes on each side of the lower part of the
cervical vertebra.” (Buckland, Bridgewater Treatise, vol. i, p. 206, and vol. ii, p. 30, 1836.)

Cuvier recognised in these lateral bones, ‘‘en forme de hache,”’ the homologues of the “ petites cétes
cervicales” of the Crocodile. (Ossemens Fossiles, 4to, tom. v, pt. ii, p. 479, 1824.) And Conybeare had

2

10 FOSSIL REPTILIA OF THE LONDON CLAY.

The purport of this modification is the same in the Crocodilia as that which seems
to be more called for in the Plesiosaurus, viz. to augment the strength of the cervical
region of the skeleton ; and this is so effectually done by the overlapping of the hatchet-
shaped ribs of this region in the Crocodilia, as shown in T. XI, that the flexibility of
the neck is much restricted, although the joint of the head allows that part to be
bent from side to side at nearly right angles with the neck. When, however, the head
is held firmly forwards by its powerful muscles, the imbricated vertebree of the neck
transmit with great effect the impulse which the strong and long tail gives to the rest
of the body in the act of swimming.

In T. IX, fig. 3 the cervical vertebra is represented minus its pleurapophyses, and it
answers accordingly to that portion of the natural segment to which the term ‘ vertebra’
is usually restricted in the dorsal region of the trunk. The exogenous processes shown
in this view of the vertebra are, p, the ‘parapophysis’ or inferior transverse process,
developed from the centrum; d, the ‘diapophysis’ or upper transverse process de-
veloped, as in most cases it is, from the neurapophysis; 2, 2, are the ‘ zygapophyses’
or ‘ oblique processes,’ which, from their function im articulating together contiguous
vertebrae, are also called ‘articular processes.’ In most of the cervical, and in some
of the dorsal, vertebree of the Crocodile, an exogenous process is developed from the
under surface of the centrum, called ‘hypapophysis ;’ it is indicated by the letters /y in
fig. 2, T. LX. In some species it is double,* and beneath the atlas it becomes ‘ auto-
genous or is developed as a separate element, ca, ev, of the subjoined Cut, fig. 8, in which
condition the part is found beneath the centrums of two or three of the anterior cervical
vertebree in the Ichthyosaurus.+

The first and second vertebre of the neck are peculiarly modified in most air-breath-

Fie. 8. ing Vertebrata, and have accordingly received the special names, the
one of ‘ atlas,’ the other of ‘ epistropheus’ or ‘ axis.’. In Comparative
Anatomy these become arbitrary terms, the properties being soon
lost which suggested those names to the human anatomist; the
‘atlas’ e. g. has no power of rotation upon the ‘axis’ in the
Crocodile, and it is only in the upright skeleton of man that the

Atlas and Axis vertebre 2° globular head is sustained upon the shoulder-like processes

of the Crocodile. of the ‘atlas.’ In the Crocodile, these vertekize are concealed by
the peculiarly prolonged angle of the lower jaw in the side view of the skeleton in T. XI,

previously extended the same homology to the “particularly prominent wing-lke appendages to the
transverse processes in many of the long-necked quadrupeds, and the long styloid processes of the cervical
vertebra of birds.” (See his admirable Memoir of June 14th, 1822, in the Geol. Trans., 2d series, vol. i, p. 384.)

* In Orocodilus basifissus, e. g., see the Quarterly Journal of the Geological Society, November 1849,
p. 381, pl. x, fig. 2.

+ This interesting discovery was communicated by its author, Sir Philip de M. Grey Egerton, Bart., F.G.S.,
to the Geological Society of London, in 1836, and is published in the fifth volume of the second series of
their Transactions, p. 187, pl. 14.

CROCODILIA. et

and a separate view of them is, therefore, given in figure 8. The pleurapophyses are
retained in both segments, as in all the other vertebre of the trunk. That of the
atlas, fig. 8, pla, is a simple slender style, articulated by the head only, to the
independently developed inferior part of the centrum, or ‘hypapophysis’ (ca, ez). The
neurapophyses (za) of the atlas retain their primitive distinctness; each rests in part
upon the proper body of the atlas (ca), in part upon the hypapophysis. The neural
spine (zs, a) is also here an independent part, and rests upon the upper extremities
of the neurapophyses. It is broad and flat, and prepares us for the further
metamorphosis of the corresponding element in the cranial vertebree.

The centrum of the atlas (ca), called the odontoid process of the epistropheus in
Human Anatomy, here supports the abnormally-advanced rib of that vertebra, which in
some Crocodilia is articulated by a bifurcate extremity, like the ribs of the succeeding
cervical vertebre ; but it is not expanded or hatchet-shaped at the free extremity.
The proper centrum of the axis vertebra (cz) is the only one in the cervical series
which does not support a rib ; it articulates by suture with its neurapophyses (zz), and
is characterised by having its anterior surface flat, and its posterior one convex.

With the exception of the two sacral vertebre, the bodies of which have one articular
surface flat and the other concave, and of the first caudal vertebra, the body of which has
both articular surfaces convex, the bodies of all the vertebree beyond the axis have the
anterior articular surface concave, and the posterior one convex, and articulate with one
another by ball-and-socket joints. This type of vertebra, which I have termed ‘ procee-
lian, * characterises all the existing genera and species of the family Crocodilia, with all
the extinct species of the tertiary periods, and also two extinct species of the Greensand
formation in New Jersey.t Here, so far as our present knowledge extends, the type
was lost, and other dispositions ‘of the articular surfaces of the centrum occur in the
vertebre of the Crocodilia of the older secondary formations. The only known
Crocodilian genus of the periods antecedent to the Chalk and Greensand deposits with
vertebre articulated together by ball-and-socket joints, have the position of the cup
and the ball the reverse of that in the modern Crocodiles, and the genus, thus cha-
racterised by vertebrz of the ‘opisthoccelian’ type, has accordingly been termed
Streptospondylus, signifying ‘vertebre reversed.’ The aspects of the zygapophyses
are, however, more constant ; the anterior ones, T. IX, fig. 3 2, look obliquely inwards ;
the posterior ones, ib. 2, obliquely outwards. In looking, therefore, upon the cut
surface of a vertical longitudinal section of a Crocodilian vertebra, the smooth,
flattened inner surface of the anterior zygapophysis is turned towards the observer,
and the convex outer surface of the posterior zygapophysis. Thus the anterior and
posterior extremity of the vertebra being determined by observation of the aspect
and direction of the zygapophyses, it is at once seen whether the body has the

* TIpos, before ; xovhos, concave.
+ Quarterly Journal of the Geological Society, November 1849.

12 FOSSIL REPTILIA OF THE LONDON CLAY.

proceelian structure, as in the true Crocodiles, T. IV, T. IX, or the opisthoccelian
structure, as in the Streptospondylus. But the most prevalent type of vertebra amongst
the Crocodilia of the secondary periods was that in which both articular surfaces of
the centrum were concave, but in a less degree than in the single concave surface of
the vertebree united by ball and socket. A section of a vertebra of this ‘amphiccelian’
type, such as existed in the Te/eosaurus and Stencosaurus, will be figured in a subsequent
Monograph. In the Jchthyosaurus, the concave surfaces are usually remarkable for
their depth, the vertebree resembling in this respect those of fishes. Some of the most
gigantic of the Crocodila of the secondary strata had one end of the vertebral centrum
flattened, and the other (hinder) end concave; this ‘ platyccelian’ type we find in the
dorsal and caudal vertebree of the gigantic Cetiosaurus.

With a few exceptions, all the modern Reptiles of the order Lacertilia have the same
proceelian type of vertebrae as the modern Crocodilia, and the same structure prevailed
as far back as the period of the Mosasauwrus, and in some smaller members of the
Lacertilian order in the Cretaceous and Wealden epochs.

Resuming the special description of the osteology of the modern 2rocodilia, we find
the proccelian type of centrum established in the third cervical, which is shorter but
broader than the second; a parapophysis is developed from the side of the centrum,
and a diapophysis from the base of the neural arch; the pleurapophysis is shorter, its
fixed extremity is bifid, articulating to the two above-named processes; its free ex-
tremity expands, and its anterior angle is directed forwards to abut against the inner
surface of the extremity of the rib of both the axis and atlas, whilst its posterior pro-
longation overlaps the rib of the fourth vertebra.

The same general characters and imbricated coadaptation of the ribs characterise
the succeeding cervical vertebrze to the seventh inclusive, the hypapophysis (Ay, fig. 2,
T. IX) progressively though slightly increasing in size. In the eighth cervical the
rib becomes elongated and slender; the anterior angle is almost or quite suppressed,
and the posterior one more developed and produced more downwards, so as to form
the body of the rib, which terminates, however, in a free point. In the ninth
cervical the rib is increased in length, but is still what would be termed a ‘ false’
or ‘ floating rib’ in anthropotomy.

In the succeeding vertebra the pleurapophysis articulates with a heemapophysis, and
the hzemal arch is completed by a hzemal spine; and by this completion of the typical
segment we distinguish the commencement of the series of dorsal vertebree. With regard
to the so-called ‘ perforation of’ the transverse process,’ this equally exists in the present
vertebra, as in the cervicals, as may be seen by comparing fig. 6, p. 5, Part I of this
Monograph, with fig. 2,T. IX, Part II; in both, the foramen is the vacuity intercepted
between the bifurcate extremity of the rib and the rest of the vertebra with which
that rib articulates; and, on the other hand, the cervical vertebrz equally show
surfaces for the articulation of ribs. Cuvier, in including the proximal portions
of the ribs with the rest of the vertebra, in his figure of a dorsal vertebra of a

CROCODILIA. i3

Crocodile,* so far follows nature, and produces a parallel to his figure of a cervical
vertebra; but the entire natural vertebra or segment includes the parts delineated
in outline in Cut 6, p. 5, Part I. In that figure is shown the semiossified bar /’
which is interposed between the pleurapophysis p/ and hemapophysis / in the
Crocodilia and some existing Lizards. The typical characters of the segment due to
the completion of both neural and hemal arches, is continued in some species of
Crocodilia to the sixteenth, in some (Crocodilus acutus) to the eighteenth vertebra. In
the Crocodilus acutus and the Alligutor lucius, the hemapophysis of the eighth dorsal
rib (seventeenth segment from the head) joins that of the antecedent vertebra. The
pleurapophyses project freely outwards, and become ‘ floating ribs’ in the eighteenth,
nineteenth, and twentieth vertebrze, in which they become rapidly shorter, and in the
last appear as mere appendages to the end of the long and broad diapophyses:
but the hemapophyses by no means disappear after the solution of their union
with their pleurapophyses ; they are essentially independent elements of the segment,
and they are continued, therefore, in pairs along the ventral surface of the abdomen
of the Crocodilia, as far as their modified homotypes the pubic bones. They are
more or less ossified, and are generally divided into two or three pieces.

Another character afforded by the hzmal arch is the more important in reference
to palzontology, as it affects the centrum and neural arch of the vertebra as well as
the pleurapophysis; and thus aids in the determination of the vertebra. The
parapophysis progressively ascends upon the side of the centrum in the two anterior
dorsal vertebrze, and disappears in the third, or, passing upon its neurapophysis, blends
with the base of the diapophysis. In this segment, therefore, the proximal end of the
rib ceases to be bifurcate, but is simply notched, the curtailed head being applied to
the end of the thickened anterior part of the transverse process, and the tubercle
abutting against its extremity ; in the five following dorsals the head and tubercle of
the rib progressively approximate and blend together, or the head disappears in the
tenth dorsal, in which the rib is simply attached to the end of the diapophysis. The
hypapophysis ceases to be developed after the third or fourth dorsal vertebrae. The
zygapophyses become gradually more horizontal, the anterior ones looking more
- directly upwards, the posterior ones downwards.

The ‘lumbar vertebre’ are those in which the diapophyses cease to support
moveable pleurapophyses, although they are elongated by the coalesced rudiments of
such which are distinct in the young Crocodilia. The development and persistent
individuality of more or fewer of these rudimental ribs determines the number of the
dorsal and lumbar vertebre respectively, and exemplifies the purely artificial character
of the distinction. The number of vertebre or segments between the skull and the
sacrum, in all the Crocodilia I have yet examined, is twenty-four. In the skeleton of

* Ossemens Fossiles, 4to, tom. v, pt. i). pl. iv, fig. 4.

14 FOSSIL REPTILIA OF THE LONDON CLAY.

a Gavial I have seen thirteen dorsal and two lumbar; in that of a Crocodilus cata-
pkractus twelve dorsal and three lumbar; in those of a Crocodilus acutus, and Alligator
luctus, eleven dorsal and four lumbar, and this is the most common number; but in
the skeleton of the Crocodile, I believe of the species called Croc. biporcatus, described
by Cuvier,* he gives five as the number of the lumbar vertebrae. But these varieties
in the development or coalescence of the stunted pleurapophysis are of little essential
moment; and only serve to show the artificial character of the ‘dorsal’ and ‘ lumbar’
vertebree. The coalescence of the rib with the diapophysis obliterates of course the
character of the ‘costal articular surfaces ; which we have seen to be common to
both dorsal and cervical vertebree. The lumbar zygapophyses have their articular
surfaces almost horizontal, and the diapophyses, if not longer, have their antero-posterior
extent somewhat increased ; they are much depressed, or flattened horizontally.

The sacral vertebree are very distinctly marked by the flatness of the coadapted
ends of their centrums: there are never more than two such vertebre in the Crocodila
recent or extinct; in the first the anterior surface of the centrum is concave, in the
second it is the posterior surface; the zygapophyses are not obliterated in either of
these sacral vertebre, so that the aspects of their articular surface—upwards in the
anterior pair, downwards in the posterior pair—determines at once the corresponding
extremity of a detached sacral vertebra. The thick and strong transverse processes
form another characteristic of these vertebre ; for a long period the suture near their
base remains to show how large a proportion is formed by the pleurapophysis. This
element articulates more with the centrum than with the diapophysis developed from
the neural arch ; ¢ it terminates by a rough, truncate, expanded extremity, which almost
or quite joins that of the similarly but more expanded rib of the other sacral vertebree.
Against these extremities is applied a supplementary costal piece, serially homologous
with the superadded piecs to the proper pleurapophysis in the dorsal vertebree (/’, fig. 6,
p- 5, Part I), but here interposing itself between the pleurapophyses and heemapophyses
of both sacral vertebrae, not of one only. This intermediate pleurapophysial piece is
called the ‘ ilium ;’ it is short, thick, very broad, and subtriangular, the lower truncated
apex forming with the connected extremities of the heemapophysis an articular cavity for
the diverging appendage, called the ‘hind leg. The hemapophysis of the anterior
sacral vertebra is called ‘pubis; it is moderately long and slender, but expanded
and flattened at its lower extremity, which is directed forwards towards that of its
fellow, and joined to it through the intermedium of a broad, cartilaginous, heemal spine,
completing the hemal canal. The posterior hemapophysis is broader, subdepressed,
and subtriangular, expanding as it approaches its fellow to complete the second heemal

* Tom. cit., p. 95. It is to be observed that Cuvier begins to count the dorsal vertebra when the rib has
changed its hatchet-shape for a styloid shape.
+ Cuvier, who well describes this structure, remarks, ‘aussi méritent-elles plutét le nom des cétes que

celui d’apophvses transverses.”’ (Tom. cit., p. 98.)

CROCODILIA. 15

arch; it is termed ‘ischium.’ The great development of all the elements of these
hemal arches, and the peculiar and distinctive forms of those that have thereby
acquired, from the earliest dawn of anatomical science, special names, relates phy-
siologically to the functions of the diverging appendage which is developed into a
potent locomotive member. This limb appertains properly, as the proportion
contributed by the ischium to the articular socket and the greater breadth of
the pleurapophysis show, to the second sacral vertebra; to which the ilium chiefly
belongs.

The first caudal vert: va, which presents a ball for articulating with a cup on the
back part of the last sacral, retains, nevertheless, the typical position of the ball on the
back part of the centrum ; it is thus biconvex, and the only vertebra of the series which
presents that structure. I have had this vertebra in three different species of extinct
Eocene Crocodilia. In the Crocodilus toliapicus,T.1V, fig. 7; in the Croc. champsoides,
T. V, fig. 10; and in the Crocodilus Hastingsie, T. IX, fig. 7.

The advantage of possessing such definite characters for a particular vertebra 1s,
that the homologous vertebra may be compared in different species, and may yield
such distinctive characters as will be hereafier pointed out in those of the three species
above cited.

The first caudal vertebra, moreover, is distinguished from the rest by having no
articular surfaces for the hemapophyses, which in the succeeding caudals form a
hzemal arch, like the neurapophyses above, by articulating directly with the centrum.
The arch so formed has its base not applied over the middle of a single centrum, »u‘
like the neural arch in the back of the tortoise and sacrum of the bird, across the
interspace between twocentrums. The first heemal arch of the tail belongs, however, to
the second caudal vertebra, but it is displaced a little backwards from its typical
position.

The detached centrum of a caudal vertebra, besides being more slender and com-
pressed, is distinguished from those of the before-described vertebree by the two articular
surfaces at the posterior border of their under surface T. IV, fig.9. The zygapophyses
become vertical as far as the sixteenth or seventeenth, beyond which the two posterior
zygapophyses coalesce in an obiique plane notched in the middle, which is received
into a wider notch at the fore part of the neural arch of the succeeding vertebra. The
sutures between the pleurapophyses and diapophyses are maintained during a long
period of the animal’s growth, and demonstrate the share which these two elements
respectively take in the formation of the transverse process. So constituted, these
processes progressively decrease in length to the fifteenth or sixteenth caudal vertebra,
and then disappear. The neural spies progressively decrease in every dimension,
save length, which is rather increased as far as the twenty-second or twenty-third

vertebra, beyond which they }egin again to shorten, and finally subside in the terminal
vertebree of the tail.

16 FOSSIL REPTILIA OF THE LONDON CLAY.

The caudal heemapophyses coalesce at their lower or distal ends, from which a
spinous process is prolonged downwards and backwards; this grows shorter towards
the end of the tail, but is compressed and somewhat expanded antero-posteriorly.
The hzemal arch so constituted has received the name of ‘ chevron bone.’

A side view of the body of a middle caudal vertebra of the Crocodilus toliapicus is
given in T. III, fig. 8, and an under view of the same in fig. 9, showing the two
hypapophysial ridges extending from the articular facets for the hemapophyses at
one end to the other end of the centrum.

The segments of the endo-skeleton composing the skull are more modified than
those of the pelvis ; but just as the vertebral pattern is best preserved in the neural
arches of the pelvis, which are called collectively ‘sacrum,’ so, also, is it in the same
arches of the skull, which are called collectively ‘cranium.’ The elements of which these
cranial arches are composed preserve, moreover, their primitive or normal individuality
more completely than in any of the vertebree of the trunk, except the atlas, and
consequently the archetypal character can be more completely demonstrated.

In fossil Crocodilia, and many other reptiles, the bones of the head are very lable
from this cause to a greater extent of dislocation and separation than happens to the
skull of the warm-blooded animal, in which a greater proportion of those primitive
bones coalesce with age. It not unfrequently happens that detached bones of the
skull of a reptile are found fossil, and the usually much modified form of these vertebral
elements renders their determination difficult. In order to diminish this difficulty,
especially as the bones of the cranium are least familiar to the palzeontologist in their
detached state, I have subjoined a side view of them, fig. 9, nearly as they are arranged
in the formation of the successive natural segments of the skull. Such figures are the
more necessary in the present state of anatomy and paleontology, since the illustrations
of the osteology of the crocodile which have hitherto been prefixed to the descriptions
of the fossil remains of the Reptilian class, as, e. g., in the great work of Cuvier,
include only figures of the bones in question as they are naturally combined together in
the entire skuil.

For the anatomical description and determination of the individual bones, as con-
stituent elements of the vertebral segments of the head, I must refer the reader to my
work ‘On the Archetype of the Vertebrate Skeleton, pp. 115-25, figs. 18-21, and
pl. 2, fig. 3; and here limit myself to an ex2mplification of the natural arrangement
and names of the bones according to the letters and numbers in figure 9.

The bones of the head of the Crocodiles, as of all other vertebrate animals, are
primarily classified into those of
The ENDO-SKELETON,
The SPLANCHNO-SKELETON, and
The Ex0-SKELETON.

CROCODILIA. 17

Fig. .9

Disarticulated bones of the Skull of an Alligator, N 1 to rv the neural arches ; H1 to Iv the hemal arches
and appendages.

The bones of the endo-skeleton of the head form naturally four segments, called

Occipital vertebra, N 1, H1;

Parietal vertebra, N 11, H11;

Frontal vertebra, N 11, H 111;

Nasal vertebra, N tv, H tv.

These segments are subdivided into the neural arches, called

Epencephalic arch (1 basioccipital, 2 exoccipital, 3 superoccipital, 4 connate
paroccipital) ;

Mesencephalic arch (5 basisphenoid, 6 alisphenoid, 7 parietal, s mastoid) ;

Prosencephalic arch (9 presphenoid, 10 orbitosphenoid, 11 frontal, 12 post-
frontal) ;

Rhinencephalic arch (13 vomer, 14 prefrontal, 15 nasal) :

and into the hzemal arches and their appendages, called

Maxillary arch (20 palatine, 21 maxillary, 22 premaxillary) and appendages
(24 pterygoid, 24° ectopterygoid, 26 malar, 27 squamosal) ;

Mandibular arch (28 tympanic, 29—32 mandible) ;

Fig. 9.

18 FOSSIL REPTILIA OF THE LONDON CLAY.

Hyoidean arch (39 epihyal, 40 ceratohyal, 41 basihyal) ;
Scapular arch (50 suprascapula, 51 scapula, 52 coracoid) and appendages
(53—58 bones of fore-limb).
The bones of the splanchno-sheleton, are
The petrosal (16) and otosteals (16’) ;
The sclerotals (17) which in most retain their primitive histological condition
as fibrous membrane.
The turbinals (18 and 19) and teeth.
The bones of the exo-skeleton, are
The lacrymals (73).
The superorbitals (present in A/iigator sclerops).

To the foregoing brief analysis of the constituent parts of the framework of the
Crocodilia, which are petrifiable or conservable in a fossil state, and from the study
and comparison of which we have to gain our insight into the nature and affinities of
the extinct Reptiles, it seems here only requisite to add a few observations on the
characteristic mode in which the bones are associated together in certain parts of the
skeleton in the present order, and especially in the skull.

With regard to the trunk, the Crocodilia are distinguished from the Lacertilia and
from all other existing orders of Reptiles, by the articulation of the vertebral ribs
(pleurapophyses) in the cervical and anterior part of the dorsal segments by a head
and tubercle to a parapophysis and diapophysis. As this double joint is associated
with a double ventricle of the heart, and as the single articulation of every rib in other
Reptiles is associated with a single ventricle of the heart, we may infer a like difference
in.the structure of the central organ of circulation in the extinct reptiles, manifesting
the above-defined modifications in the proximal joints of the ribs.

The sacrum consists of two vertebre only, in Crocodilia as in Lacertiha: they are
modified in the present order, as before described, p. 14.

The skull consists, as above defined, of four segments. The hinder or occipital
surface of the skull presents, in the Crocodiia as in the Lacertilia, a single convex occipital
condyle, formed principally by the basioccipital, and not showing the trefoil character
which it bears in the Chelonia (Part I, T. XV, fig. 4), in which the exoccipitals con-
tribute equal shares to its formation. In the Batrachia, the exoccipitals exclusively form
the joint with the atlas, and there are accordingly two condyles. The occipital region
of the crocodilian skull is remarkable for its solidity and complete ossification, and for
the great extent of the surface which descends below the condyle. (T. VI, fig. 2.)
In the Zacertilia, a wide vacuity is left between the mastoid, exoccipital, and par-
occipital: but in the Crecodilia this is reduced to the small depressions or foramina
near 3, fig. 2, T. VI. The tympanic pedicles (28) extend outwards and downwards,
firmly wedged between the paroccipital, mastoid, and squamosal; in the Lacertians

CROCODILIA. 19

these pedicles are suspended vertically from the point of union of the mastoid and
paroccipital.

The chief foramen in the occipital region is that called ‘foramen magnum’ (between
2 and 2, in fig. 2, T. VI), through which the nervous axis is continued from the skull.
On each side of the foramen magnum isa small hole, called ‘ precondyloid foramen,’ for
the exit of the hypoglossal nerve. External to this is a larger foramen, marked x in
fig. 2, for the transmission of the nervus vagus and a vein. Below this is the
‘carotid foramen’ c. All these are perforated in the exoccipital. Below the condyle
there is usually a foramen, and sometimes two, for the transmission of blood-vessels.
Lower down, at the suture between the basioccipital and basisphenoid, is a larger and
more constant median foramen, indicated by the dotted line from e ¢; it is the bony
outlet of a median system of eustachian tubes, peculiar to the Crocodilia. On each
side of the median eustachian foramen, and in the same suture, is a smaller foramen,
which is the bony orifice of the ordinary lateral eustachian tube. The membranous
continuations of the lateral eustachian tubes unite with the shorter continuation from
the median tube, and all three terminate by a common valvular aperture, upon the
middle line of the faucial palate, behind the posterior or palatal nostril. The large,
bony aperture of this nostril is formed by the pterygoids (24 in fig. 2). The carotid
canal, c, opens by a short bony tube into the tympanic cavity, and is described as the
‘eustachian canal’ in the ‘Lecons d’Anatomie comparée’ of Cuvier. The artery
crosses the tympanic cavity, and enters a bony canal at its fore part, which conducts
to the ‘sella turcica’ in the interior of the cranium.

The median eustachian foramen is described by Cuvier as the ‘arterial foramen,*
the canal from which divides and terminates in the ‘sella turcica. | By MM. Bronn,
Kaup, and De Blainville, the median eustachian foramen is contended to be the bony
aperture of the posterior nostrils.

The results of the dissections and injections of recent Crocodiles and Alligators, by
which I have been able to rectify the discrepant opinions regarding the carotid,
eustachian, and naso-palatal foramina, and which have led to the discovery of a third
median eustachian canal, or rather system of canals, between the tympanic cavities and
fauces, peculiar to the Crocodilian Reptiles, are given in detail in the ‘ Philosophical
Transactions’ for 1850. The complexity of the superadded system has doubtless
chiefly contributed to mislead the justly-esteemed authorities who have believed that
they saw in it characters of the carotid canals or of the posterior nasal passages. The
eustachian apparatus in the Crocodilia may be briefly described as follows: From the
floor of each tympanic cavity two air-passages are continued ; the canal from the fore part
of the cavity extends downwards, backwards, and inwards, in the basisphenoid, which

* Ossemens Fossiles, tom. v, pt. i, p. 133.
falb= pe Ae.
¢t Abhandlungen iiber die Gavialartigen Reptilien der Lias-formation, folio, 1841, pp. 12, 16, 44.

20 FOSSIL REPTILIA OF THE LONDON CLAY.

unites with its fellow from the opposite tympanum, to form a short median canal, which
descends backwards to the suture between the basisphenoid and the basioccipital, where
it joins the median canal formed by the union of the two air-passages from the back
part of the floor of the tympanum, which traverse the basioccipital. ‘The common canal
formed by the junction of the two median canals descends along the suture to the
median foramen e/, fig. 2, T. VI. The air-passage from the back part of the tym-
panum, which traverses the basioccipital, swells out into a rhomboidal sinus in its
convergent course towards its fellow, and from this sinus is continued the normal
lateral eustachian canal, which, on each side, terminates below in the small aperture,
external to the median eustachian foramen.

That part of the outer surface of the skull which is covered by the common
integument is more or less sculptured with wrinkles and pits in the Crocodila: the
modifications of this pattern are shown in T. I, fig. 1, in the nilotic Crocodile, and
in T. VI, in the eocene Crocodile from Hordwell. The flat platform of the upper
surface of the cranium is perforated by two large apertures, surrounded by the bones
numbered 7, 8, 11, 12; these apertures are the upper outlets of the temporal fossa,
divided from the lower and lateral outlets by the conjoined prolongations of the
mastoid s and postfrontal 12: if ossification were continued thence to the parietal 7,
the temporal fossze would be roofed over by bone, as in the Chelones. In old
Crocodiles and Alligators there is an approximation to this structure, and the upper
temporal apertures are much diminished in size. In the Gavials (T. XI, fig. 1 a) they
remain more widely open, and, in the fossil Gavials of the secondary strata, they are still
wider, as seen in T. XI, fig. 2a; by which the structure of the cranium approaches more
nearly to that of the Lacertian reptiles, where the temporal fossa is either not divided
into an upper and lateral outlet, or is bridged over by a very slender longitudinal bar
from the postfrontal to the mastoid. The lateral outlets of the temporal fossee (T. VI,
fig. 1) are divided from the orbits by a bar of bone developed from the postfrontal (12)
and malar (26), and against the inner side of the base of which the ectopterygoid
abuts; the posterior boundary of the fossa is made by the tympanic (28) and squamosal
(27). The orbits, having the postfronto-malar bar (12, 26) behind, are surrounded in
the rest of their circumference by the frontal (11), the prefrontal (14), the lachrymal (73),
and the malar (26). The supraorbital or palpebral ossicle is rarely preserved in fossil
specimens.

The facial or rostral part of the skull anterior to the orbit, is of great extent, broad
and flat in the Alligators and some Crocodiles, narrower, rounder, and longer in other
Crocodiles, always most narrow, cylindrical, and elongated in the Gavials. The
anterior or external nostril is single, and is perforated in the middle of the anterior
terminal expansion of the upper jaw. This expansion is least marked in the broad-
headed species (compare T. VI, fig. 1, with T. III, fig. 1); in existing Crocodiles
and Alligators the points of the nasal bones penetrate its hind border, as at 15, fig. 1,

CROCODILIA. ZI

T. I. In the Gavials (T. XI, fig. 1a@) the nasals (w) terminate a long way from the
nostril. The Crocodilia resemble the Chelonia in the single median nostril.* In the
Lacertiia there is a pair of nostrils, one on each side the median plane, which is
occupied by a bridge of bone extending from the usually single premaxillary to the
nasals. The plane of the single nostril is almost horizontal in all existing and tertiary
Crocodilia.

On the inferior or palatal surface of the skull (T. VII, fig. 2), the most anterior
aperture is the circular prepalatal foramen surrounded by the premaxillaries 22; then
follows an extensive smooth, horizontal, bony plate, formed by the premaxillaries (22),
the maxillaries (21), and the palatines (20). The postpalatal apertures are always large
in the Crocodilia, and are bounded by the palatines (20), maxillaries (21), pterygoids (24),
and ectopterygoids (25). The posterior aperture of the nostril is formed wholly by the
pterygoids; it is shown in T. VI, fig. 3, between the bones marked 24. Behind it is
the median and lateral eustachian foramen already described, as belonging rather to
the posterior region of the head.

The scapulo-coracoid arch, both elements (fig. 9, 51, 52) of which retain the form of
strong and thick vertebral and sternal ribs in the crocodile, is applied in the skeleton of
that animal over the anterior thoracic hemal arches (T. XI). Viewed as a more robust
heemal arch, it is obviously out of place in reference to the rest of its vertebral segment.
If we seek to determine that segment by the mode in which we restore to their centrums
the less displaced neural arches of the antecedent vertebre of the cranium or in the
sacrum of the bird,t we proceed to examine the vertebre before and behind the dis-
placed arch, with the view to discover the one which needs it, in order to be made
typically complete. Finding no centrum and neural arch without its pleurapophyses
from the scapula to the pelvis, we give up our search in that direction; and in the
opposite direction we find no vertebra without its ribs until we reach the occiput :
there we have centrum and neural arch, with coalesced parapophyses—the elements
answering to those included in the arch N1, fig. 9—but without the arch H1; which
arch can only be supplied, without destroying the typical completeness of antecedent
cranial segments, by a restoration of the bones 50—52, to the place which they
naturally occupy in the skeleton of the fish. And since anatomists are generally
agreed to regard the bones 50—s2 in the crocodile as specially homologous with
those so numbered in the fish,f we must conclude that they are likewise homologous
im a higher sense; that in the fish, the scapulo-coracoid arch is in its natural or
typical position, whereas in the crocodile it has been displaced for a special purpose.
Thus, agreeably with a general principle, we perceive that, as the lower vertebrate

* Ina skeleton of the Alligator lucius in the Museum of the Royal College of Surgeons, a slender bar
of bone is continued from the nasals to the premaxillary, across the median nasal aperture, as it is in the
skull of the same species figured in the ‘Ossemens Fossiles,’ tom. v, pt. ii, pl. i, fig. 8.

+ See ‘On the Archetype and Homologies of the Vertebrate Skeleton,’ p. 117, p. 159.

PeOpwcits tesa, pe Lz.

U2) FOSSIL REPTILIA OF THE LONDON CLAY.

animal illustrates the closer adhesion to the archetype by the natural articulation of the
scapulo-coracoid arch to the occiput, so the higher vertebrate manifests the superior
influence of the antagonising power of adaptive modification by the removal of that arch
from its proper segment.

The anthropotomist, by his mode of counting and defining the dorsal vertebrae and
ribs, admits, unconsciously perhaps, the important principle in general homology
which is here exemplified, and whicl.. pursued to its legitimate consequences and
further applied, demonstrates that the scapula is the modified rib of that centrum and
neural arch which he calls the ‘occipital bone,’ and that the change of place which
chiefly masks that relation (for a very elementary acquaintance with comparative
anatomy shows how little mere form and proportion affect the homological characters
of bones) differs only in extent and not in kind from the modification which makes a
minor amount of comparative observation requisite, in order to determine the relation
of the shifted dorsal rib to its proper centrum in the human skeleton.

With reference, therefore, to the occipital vertebra of the crocodile, if the com-
paratively well-developed and permanently distinct ribs of all the cervical vertebre
prove the scapular arch to belong to none of those segments, and, if that heemal arch be
required to complete the occipital segment, which it actually does complete in fishes, then
the same conclusion must apply to the same arch in other animals, and we must regard
the occipital vertebra of the tortoise as completed below by its scapulo-coracoid arch
and not, as Bojanus supposed, by its hyoidean arch.*

Having thus endeavoured to show what the scapular arch of the crocodile is, I
proceed to point out the characteristic form of its chief elements. The upper and
principal part of the scapula (51, fig. 9) is flattened, and gradually becomes narrower to
the part called its neck, which is rounded, bent inwards, and then suddenly expanded to
form a rough articular surface for the coracoid, and a portion of a smoother surface for the
shoulder-joint. The contiguous end of the coracoid (52) presents a similar form, having
not only the rough surface for its junction with the scapula, but contributing, also, one
half of the cavity for the head of the humerus. It is perforated near the interspace
between these two surfaces. As it recedes from them it contrac's, then expands and
becomes flattened, terminating in a somewhat broader margin than the base of the

* Anatome Testudinis Europea, fol., 1819, p. 44. Geoffroy St. Hila:ve selected the opercular and sub-
opercular bones to form the inverted arch of his seventh (occipital) cranial vertebra, and took no account of
the instructive natural connexions .nd relative position of the hyoidean and scapular arches in fishes. With
regard to the scapular arch, he alludes to its articulation with the skull in the lowest of the vertebrate
classes as an ‘amalgame inattendue’ (Anatomie Philosophique, p. 481): and elsewhere describes it as a
“disposition veritablement trés singuliére, et que le manque absolu du cou et une combiuaison des pieces
du sternum avec celles de la téte pouvoient seules rendre possible.” —-Annales du Muséum, ix, p. 361. Adue
appreciation of the law of vegetative uniformity or repetition, and of the ratio of its prevalence and power to
the grade of organization of the species, was, perhaps, essential in order to discern the true signitication of
the connexion of the scapular arch in fishes.

CROCODILIA. 23}

scapula, which margin is morticed into a groove at the anterior border of the broad
rhomboidal cartilage continued beyond the ossified part of the manubrium, which forms
the key-bone of the scapular arch. The anterior locomotive extremity is the
diverging appendage of the arch, under one of its numerous modes and grades of
- development.*

The proximal element of this appendage or that nearest the arch, is called the
‘humerus’ (53, fig. 9): its head is subcompressed and convex; its shaft bent in two
directions, with a deltoid crest developed from its upper and fore part; its distal end
is transversely extended, and divided anteriorly into two condyles. The shaft of
this bone has a medullary cavity, but relatively smaller than in the mammalian
humerus.

The second segment of the limb consists of two bones: the larger one (54) is called
the ‘ulna:’ it articulates with the outer condyle of the humerus by an oval facet, the
thick convex border of which swells a little out behind, and forms a kind of rudimental
‘olecranon ;’ the shaft of the ulna is compressed transversely, and curves slightly out-
wards ; the distal end is much less than the proximal one, and is most produced at the
radial side.

The radius (55) has an oval head; its shaft is cylindrical; its distal end oblong and
subcompressed.

The small bones (56) which intervene between these and the row of five longer
bones, are called ‘ carpals: they are four in number in the Crocodilia. One seems to
be a continuation of the radius, another of the ulna; these two are the principal
carpals; they are compressed in the middle and expanded at their two extremities ;
that on the radial side of the wrist is the largest. A third small ossicle projects
slightly backwards from the proximal end of the ulnar metacarpal: it answers to the
bone called ‘pisiforme’ in the human wrist. The fourth ossicle is interposed between
the ulnar carpal and the metacarpals of the three ulnar digits. .

These five terminal jointed rays of the appendage are counted from the radial to
the ulnar side, and have received special names: the first is called ‘ pollex,’ the second
‘index, the third ‘ medius,’ the fourth ‘annularis,’ and the fifth ‘minimus.’ The first
joint of each digit is called ‘metacarpal;’ the others are termed ‘phalanx.’ In the
Crocodilia the pollex has two phalanges, the index three, the medius four, the annularis
four, and the minimus three. The terminal phalanges, which are modified to support
claws, are called ‘ ungual’ phalanges.

As the above-described bones of the scapular extremity are developments of the
appendage of the scapular arch, which is the hemal arch of the occipital vertebra, it
follows, that, like the branchiostegal rays and opercular bones in fishes, they are
essentially bones of the head.

The diverging appendage of the pelvic arch being a repetition of the same element

* See my Discourse ‘ On the Nature of Limbs,’ 8vo, Van Voorst, 1849, pp. 64-70.

24 FOSSIL REPTILIA OF THE LONDON CLAY.

as the appendage of the scapular arch modified and developed for a similar office,
a close resemblance is maintained in the subdivisions of the framework of both limbs.
The first bone of the pelvic limb, called the ‘femur,’ is longer than the humerus, and,
like it, presents an enlargement of both extremities, with a double curvature of the
intervening shaft, but the directions are the reverse of those of the humerus, as may be
seen in T. XI, where the upper or proximal half of the femur is concave, and the distal
half convex, anteriorly. The head of the femur is compressed from side to side, not
from before backwards as in the humerus; a pyramidal protuberance from the mner
surface of its upper fourth represents a ‘trochanter; the distal end is expanded
transversely, and divided at its back part into two condyles.

The next segment or ‘leg,’ includes, like the corresponding segment of the forelimb
called ‘forearm,’ two bones. The largest of these is the ‘tibia,’ and answers to the
radius. It presents a large, triangular head to the femur; it terminates below by an
oblique crescent with a convex surface.

The ‘ fibula’ is much compressed above; its shaft is slender and cylindrical, its
lower end is enlarged and triangular.

All these long bones have a narrow medullary cavity.

The group of small bones which succeed those of the leg, are the tarsals; they are
four in number, and have each a special name. The ‘astragalus’ articulates with the
tibia, and supports the first and part of the second toe. It is figured in Cuvier’s
‘Ossemens Fossiles,’ tom. v, pt. ii, pl. iv, figs. 19 4, B, C, D. The ‘calcaneum’ inter-
venes between the fibula and the ossicle supporting the two outer toes; it has a short
but strong posterior tuberosity.

The ossicle referred to represents the bone called ‘cuboid’ in the human tarsus.
A smaller ossicle, wedged between the astragalus and the metatarsals of the second
and third toes is the ‘ ectocuneiform.’

Four toes only are normally developed in the hind-foot of the Crocodiha ; the fifth
is represented by a stunted rudiment of its metatarsal, which is articulated to the
cuboid and to the base of the fourth metatarsal.

The four normal metatarsals are much longer than the corresponding metacarpals.
That of the first or innermost toe is the shortest and strongest; it supports two
phalanges. The other three metatarsals are of nearly equal length, but progressively
diminish in thickness from the second to the fourth. The second metatarsal supports
three phalanges ; the third four; and the fourth also has four phalanges, but does
‘not support a claw. The fifth digit is represented by a rudiment of its metatarsal in
the form of a flattened triangular plate of bone, attached to the outer side of the
cuboid, and slightly curved at its pointed and prominent end.

In the skull of the Crocodile, as of most other vertebrates, there are intercalated
a few bones, or ossified parts of special organs, which, as is shown in the classed
Table of the bones of the head, do not belong to the vertebral system of bones.

CROCODILIA. 25

The bone anterior to the orbit, marked 73 in fig. 9, and in T. I and T. VI is perforated
at its orbital border by the duct of the lachrymal gland, whence it is termed the
‘lachrymal bone, and its facial part extends forwards between the bones marked
14, 15, 21, and 26 in the same plates. In many Crocodilia there is a bone at the
upper border of the orbit, which extends into the substance of the upper eyelid; it is
called ‘ superorbital.’ In the Crocodilus palpebrosus there are two of these ossicles.

Both the lachrymal and superorbital bones answer to a series of bones found com-
monly in fishes, and called ‘ suborbitals’ and ‘superorbitals.’ The lachrymal is the
most anterior of the suborbital series, and is the largest in fishes; it is also the most
constant in the vertebrate series, and is grooved or perforated by a mucous duct.
These ossicles appertain to the dermal or muco-dermal system or ‘ exoskeleton.’

The little slender bone, marked 16’ in fig. 9, has one of its extremities in the form
of a long, narrow, elliptic plate, which is applied to the ‘ feuestra ovalis’ of the internal
ear; from this plate extends a long and slender bony stem, which grows somewhat
cartilaginous, expands and bends down, as it approaches the tympanum or ear-drum, to
which it is attached.

The cartilaginous capsule of the labyrinth or internal ear is partially ossified
by sinuous plates of bone connate with the neurapophyses (fig. 10, 2 and 6), between
which that organ is lodged; I apply the term ‘ petrosal’ to the principal and most
independent of those ossifications of the ear-capsule, to that, e. g., which retains
some mobility after it has con-
tracted a partial anchylosis to
the exoccipital (2), and which
appears upon the inner surface
of the cranial walls at the part
marked 16 in the subjoined
Cut 10, between 2 and 6. It
is the only independent bone
on that surface of the cranium
which, in my opinion, answers
to the ‘ petrous portion of the
temporal’ in human anatomy,
and to which the term ‘rocher
can be properly applied, in the language of the French comparative anatomists. Cuvier,
however, restricts that name to the ‘alisphenoid’ (6, figs. 9, 10) in the Crocodiles.

The ossicles, (16 and 16’, fig. 9), together with the partial ossifications in the sclerotic
capsule of the organ of sight, (17, fig. 9)—always more distinct in Chelonia than in
Crocodilia— belong to that category of visceral bones to which the term ‘ splanchno-
skeleton’ has been given; they also are foreign to the true vertebrate system of the
skeleton.

Vertical longitudinal section of the cranium of a Crocodile
(Crocodilus acutus).

4

26 FOSSIL REPTILIA OF THE LONDON CLAY.

The teeth——The most readily recognisable character by which the existing
Crocodilians are classified and grouped in appropriate genera, are derived from
modifications of the dental system.

In the Caimans (genus Alligator) the teeth vary in number from ao to et
the fourth tooth of the lower jaw is received into a cavity of the alveolar surface of the
upper Jaw, where it is concealed when the mouth is shut. In T. VIII, fig. 2, these
pits are shown behind the last premaxillary tooth e, in an Eocene Alligator from
Hordwell. In old individuals of the existing species of Alligator, the upper jaw is
perforated by the large inferior teeth in question, and the fossz are converted into
foramina.

In the Crocodiles (genus Crocodilus) the fourth tooth in the lower jaw is received
anto a notch excavated in the side of the alveolar border of the upper jaw, as in fig. 1,
T. VIII, behind the tooth e, and is visible externally when the mouth is closed, as in
T. VII, fig. 1. In most Crocodiles, also, the first tooth in the lower jaw perforates
the premaxillary bone when the mouth is closed, as in T. I, between the teeth
marked a and 6.

In the two preceding genera the alveolar borders of the jaw have an uneven or
wavy contour, and the teeth are of an unequal size.

In the Gavials, (genus Gavialis) the teeth are nearly équal in size and similar in
form in both jaws, and the first as well as the fourth tooth in the lower jaw, passes
into a groove in the margin of the upper jaw when the mouth is closed, T. XI.

In the Alligators and Crocodiles the teeth are more unequal in size, and less regular
im arrangement, and more diversified in form than in the Gavials: witness the strong
thick conical laniary teeth at the fore part of the jaw, as shown in T. VII and T. III, fig. 6,
as contrasted with the blunt mammillate summits of the posterior teeth, as shown in
T. V, fig. 12. The teeth of the Gavial are subequal, most of them are long, slender,
pointed, subcompressed from before backwards, with a trenchant edge on the right
and left sides, between which a few faint longitudinal ridges traverse the basal part of
the enamelled crown.

The teeth of both the existing and extinct Crocodilian reptiles consist of a body of
compact dentine forming a crown covered by a coat of enamel, and a root invested by
a moderately thick layer of cement. The root slightly enlarges, or maintains the same
breadth to its base, which is deeply excavated by a conical pulp-cavity extending
into the crown, and is commonly either perforated or notched at its concave or inner
side.

The dentinal tubes in the crown of a fully-developed tooth form short curvatures at
their commencement at the surface of the pulp-cavity, and then proceed nearly straight
to the periphery of the crown; they very soon bifurcate, the divisions slightly
diverging; then continuing their course with gentle parallel undulations, they

CROCODILIA. 27

subdivide near the enamel, and terminate in fine and irregular branches, which
anastomose generally by the medium of cells. The dentinal tubes send off from both
sides, throughout their progress, minute branches into the intervening substance, and
terminate in the dentinal cells. These cells are subhexagonal, about ;+, of an
inch in diameter, and are traversed by from ten to fourteen of the dentinal tubes; they
are usually arranged in planes parallel with the periphery of the crown, near which
they are most conspicucus, and towards which their best defined outline is directed:
they combine with the rarallel curvatures of the dentinal tubes to form the striae,
visible in sections of the teeth by the naked eye, which cause the stratified appearance
of the dentine as if it were composed of a succession of superimpos2:d cones. The
diameter of the dentinal tube before the first bifurcation is ~,1,,th of an inch,
both the trunks and bifurcations of the tubes have interspaces equal to four of their
respective diameters.

The enamel viewed in a transverse section of the crown presents some delicate
strie parallel with its surface, whilst the appearance of fibres vertical to that surface
is only to be detected, and these faintly, on the fractured edge. It is a very compact
and dense substance; the dark brownish tint is strongly marked in the middle of the
enamel when viewed by transmitted light.

The cells with which the fine tubes of the basal cement communicate, are oblong,
about =1,,th of an inch across their long axis, which is transverse to that of the tooth ;
the inter-communicating tubes, which radiate from the cells, giving them a stellate figure.
I have entered into these particulars of the microscopic texture of the teeth of the
Crocodile because it will be seen in the sequel that important modifications of
the dental tissues characterise some of the extinct Repfilia.

In the black Alligator of Guiana the first fourteen teeth of the lower jaw are
implanted in distinct sockets, the remaining posterior teeth are lodged close together
in a continuous groove, in which the divisions for sockets are faintly indicated
by vertical ridges, as in the jaws of the Ichthyosaurs. A thin compact floor of bone
separates this groove, and the sockets anterior to it, from the large cavity of the ramus
of the jaw; it is pierced by blood-vessels for the supply of the pulps of the growing
teeth and the vascular dentiparous membrane which lines the alveolar cavities.

The tooth-germ is developed from the membrane covering the angle between
the floor and the inner wall of the socket. It becomes in this situation completely
enveloped by its capsule, and an enamel-organ is forined at the inner surface of
the capsule before the young tooth penetrates the interior of the pulp-cavity of its

predecessor.

The matrix of the young growing tooth affects, by its pressure, the mner wall of
the socket, and forms for itself a shallow recess; at the same time it attacks the side
of the base of the contained tooth; then, gaining a more extensive attachment by its
basis and increased size, it penctrates the large pulp-cavity of the previously formed

28 FOSSIL REPTILIA OF THE LONDON CLAY.

tooth, either by a circular or semicircular perforation. The size of the calcified part
of the tooth matrix which has produced the corresponding absorption of the previously
formed tooth on the one side, and of the alveolar process on the other, is represented
in the second exposed alveolus of the portion of jaw figured in Pl. 75, fig. 4, of my
‘Odontography,’ the tooth marked a in that figure, having been displaced and turned
round to show the effects of the stimulus of the pressure. The size of the perforation
in the tooth, and of the depression in the jaw, proves them to have been, in great
part, caused by the soft matrix, exciting dissolution and absorbent action, and not by
mere mechanical force. The resistance of the wall of the pulp-cavity having been thus
overcome, the growing tooth and its matrix recede from the temporary alveolar
depression, and sink into the substance of the pulp contained in the cavity of the fully-
formed tooth. As the new tooth grows, the pulp of the old one is removed; the old
tooth itself is next attacked, and the crown being undermined by the absorption of the
inner surface of its base, may be broken off by a slight external force, when the point
of the new tooth is exposed.

The new tooth disembarrasses itself of the cylindrical base of its predecessor, with
which it is sheathed, by maintaining the excitement of the absorbent process so long
as the cement of the old fang retains any vital connexion with the periosteum of the
socket; but the frail remains of the old cylinder, thus reduced, are sometimes lifted off
the socket upon the crown of the new tooth, when they are speedily removed by the
action of the jaws. This is, however, the only part of the process which is immediately
produced by mechanical force: an attentive observation of the more important pre-
vious stages of growth, teaches that the pressure of the growing tooth operates upon
the one to be displaced only through the medium of the vital dissolvent and absorbent
action which it has excited.

Most of the stages in the development and succession of the teeth of the Crocodiles
are described by Cuvier* with his wonted clearness and accuracy ; but the mechanical
explanation of the expulsion of the old tooth, which Cuvier adopts from M. Tenon, is
opposed by the disproportion of the hard part of the new tooth to the vacuity in the
walls of the old one, and by the fact that the matter impressing—viz. the uncalcified
part of the tooth-matrix—is less dense than the part impressed.

No sooner has the young tooth penetrated the interior of the old one, than another
germ begins to be developed from the angle between the base of the young tooth and
the inner alveolar process, or in the same relative position as that in which its imme-
diate predecessor began to rise, and the processes of succession and displacement
are carried on, uninterruptedly, throughout the long life of these cold-blooded
carnivorous reptiles.

From the period of exclusion from the egg, the teeth of the crocodile succeed each
other in the vertical direction; none are added from behind forwards, like the true

* Op. cit., pp. 90-3.

CROCODILIA. 29

molars in Mammalia. It follows, therefore, that the number of the teeth of the cro-
codile is as great when it first sees the ight as when it has acquired its full size; and,
owing to the rapidity of the succession, the cavity at the base of the fully-formed
tooth is never consolidated.

The fossil jaws of the extinct Crocodilians demonstrate that the same law regulated
the succession of the teeth at the ancient epochs when those highly organized reptiles
prevailed in greatest numbers, and under the most varied generic and specific modi-
fications, as at the present period, when they are reduced to a single family, composed
of so few and slightly varied species, as to have constituted in the Systema Nature of
Linnzus, a small fraction of the genus Lacerta.

CrocopiLus ToLtaPicus, Owen. Tab. Il, fig. 1, and Tab. II 2.

Syn. CrocopiLe DE Sueppy (?), Cuvier. Ossemens Fossiles, 4to, tom. vy, pt. ii, p. 165.
CrocopiLus SpenceERI, Buckland. Bridgewater Treatise, vol. i, p. 251. ‘Crocodile
with a short and broad snout.” Vol. ii, p. 36, pl. 25’, fig. 1.
— —- Owen. Reports of the British Association, 1841, p. 65.

In proceeding to the comparison, and preparing for the description of the British
fossil Crocodilia, 1 endeavoured, in the first place, to obtain the bones of the species
which now exists in a locality nearest to Great Britain, and also of an individual of that
same species which had lived at a remote period; and I have been favoured by the
kindness of my esteemed friend Philip Duncan, Esq., Fellow of New College, Oxford, and
Conservator of the Ashmolean Museum, with the opportunity of examining the bones of a
mummified Crocodile from a sarcophagus at Thebes, in that collection at Oxford. Two
views of the skull of this old Egyptian Crocodile are given in T. I. The total length
of the skull from the bone marked 2s to the end of 22, is twice the breadth of the back
part of the skull. The upper apertures of the temporal fossee are subcircular ; the point of
the squamosal (27) projects into the lateral aperture. The breadth of the back part of the
sculptured cranial platform (8,8), is less by one fourth than the breadth of the skull anterior
to the orbits. The breadth of the interorbital space is nearly equal to the transverse
diameter of the orbit. The points of the nasals (15) project into the external nostril.
The postpalatal apertures reach as far forwards as the seventh tooth, counting from
the hindmost; there are nineteen alveoli on each side of the upper jaw, the five
anterior teeth being lodged in the premaxillary, which is perforated by the first tooth
of the lower jaw.

Geoffroy St. Hilaire has applied the old Egyptian name Zovyoc to the mummified
Crocodiles of that country; but there is no good specific character which distinguishes
them from the modern Crocodiles of the Nile, to which Cuvier has given the name of
Crocodilus vulgaris.

Cuvier appears to have first called the attention of paleontologists to the remains

30 FOSSIL REPTILIA OF THE LONDON CLAY.

of Crocodilia in the Eocene clay forming the Isle of Sheppy, in the last volume of the
second edition of his great work on the ‘Ossemens Fossiles,’ p. 165, 1824. He there
specifies a third cervical vertebra, which was obtained by M. G. A. Deluc, at Sheppy,
and of which Cuvier made a drawing at Geneva; he says it much resembles the
corresponding vertebra in one of our living Crocodiles, and might have come from an
individual about five feet in length. ‘“M. Deluc,” he adds, ‘found very near it a
much smaller vertebra, which I recognised as belonging to a monitor or some allied
genus.’ *

Our knowledge of the Eocene Crocodiles of Sheppy received a remarkable accession
at the publication of the highly imteresting and instructive ‘ Bridgewater Treatise’ of
Dr. Buckland, in which he states that “ true Crocodiles, with a short and broad snout,
like that of the Caiman and the Alligator, appear, for the first time, in strata of the
tertiary periods, in which the remains of mammalia abound. . . . One of these,” he adds,
‘found by Mr. Spencer in the London Clay of the Isle of Sheppy, is engraved Pl. 25’,
fig. ],” and the name ‘ Crocodilus Spencer?’ is appended to that figure.

In preparing my ‘ Report on British Fossil Reptiles’ for the British Association in
1841, I examined the original specimen figured by Dr. Buckland, in which unfortunately
the end of the snout with the intermaxillaries and an indeterminate proportion of the
maxillaries having been broken off and lost, no exact idea could be formed of the pro-
portions of the facial or rostral part of the skull.

In a larger specimen of the fossil skull of a Crocodile from Sheppy, in the British
Museum, T. II 4, the whole of the upper, as well as the lower jaw, were preserved, and
as the proportions of the snout agreed with those of some true Crocodiles, and differed
in an equal degree with those species from the Gavial; and as, like the Crocodiles and
Caimans, it presented the more important distinction of a different composition of that
part of the skull, I retained for the specimen in that ‘Report’ the name of Crocodilus
Spencert, proposed by the author of the Bridgewater Treatise for the Sheppy Crocodile,
so differing from the Gavial.

The able keeper of the Mineralogical Department of the British Museum, Charles
Konig, K.H., F.R.S., to whom I am indebted for every facility in describing and figuring
this specimen, has suggested that the name by which Baron Cuvier first indicated the
existence of a true Crocodile in the Eocene clay of Sheppy, should have the priority,
and I adopt, therefore, the name Crocodilus toliapicus, which he has attached to the
specimen in question, and with the more readiness since I have now reason to doubt
whether the mutilated cranium, figured in the ‘ Bridgewater Treatise,’ belongs to the
same species.

* Could this have been a vertebra of the large serpent, which I have subsequently described under the
name of Palzophis ? I have not as yet met with a single lacertian vertebra from Sheppy. If the collection
of M. Deluc be still preserved at Geneva, the vertebra in question might be compared with the figures of the
Paleophus toliapicus, ‘Ophidia,’ T. XV.

CROCODILIA. 31

The more entire fossil skull in question presents the following dimensions :

Feet. Inches. Lines.
Total length from the hindmost part of the lower jaw. 2 2 0

Breadth between the articular ends of the tympanics 0 10 0
Do. across the orbits 0 7 6
Do. of the intertemporal space 0 0 9
Do. of the interorbital space : : 0 1 4

From the articular end of the tympanic to the orbit 0 8 6

From the occipital condyle to the orbit . 0 7 0

From the orbit to the external nostril : 0 14 0

Breadth of the cranium five inches in advance of fe orbits 0 3 8
Do. across the external nostril ; : 0 2 8

Depth of the lower jaw at the vacuity between the angular

and surangular . : : 5 : : - 0 3 6

Length of that vacuity : 0 3 0

Breadth of the base of one of the lieder iaaxillaby eset 5 0 0 8

This remarkably fine fossil skull, which is figured one third of its natural size in
T. Il, fig. 1, presents proportions which come nearest to those of the Crocodilus
acutus, being longer in proportion to its basal breadth than in the Crocodilus Suchus, n
which the diameter between the articular ends of the tympanis (28) is just half the length
of the entire skull. The interorbital space in the Crocodilus toliapicus is relatively
narrower and flatter than in the Croc. acutus or Croc. Suchus, and the facial part of the
skull becomes narrower before the expansion of the upper jaw, at the figure 15, than it
does in either of those species. The narrow elongated nasals on which the figure 15 is
placed, extend forwards to the external nostril (22), as in the true Crocodiles, and the

alveolar border is festooned as is. shown in the side view in T. II 4. The teeth
22—22

e
20—20
than in the recent species above cited, and resemble in this respect the teeth of the
Crocodilus Schlegel of S. Miiller, which is from Borneo. The extent of the symphysis

of the lower jaw is greater in the Crocodilus toliapicus than in the Croc. acutus, and still

=84 in number: they are more uniform in size, and more regularly spaced

greater than in the Croc. Suchus; the Sheppy species in this respect more nearly
resembles the living species from Borneo above cited.

CROCODILUS CHAMPSOIDES, Owen. Tab. III. (Tab. IL, fig. 2’)

Syn. CxocopiLe DE Suepry (?), Cuvier. Loc. cit.
Crococitus Spencert, or “Crocodile with a short and broad snout”’ (?) Buckland.
Bridgewater Treatise, vol. ii, pl. xxv, fig. 1.

The fossil skull already described establishes the fact of the existence of a true
Crocodile in the London Clay at Sheppy, but not of a species with a short and broad
snout ; the present specimen equally demonstrates the presence at the earliest period
of the Tertiary geological epoch of Crocodilia with those modifications of the cranial

32 FOSSIL REPTILIA OF THE LONDON CLAY.

and dental structure on which the characters of the restricted genus Crocodilus of
modern Zoology are founded; but they are associated with a general form of the head
which approaches more nearly to the Gavials than does that of the Crocodilus toliapicus,
and which are most nearly paralleled amongst the known existing true Crocodiles by
the Crocodilus Schlegeht. This Bornean species was, in fact, originally described as a
new species of Gavial, but the nasal bones, as in the fossil from Sheppy figured in
T. II, 15, extend to the hind border of the exteraal nostril.

The fine subject of T. IIJ, forms part of the collection of J. S. Bowerbank, Esq.
F.R.S., which is well known for its rich and varied illustrations of the fossils of the
Isle of Sheppy.

The following are some of its admeasurements :

Feet. Inches. Lines.
Total length from the occipital condyle to the end of the

premaxillaries . 6 : 6 6 : : 1 4 0
Breadth across the hinder angles of the supracranial platform 0 4 0
Do. across the orbits 0 5 0
Do. of the intertemporal space 0 0 4
Do. of the interorbital space . 0 1 0
Do. across the external nostril : ; : , 0 2 0
From the occipital condyle to the orbit 0 3 4
From the orbit to the external nostril . : : : i) 10 0

The skull yielding the above dimensions is much smaller than that of the Urocodilus
tohapicus, T. IL; but it cannot have belonged to a younger individual of the same
species, because, in existing Crocodiles, the part of the skull anterior to the orbits is
proportionally shorter in the young than in the old individuals, as may be seen by -
comparing the figures which Cuvier has given of the skulls of three individuals of
different ages of the Croccdilus biporcatus, in figures 4, 18, and 19, of plate 1 of the last
volume of the ‘ Ossemens Fossiles,’ 4to, 1824; whereas the part of the skull anterior to
the orbits is relatively longer and more slender in the smaller fossil skull now described
than in the larger one on which the species Croc. toliapicus is founded. We have,
therefore, satisfactory proof that two species of true Crocodile existed during the de-
position of the Eocene Clay at the actual mouth of the Thames, and have left their
remains in that locality.

Their specific distinction is further illustrated by the different forms and propor-
tions of particular parts of the skull. The alveolar border is more nearly straight ; the
transverse expansion of the maxillaries (21) is less, whilst that of the premaxillaries
(22) is greater: the interorbital space is broader and more concave. The teeth are
more uniform in size, are more regularly spaced, and are wider apart : they are, likewise,
upon the whole, larger in proportion to the size of the jaw. Figure 5, T. II, shows
the crown of a new tooth just emerging from the second socket of the maxillary bone
ef the natural size ; figure 6 is the fourth tooth of the premaxillary, fully formed ; fig. 7

CROCODILIA. 33

is the displaced tooth which is cemented by the matrix to the palatal surface of the
premaxillary in fig. 2. The enamelled crown shows the fine raised longitudinal ridges
better developed than one usually sees them in modern Crocodiles. There are twenty-
one alveoli on each side of the upper jaw.

In all the particulars in which the skull under description differs from that of the
Crocodilus toliapicus, it departs further from the nilotic crocodile, and resembles more
the Gavial-like Crocodile of Borneo; and as one of the old Egyptian names of the
Crocodile, Champsa, has been applied generically to the Gavials by some recent
Erpetologists, I have adopted the term ‘ Champsoides’ to signify the resemblance of
the present extinct species of Eocene Crocodile to the Gavials.

The basioccipital condyle, together with the condyloid processes of the exoccipital,
project backwards in the Croc. champsoides farther than in any modern Crocodile; and
the supraoccipital 3, fig. 4, T. III, descends nearer to the foramen magnum.

The upper jaw is more depressed, and the suborbital part of the maxillary bone
is much less inclined to the vertical in the present skull than in the original of Dr.
Buckland’s figure of the Crocodilus Spencert, which in other respects more nearly
resembles the Croc. champsoides than the Croc. toliapicus; the difference above specified
seems to be greater than can be accounted for by any accidental pressure to which
the fossil skull figured in T. III can have been subjected. The mutilated skull to
which the term Croc. Spenceri was originally applied, is defective, as I have said, in the
facial or maxillary portion which is requisite for its unequivocal determination to
either of the two species which the more perfect specimens since acquired have
proved to have existed at the Eocene tertiary period. The form of the mutilated
portion of skull, and the figure of it given in Pl. 25’ of the ‘ Bridgewater Treatise,’ might
well appear to indicate a short and broad snouted species of true Crocodile; but if it
be not distinct from the two better represented species above described, I should be
more inclined to refer it to that which has the longest and narrowest snout, from the
conformity of the characters of the part of the skull which is preserved. A view of the
palatal surface of the specimen in question is given in T. II, fig. 2.

Crocodilian vertebre referable to the two foregoing species of Sheppy Crocodiles.

Not more than two species of Crocodile are indicated by the detached vertebree
from Sheppy; but the different proportions of the homologous cervical vertebre,
fig.3 and 7, T. V, and of the characteristic biconvex caudal vertebra, fig. 7, T. IV, and
fig. 10, T. V, would have determined the fact of there being two distinct species,
had their cranial characters, which are so satisfactorily demonstrated in T. II 4 and
T. III, remained unknown. I refer, provisionally, the shorter and thicker vertebre to
the Crocodilus tolapicus with the shorter and thicker snout, and the longer and thinner
vertebra to the Croc. champsoides with the snout of similar proportions.

5

34 FOSSIL REPTILIA OF THE LONDON CLAY.

Vertebre of the CROCODILUS TOLIAPIcUS, Tab. IV and Tab. V, fig. 1, 2, 3, 5, 6.

The vertebra, fig. 1, 2, T. V, is the fourth cervical; it differs from that of the
Crocodilus acutus, Croc. Suchus, and Croc. biporcatus, in the greater breadth and squareness
of the base of the hypapophysis (fig. 27%), which extends almost to the bases of the
parapophyses p ; the vertical diameter of the parapophyses is greater in comparison with
their antero-posterior extent in the fossil than in the above-cited recent Crocodiles ;
the neurapophyses are thicker, and terminate in a more rounded border both before
and behind; their bases extend inwards, and meet above the centrum, whilst a narrow
groove divides them in the recent Crocodiles above cited; the length of the centrum
is greater in proportion to the height and breadth in the fossil vertebra. In other
respects the correspondence is very close, and the modern crocoailian characters are
closely repeated. ‘Traces of the suture between the centrum and neurapophysis remain,
as shown at 2, 2, fig. 1. The diapophysis d, and the upper portion of the neural arch,
with the zygapophyses and neural spine, have been broken away ; the borders of the
articular ends of the centrum have been worn away.

The vertebra (fig. 3, T. V) is the sixth cervical: in this specimen the base of the
iypapophysis is contracted laterally and extended antero-posteriorly ; the side of the
centrum above the parapophysis (7) has become less concave; the vertebra has increased
more in thickness than in length; in these changes it corresponds with the modern
Crocodiles ; it has been mutilated and worn in almost the same manner and degree as
the fourth cervical.

The vertebra (fig. 1, 2, T. IV) is a seventh cervical of a smaller individual of the
Crocodilus toliapicus. The hypapophysis has become more compressed and more
extended antero-posteriorly ; the paraporhysis has become shortened antero-posteriorly,
and increased in vertical diameter. ‘The anterior concave surface of the centrum
(fig. 1) is more circular, less extended transversely, than in the corresponding vertebra
of the recent Crocodiles compared with the fossil.

Fig. 3, 4, T. IV, are two views of the eighth cervical of an individual of about
the same size as that to which the fourth and sixth cervicais in T. V belong. Fig. 4,
exemplifies the same difference which fig. | presents in regard to the more circular
contour of the anterior concave surface of the centrum as compared with recent
Crocodiles ; the bases of the neurapophyses are thicker and more rounded anteriorly ;
the neural canal is rather more contracted; the base of the hypapophysis more extended
in the axis of the vertebra (see fig. 3) than in the recent Crocodiles compared. The
parapophyses have now risen, as in those Crocodiles, to the suture of the neuvapophysis,
and the diapophysis springs ott at some distance above that suture.

Fig. 6, T.1V, shows the under surface of a dorsal vertebra, in which the
hypapophysis ceases to be developed (probably the fourth or fifth).

Fig. 5, T. IV, gives the same view of one of the lumbar vertebre, showing the

CROCODILIA. 35

elongation of the centrum, and the broad bases of the depressed diapophyses ; there is an
indication of two longitudinal risings towards the back part of the under surface of the
centrum.

Fig. 5 and 6, T. V, give two views of the anterior sacral vertebra of the Crocodilus
tohapicus ; it is concave and much expanded transversely at its fore part (fig. 5),
flattened and contracted behind. ‘Traces of the suture remain to show the proportion
of the anterior articular surface which is formed by the base uf the pleurapophysis p ;
and fig. 6 shows the extension of that base from the side of the centrum upon the
diapophysis or overhanging base of the neurapophysis; the under surface of the
centrum of this vertebra has a slight median longitudinal rising.

Fig. 7, T. IV, gives a side view of the characteristic, biconvex, anterior caudal
vertebra of the Crocodilus toliapicus.

Fig. 8, 9, T. IV, give two views of a middle caudal vertebra: in fig. 9 are shown
the characteristic hypapophysial ridges extending from the articular surfaces for the
heemapophyses at the hind part of that aspect of the centrum: in fig. 8 the processes
of the neural arch are restored in outline; a thick and low ridge extends from the
middle of the side of the centrum to the base of the transverse process which it
strengthens, like an underpropping buttress.

Vertebre of the CROCODILUS CHAMPSOIDES.

Fig. 7 and 8, T. V, give two views of the third cervical vertebra of the above-
named gavial-like Crocodile, which vertebra, besides its longer and more slender propor-
tions, differs in the smaller size of its hypapophysis from the corresponding vertebra in
any existing species of Crocodile or Gavial: the process in question being in the form of
a low crescentic ridge, as shown at figure 8, between the bases of the parapophyses (7).

Both parapophyses terminate by a convex surface, which appears to have been
anatural one. Between the parapophysis (p) and diapophysis (d), fig. 7, the side of the
centrum is more deeply excavated than in the Crocodilus toliapicus. The centrum
contributes a small part to the base of the diapophysis, as in the third cervical vertebra
of modern Crocodiles. The neurapophysis are thinner than in the Croc. toliapicus, and
their bases do not joi one another above the centrum. The longitudinal ridge
extending from the anterior to the posterior zygapophysis is sharply defined.

Fig. 4,T. V, is the first dorsal vertebra of the Crocodilus champsoides, in which, as
in existing Crocodiles, the parapophysis (7) has passed almost wholly from the centrum
upon the neurapophysis, the diapophysis (¢d) having been subject to a corresponding
ascent. ‘The base of the compressed hypapophysis extends over the anterior third of
the middle line of the under surface of the centrum. There is a remarkable transverse
constriction at the base of the posterior ball of the centrum, as if a string had been tied
round that part when it was soft, and there is no appearance of this groove having been
produced by any erosion of the fossil, or being otherwise than natural.

36 FOSSIL REPTILIA OF THE LONDON CLAY.

The same character is repeated, though with less force, in the posterior dorsal
vertebra, fig. 9,T. V, and, together with the general proportions of the specimen,
supports the reference of that vertebra to the Crocodilus champsoides. There is a slignt
longitudinal depression at the middle of the side of the centrum near the suture with
the neurapophysis (7, 7).

Fig. 10 is a side view of the first caudal vertebra of the Crocodilus champsoides .
besides being longer and more slender than that vertebra is in the Croc. toliapicus, the
inferior surface of the centrum is less concave from before backwards.

The evidences of Crocodilian reptiles from the deposits at Sheppy less characteristic
of particular species than those above described, are abundant. Mr. Bowerbank
possesses numerous rolled and fractured vertebre, condyloid extremities, and other
portions of long bones; with fragments of jaws and teeth.

Mr. J. Whickham Flower, F.G.S., has transmitted to me some fragments of the
skull of a Crocodile from Sheppy, including the articular end of the tympanic hone,
equalling in size that of a Crocodilus biporcatus the skull of which measures two feet
eight inches in length.

Mr. Leifchild, C.E., possesses a considerable portion of the lower jaw of a Crocodile
of at least equal dimensions, also from Sheppy, showing the angle of union of the rami
of the lower jaw which corresponds with that in the Crocodilus toliapicus, Pl. 2.

In the museum of my esteemed and lamented friend, the late Frederic Dixon, Esq.,
F.G.S., at Worthing, is preserved a portion of the fossilized skeleton of a Crocodile, from
the Eocene clay at Bognor, in Sussex ; it consists of a chain of eight vertebree, including
the lumbar, sacral, and the biconvex first caudal, which are represented of their
natural size in tab. xv, of Mr. Dixon’s beautiful and valuable work on the ‘ Geology of
Sussex.’ A dorso-lateral bony scute adheres to the same mass of clay close to the
vertebree, and doubtless belonged to the same individual. The proportions of the
vertebre agree with those of the Crocodilus toliapicus. ‘This fine specimen was dis-
covered, and presented to Mr. Dixon, by the Rev. John Austin, M.A., Rector of
Pulbrough, Sussex. Mr. Dixon had also obtained from the same locality a posterior
cervical vertebra of a Crocodile, similar in its general characters to those above
mentioned, but belonging to a larger individual. The length of the body of this
vertebra is two inches and a half.

I have examined some remains of Crocodilia from the London Clay at Hackney; but as
these also are not sufficiently perfect or characteristic for decided specific determination, no
adequate advantage would be obtained by a particular description, or by figures of them.

The chief conclusion arrived at from the study of the Crocodilian fossils from the
Island of Sheppy has been the proof, by the specimens selected for depiction in the
present work, that at least two species of true Crocodile have left their remains in
that locality; that neither of these had a short and broad snout like the Caimans, but
that one of them—the Croc. champsoides—much more nearly resembled the Gavial of

CROCODILIA. 37

the Ganges in the proportion of that part of the skull; although, in its composition,
especially as regards the length and connexions of the nasal bones, it is a true Crocodile.

Amongst the existing species of Crocodile the Croc. acutus of the West Indies
offers the nearest approach to the Croc. toliapicus, and the Croc. Schlegelii of Borneo
most resembles the Croc. champsoides. But there are well-marked characters in both
the skull and the vertebrze which specifically distinguish the two fossil Crocodiles of
Sheppy from their above-cited nearest existing congeners.

Crocopitus Hastinest&, Owen. Tab. VI, VII, VIII, IX, and T. XII, fig. 2 and 5.
Reports of the British Association, 1847, p. 65.

That Crocodiles with proportions of the jaws assigned to the Eocene species noticed
in Dr. Buckland’s ‘ Bridgewater Treatise’ and especially adapted for grappling with
strong mammiferous animals, actually existed at that ancient tertiary epoch, and have left
their remains in this island, is shown by the singularly perfect fossil skull figured in
the above-cited plates. This specimen was discovered by the Marchioness of Hastings,
in the HKocene fresh-water deposits of the Hordle Cliffs in Hampshire, which her
Ladyship has described in the volume of ‘ Reports of the British Association’ above
cited, (p. 63).

When the specimen was originally exposed, it was in the same extremely fragile
and crumbling state as the beautiful carapaces of Zrzonyx obtained by Lady Hastings from
the same locality, and described and figured in the First Part of this Monograph on the
Chelonia ; but thanks to the skill and care with which the noble and accomplished
discoverer readjusted and cemented the numerous detached fragments of those
specimens, the present unique fossil has been in like manner restored as nearly to its
original state as is represented in the plates; and all the requisite characters for deter-
mining the nature and affinities of the species, can now be studied with the same facility
as in the skulls of existing Crocodiles.

If the reader will compare the plates above cited with the section of Cuvier’s
‘Ossemens Fossiles, in which the distinctions between the Alligators and Crocodiles
are specified,* he will see, (in fig. 1, T. VII) for example, that the fourth tooth or
canine of the lower jaw is not received into a circumscribed cavity of the upper jaw,

* «Tes tétes des caimans, outre le nombre des dents, et surtout la maniére dont la quatriéme d’en bas
est recue, outre les différences qui dependent de la circonscription totale, se distinguent de celles des
Crocodiles proprement dits, 1°, parce que le frontal antérieur et le lacrymal descendent beaucoup moins sur
le museau ; 2°, en ce que les trous percés a la face supérieure du crane, entre le frontal postérieur, le pariétal
et le mastoidien, y sont beaucoup plus petits, souvent méme y disparaissent tout-a-fait, comme dans le
caiman a paupiéres osseuses; 3°, en ce que l’on apercoit une partie du vomer dans le palais, entre les
intermaxillaires et les maxillaires; 4°, en ce que les palatins avancent plus dans ce méme palais, et s’y
élargissent en ayant; 5°, en ce que les narines postérieures y sont plus larges que longues, etc.” (tom. v,
pt. ii, p. 105.)

38 FOSSIL REPTILIA OF THE LONDON CLAY

but is applicd to a groove upon the side of the upper jaw, and is exposed. Fig. 1,
T. VI, shows that the prefrontal (14) and lachrymal (73) bones, instead of descending
much less upon the facial part of the skull, extend much more, and advance nearer to
the end of the muzzle than in any Alligator, or even than in any actual species of broad-
nosed Crocodile.

The vacuities left between the postfrontal (12), the parietal (7), and the mastoid (8)
(T. VI, fig. 1, and T. II, fig. 3), are as wide as in the skull of a Crocodilus biporcatus
of equal size, and are larger than in the Alligator lucius or All. sclerops. Fig. 2,T. VII,
shows that no part of the vomer is visible between the premaxillaries (22) and maxillaries
(21), or elsewhere on the palate. But the palatine expansion of the vomer is nota
constant character ; it is wanting, for example, in the Alligator lucius of North America.
The palatines (20) are not more advanced in the fossil in question than they are in the
true Crocodiles, and their anterior portion does not expand to its anterior truncated
termination. The posterior nostril, the entire contour of which is shown in the portion
of the skull of the same species figured in T. VI, fig. 3, is longer than it is broad.

There is but one character in which the fossil skull in question differs from the
true Crocodile, and agrees with most species of Alligator; it is in the reception of
the two anterior teeth of the lower jaw into cavities of the premaxillaries, shown in
fig. 2, T. VII, which are not perforated ; so that there are no foramina anterior to the
bony nostril, as in T. I, in the bone marked 22. These foramina are not, however,
absent in all Alligators; the skull of the Alligator sclerops, figured by Cuvier (tom. cit.,
pl. i, fig. 7), shows them, as do all the species of true Crocodile the skulls of which are
figured in the same plate. There is one character by which the Crocodilus Hastingsie
differs from all known species of both Crocodile and Alligator: it is that afforded by
the broad and short nasal bones (15, fig. !, T. VI), which do not reach the external
nostril; this being formed, as in the Gavials, exclusively by the premaxillaries 22.

In the general proportions, however, of the skull of Croc. Hastingsia, especially
the great breadth, shortness, and flatness of the obtusely-rounded snout, it resembles
that of the Alligators more than that of any known species of true Crocodile, the length
from the tympanic condyle to the end of the snout being to the breadth taken at the
condyles as 16 to 9.

The following are dimensions of the fossil in question :

Feet. Inches. Lines.
Length of skull from the angle of the lower jaw to the

end of “the snout i 1 6 6

Do. from the tympanic condyle to ditto. 1 4 6

Do. do. to the orbit 0 5 4

Do. from the orbit to the external nostril . 0 7 0
Breadth of the skull across the tympanic condyles 0 9 3
Do. the orbits ; 0 7 0

Do. the external nostril 0 4 0

Longest diameter of upper temporal aperture 0 1 9
Do. the post-palatal vacuities 0 4 9

Depth of the lower jaw at the posterior vacuity . 0 3 0
Depth of the occipital region 6 0 4 3

CROCODILIA. 39

The occipital region of the skull (T. VI, fig. 2), in the proportion of its breadth to
the depth of the lateral parts formed by the conjoined paroccipitals (4) and mastoids (s),
resembles that of the true Crocodiles rather than that of the Alligators, in which that
region is proportionally deeper than in the Crocodiles; the vertical extent of the
supraoccipital is less, and that of the conjoined parts of the exoccipitals above the foramen
magnum is greater; the vertical extent of the descending part of the basioccipital is
also greater in proportion to its breadth than in the Alligators. The proportion of the
basisphenoid (5) and of the conjoined parts of the pterygoids (24) which appear in this
view (fig. 2),is less than in the Alligators, but is greater than in most Crocodiles, thus
presenting an intermediatc character; but the entire exclusion of any part of the
posterior nostril from this view is a character of the Alligators, and is due to the
horizontal plane of that aperture in them, and to its position in advance cf the posterior
border of the pterygoids, from which it is partitioned off usually by a bony ridge.
The posterior nostril has the same position and aspect in the Crocodilus Hastingsia,
and these characters of the posterior nostril are perhaps more distinctive between
Alligator and Crocodile than the shape of the aperture, in which the present fossil differs
both from the Alligators and from most of the Crocodiles with which I have compared
it. The backward extension of the exoccipitals and of the basioccipital condyle, is
such as to bring both parts into view in looking directly upon the middle of the upper
surface of the skull, asin T .VI, fig. 1. In this character the fossil resembles the
Crocodiles more than the Alligators, but the projection is greater than in existing
Crocodiles, and equals that in the Sheppy Crocodilus champsoides.

On the upper surface of the skull a distinctive character has been pointed out by
Cuvier in the different proportions of the supra-temporal apertures in the Alligators
and Crocodiles. The horizontal platform in which these apertures are perforated, is
also square in the Alligators ; the mastoidal angles being less produced outwards and
backwards, and the postfrontal angles less rounded off; this difference is shown in the
skulls figured in Cuvier’s pl. i, tom. cit. The Croc. Hastingsig, both by the obtuseness
of the postfrontal angles, and the acuteness and production of the mastoidal angles,
resembles the Crocodiles, as well as by the size of the supra-temporal apertures; these
are ovate with the small end turned forwards and a little outwards.

Another character may be noticed in the figures of the skulls of the three species
of Alligators as compared with those of the three species of Crocodile in Cuvier’s
pl. i, viz. the larger proportional size of the orbits in the former, in which the orbit
much exceeds in size the lateral temporal aperture. In the Al/gator niger, also, I find
the orbits enormous, and it is the encroachment of the narrow anterior part of the orbital
cavity upon the facial part of the prefrontal and lachrymal, that renders that part of those
bones relatively shorter in the Alligators. Inthe Crocodilus Hastingsia the proportions of
the lateral temporal apertures (T. VI, fig. 1, 12, 26) and orbital (11, 14, 73) apertures,
are like those in the species of Crocodile in which the orbits are smallest. The extent of

40 FOSSIL REPTILIA OF THE LONDON CLAY.

the facial part of the prefrontal (14) and lachrymal (73) is greater in the Croc. Hastingsie
than in any existing species of true Crocodile. Another characteristic of the present
fossil presented by the upper surface of the skull, is the shortness as well as breadth of the
nasal bones, and their almost truncate anterior termination at nearly one inch from the
external nostril. In all the Alligators’ skulls that I have examined or seen figured,
the nasal bones are broadest at their posterior third part, and converge to a point
anteriorly, where in the A/igator lucius, e. g., they extend across the nasal aperture.
The interorbital space is slightly concave in the Crocodilus Hastingsie ; two broad
and slightly elevated longitudinal tracts are continued forwards upon the face from the
fore part of the orbits; but they are not developed into ridges, as in the Croc. biporcatus.
The maxillaries swell out a little in advance of the middle of the nasals, and then
contract to the crocodilian constriction at the suture with the premaxillaries, where the
tips of the lower canines appear in the upper view (fig. 1, T. VI), and their whole
crown is exposed in the side view (fig. 1, T. VII). The conjoined parts of the
premaxillaries send a short pointed projection into the back part of the external nostril.
On the under or palatal surface of the skull (T. VII, fig. 2) the maxillo-premaxillary
suture runs almost transversely across, as in the Crocodilus rhombzfer, figured by Cuvier
in pl. iii, fig. 2, of the volume above cited. There is no appearance of the vomer
upon the palate. The palatal bones (20), though somewhat broader anteriorly, and more
abruptly truncate than in any existing Crocodile that I have seen, are more like those
bones in the true Crocodiles than in the Alligators. The portion between the post-
palatal vacuities is longer and narrower; the posterior end of the palatines is nar-
rower, and the part of the bone anterior to the notch receiving the posterior angle of
the palatal plate of the maxillary does not expand in advancing forwards, as it does m
the Alligators: in the Alligator niger this expansion is greater than in the All. Lucius,
and the posterior ends of the palatines are also remarkably expanded, and applied to
the anterior borders of the pterygoids almost as far as their articulation with the
ectopterygoids, the postpalatal vacuities not at all encroaching on the pterygoids, as they
are seen to do at 24, T. VII, fig. 2, and also in the figure of the Crocodilus rhombifer above
cited, and in other true Crocodiles. The form of the pterygoids (24, T. VI, fig. 2) is
peculiar in the Crocodilus [astingsie : they are contracted anteriorly, and send forwards
a short truncated process to meet the narrow posterior ends of the palatines (20); and
the same character being repeated in anotier skull of the same species, from Hordle,
also in the collection of Lady Hastings, iz. which this part of the bony palate (T. VI,
fig. 3) is more perfect than in the subject of T. VII, fig. 2, it may be regarded with
some confidence as specific. In tie Crocodilus champsoides of Sheppy it will be seen,
by fig. 2, T. II, that the pterygoids (24, 24) are not produced where they join the
palatines (20). In the Alligators, the posterior border of the conjoined pterygoids is
deeply notched behind the posterior nostrils, the angles of the notch being slightly
extended backwards: in most Crocodiles, the sides of the notch are so developed that

CROCODILIA. Al

it does not sink deeper than the line of the posterior border of the pterygoids ; and this
modification is exaggerated in the Crocodilus Hastingsi@ (T. VI, fig. 3) in which the notch
in question is merely the interval between two slender diverging processes from the
middle of the back part of the pterygoids. 24. The posterior aperture of the nasal
passages is wholly surrounded in the Crocodilus Hastingsie by the horizontal plate of
the pterygoids, and has the same position and aspect as in the Alligators ; but its form
is heart-shaped, with the apex directed backwards, and the antero-posterior diameter
exceeding the transverse one. I have not met with this form of the posterior nostril
in any other species of Crocodilian; but it is repeated in two individuals of the Croc.
Hastingsig, and may be regarded as a specific character.

The ectopterygoid, 25, T. VI, fig. 3, T. I], fig. 2 (d fig. 2,.pl. ii, ‘Ossemens
Fossiles,’ t. v, pt. ii) articulates with a larger proportion of the outer surface of the
pterygoids (24) in the Crocodiles than in the Alligators: it agrees with the Crocodiles
in the extent of this articulation in the Croc. Hastingsia.

The number of teeth in this species is a = 84.

In the upper jaw the fourth, ninth, and tenth are the largest; and the fifteenth
and sixteenth exceed in size those immediately before and behind them. The alveolar
border of the jaw increases in depth to form the sockets requisite for firmly lodging
these larger teeth, and gives rise to the festooned outline of the jaw, which is found in
all Crocodiles and Alligators in proportion as the teeth are unequal in size.

The lower jaw presents the same compound structure as that in the Crocodilia,
with the general form characteristic of that in the Alligators and in most of the true
Crocodiles: the symphysis, e. g. is as short as Crocodilus biporcatus and the Alligator
niger, in which it extends as far back as the interval between the fourth and fifth
socket. This is the relative position of the back end of the symphysis in a fine and
perfect under jaw of the Crocodilus Hastingsig in the collection of the Marchioness of
Hastings. Ina portion of the under jaw of apparently the same species of Crocodile.
from the same locality, in the collection of Searles Wood, Esq., F. G. S., the symphysis
terminates opposite the interval between the third and fourth tooth.

The chief distinction observable between the modern Crocodiles and Alligators in
the lower jaw is the greater relative size of the vacuity between the angular (30) and
surangular (29) pieces, and the greater relative depth of the ramus at that part, in the
Alligators. In these characters the lower jaw of the present species more resembles
that of the true Crocodiles; although, as the vacuity in question is somewhat larger,
a slight affinity to the Alligator might be inferred from that circumstance. The
comparative figures of the hinder third of the mandibular ramus in T. XII, fig. 4, 5, 6,
will exemplify the difference in question, and the degree of proximity to the crocodilian
and alligatorial characters respectively.

With regard to another character deducible from the relation of the backwardly-
produced angle of the jaw to the articular surface, the Crocodilus Hastingsie@ more

6

A2 FOSSIL REPTILIA OF THE LONDON CLAY.

decidedly resembles the Alligator: I allude to the depth of the excavation between
the articular cavity (29) and the end of the angular bone (30), and to the lower or
higher level of the angle itself: the fossil jaw (fig. 5) resembies the Alligator (fig. 6)
in this respect more than the Crocodile (fig. 4).

The alveoli are twenty in number in each ramus of the Crocodilus Hastingsie : the
third and fourth are large, of equal size, and close together ; behind these the eleventh,
twelfth, and thirteenth are the largest, and the alveolar ridge is raised to support them ;
after the seventeenth the summits of the crowns of the teeth become obtuse, and the
crowns mammilloid, and divided by a constriction or neck from the fang; they each,
however, have a separate socket, as in the Crocodiles, the septa not being complete
at the hinder termination of the dental series, as in the d//iigator niger figured in my
‘Odontography.’*

Fig. 3, T. II, gives a representation, of the natural size, of the cranial platform
of a young Crocodilus Hastingsi@ in the collection of Searles Wood, Esq.; the hemi-
spheric depressions in the surface of the bone are more regular, distinct, and relatively
larger, and the interorbital part of the frontal is narrower, concomitantly with the
larger proportional eyeballs and orbits of the young animal. The relatively larger
supratemporal apertures form another character of nonage; but there is no ground for
deducing a specific distinction from any of the differences observable between this
part of the young crocodile’s cranium and the corresponding part of that of the more
mature specimen (T. VI).

ALLIGATOR HANTONIENSIS, Wood. Tab. VIII, fig. 2.

London Journal of Paleontology and Geology.

On reviewing the characters of the skull of the Crocodilus Hastingsi@ we perceive
that they combine to a certain extent those which have been attributed to the genus
Crocodilus and the genus Alligator ; in general form it resembles most the latter, but
agrees with the former in some of the particulars that have been regarded by Cuvier
and other paleontologists as characteristic of the true Crocodiles. I allude more
particularly to the exposed position of the inferior canes when the mouth is shut.
Respecting which, however, I am disposed to ask, whether this be truly a distinctive
character of importance? One sees that it needs but a slight extension of ossification from
the outer border of the groove to convert it intoa pit; yet the character has never been
found to fail as discriminative of the several species of existing Crocodiles and Alligators
hitherto determined. It constitutes, however, the only difference between the skulls of
the Crocodilus Hastingsie in the collectidn of the Marchioness of Hastings and that fine
portion of skull now, by the kindness of Mr. Searles Wood, before me, on which he
has founded the species named at the head of the present section. So closely, in fact,

* Tom. ii, pl. Ixxv, fig. 3.

CROCODILIA. ~ 43

do those specimens from the same rich locality correspond, that any other comparative
view than that given in T. VIII appeared superfluous. In both the broad nasal bones
terminate at the same distance from the external nostril, which is accordingly formed
exclusively by the premaxillaries ; in both, the palate-bones present the same narrow,
truncate posterior ends, and the same equal breadth of their anterior portions included
between the maxillaries; only these terminate rather more obliquely in Mr. Wood’s
specimen, their anterior ends forming together a very obtuse angle directed forwards.
But this is comparatively an unimportant difference, and I regard as equally insignifi-
cant the slight interruption of the transverse line of the maxillo-premaxillary suture,
at the middle part, which will be seen by comparing fig. 2 with fig. 1, in T. VIII.
The teeth are the same in number, arrangement, and proportion in the Alligator
Hlantoniensis as in the Crocodilus Hastingsig, and the alveolar border of the jaws
describes the same sinuous course.

Had the complete fossil skull first submitted to my inspection at the meeting of
the British Association at Oxford presented the same fosse for the reception of the
lower canines which exist in fig. 2, T. VIII, I should have referred it to the Alligators,
notwithstanding the crocodilian characters of the small orbits, the long facial plates of
the prefrontal and lachrymal, the wide supratemporal apertures, the non-expansion of
the fore part of the palatines, and the non-appearance of the vomer on the palate, with
other minor marks of the like affinity. For all these characters arise out of secondary
modifications, and are presented in different degrees in the different species of
Crocodile, and are rather of a specific than a generic value. They determine the
judgment by the extent of their concurrence rather than by their individual intrinsic
worth, and for that reason, therefore, the exposed position of the lower canine in the
lateral groove of the upper jaw inclined the balance in favour of a reference of the
previously-described fossil to the true Crocodiles. One cannot, indeed, attach any real
generic importance to the modification of the upper jaw in relation to the lower canines.
In three examples, however, in the collection of the Marchioness of Hastings, the
crocodilian modification of this character is repeated, as it is shown in T. VII, fig. 1;
and we have to choose, therefore, between the conclusion that Mr. Wood’s specimen
(T. VIII, fig. 2) presents an accidental variety in this respect, or to view the fossz in
the upper jaw as indicative of not only a different species but a distinct genus from the
Crocodilus Hastingsig. I should be glad to have more evidence on this point, and
especially the opportunity of comparing the posterior nostrils, the orbits, the supra-
temporal apertures, and the occipital part of the skull of a specimen from Hordwell,
repeating the alligatorial character of the fossze in the upper jaw for the lower canines.
I am disposed to regard this character, notwithstanding its constancy in the living
species of Alligator, asa mere variety in the Hordwell fossil; but pending the acquisition
of further evidence, it seems best to record this fossil under the title proposed for it by
the able geologist by whom it was discovered.

44. FOSSIL REPTILIA OF THE LONDON CLAY.

CrocopiLus HASTINGSIA.

Vertebre referable to the CRocopiLus Hastinesi@, Tab. IX.

The fossil crocodilian vertebree obtained from the Eocene sand at Hordle, notwith-
standing the comparatively limited extent of the researches in that interesting
formation, are at least as abundant as those which have been discovered at Sheppy,
but they do not, as at that locality, indicate two distinct species ; all that have, hitherto,
been found belong to one and the same kind of Crocodile, and from their robust
proportions, would seem to have come from a species with a short and broad muzzle,
1ike that of the Crocodile or Alligator, the fossil skulls of which have been described.

Perhaps the most perfect fossil reptilian vertebra that has hitherto been discovered
is the one figured, of the natural size, in T. IX, fig. 1, 2, and 3. It is the fifth
cervical vertebra. As compared with that of the Crocodilus toliapicus (T. V, fig.
1, 2), which it resembles in size, the hypapophysis, /y (fig. 2, T. IX), is much more
compressed, and the under part of the centrum is more extensively and deeply exca-
vated between it and the parapophyses (p); it is also excavated on each side behind
the base of the hypapophysis, from which a progressively widening smooth ridge is
continued to near the posterior surface of the centrum. The interspace at the side of
the vertebra, between the parapophysis and diapophysis, is smaller but deeper in the
Crocodilus Hastingsie. 'Theneurapophyses meet above the centrum in both; but in the
Crocodilus Hastingsi@ they are thicker anteriorly and thinner at their posterior border,
and the neural canal (fig. 2, ~) is more contracted than in the Crocodilus toliapicus.

As compared with the cervical vertebra of the Crocodilus champsotdes from Sheppy,
the present vertebra differs in the form of the hypapophysis in a greater degree than from
the Crocodilus toliapicus. Fig.8,T. V, shows as little as does fig. 2 in the same plate,
the median ridge and lateral excavations of the under part of the centrum which charac-
terise the present vertebra of the Crocodilus Hastingsie. The Crocodilus champsoides
resembles the Crocodilus Hastingsia in the character of the proportion and depression
of that part of the side of the centrum forming the interspace between the par-
apophysis and diapophysis; but the antero-posterior extent of the parapophysis is
relatively less in that Sheppy species. The outer surfaces of the neurapophyses in the
Crocodilus Hastingsie slope or converge towards each other from before backwards, in
a much greater degree than in either of the Sheppy species. I have not observed in
any recent Crocodile or Alligator the median ridge, continued backwards from the
hypapophysis and the lateral depressions, so strongly developed, as in the Crocodilus
Hastingsie. The fore part of the neurapophyses is relatively thicker in this than in the
recent species. The pleurapophyses pi, (figs. 1, 2), are well developed both forwards and
backwards, and the latter productions are expanded and excavated above for the reception
of the fore part of the succeeding cervical rib. The zygapophyses (¢) are thicker at their

CROCODILIA. A5

base, especially the hinder pair, where the base fills up the entire interval between the
articular surface and the base of the spine (see fig. 2). There is the usual deep exca-
vation at the fore and back part of the base of the spine (ws) for the insertion of the
interspinal ligaments. The neural spine is compressed, moderately long, straight and
truncate at its summit.

Although the hypapophysis maintains its characteristic form with much constancy
in the homologous vertebre of the same species of Crocodile, it varies in different
cervical vertebre of the same individual in certain existing species. It is, for example,
shorter and thicker in the third and fourth vertebre than in the succeeding ones in the
Crocodilus acutus; whilst in the Crocodilus biporcatus the hypapophysis of the third
cervical is more compressed than that of the sixth. The greatest difference is, how-
ever, presented, as far as I have yet made the comparison, by the cervical vertebre of
the Alligator lucius, in respect of the hypapophysis, which is broad and short in the
third and fourth cervicals, but becomes Jong and slender in the succeeding cervicals.
The small vertebral centrum (fig. 4, T. IX) resembles, in its broad and stunted
hypapophysis, that of the third cervical vertebra of the Alligator, but with an indication
of a median rising and lateral depressions, behind that process, like those which are
more decisively shown in the fifth cervical vertebra of the larger individual of the Croco-
dilus Hastingsie, to which species I believe the specimen fig. 4 to belong. It is the
homologous vertebra with fig. 8, T. V, and well illustrates the different proportions
of the bones in different species of Crocodile.

Fig. 6 gives a view of the anterior surface of the first sacral vertebra of the
Crocodilus Hastingsiea : the under surface of the centrum has ceased to develope the
median ridge; the short and thick ribs (p/) have completely coalesced with both the
centrum and neuralarch. The anterior concavity has a fuller and more exact elliptical
form than that of the Crocodilus toliapicus (fig. 5,T. V); the anterior zygapophyses
do not project over the rim of that concavity; but, like those of the Alligator and
Crocodile, they are more transversely extended than in the Gavial.

The general proportions of the first caudal vertebra (fig. 7, T. 1X) are intermediate
between those of the Crocodilus toliapicus (fig. 7,T.1V) and of the Crocodilus champsoides
(fic. 10, T. V): the under surface of the centrum is flat, not concave, lengthwise, as
in both the Sheppy Crocodiles; the side of the centrum is irregularly tuberculate, not
smooth, and concave lengthwise; the broad and high neural spine is deeply grooved at its
fore part: a smaller proportion of the hinder end of the centrum (fig. 5) is occupied by
the articular ball than we find in the antecedent vertebre.

As none of the other numerous vertebre and portions of vertebre give any indi-
cations of a different species from the Crocodilus Hastingsig, or add any material
characters to those of that species which have been deduced from the parts of the
skeleton already described, I refrain from trespassing on the reader’s attention or
occupying further space by their description or figures.

46 FOSSIL REPTILIA OF THE LONDON CLAY.

(renus—GAVIALIS, Oppel.

GAVIALIS Dixont, Owen. Tub. X.

The characters of the genus Gavialis are much more strongly marked than are those
which distinguish the Alligators from the Crocodiles, and leave no ambiguity in the
conclusions that may be deduced from them. The present interesting addition to the
catalogue of British Fossil Reptiles, is due to the discovery in the Eocene deposits at
Bracklesham, by my lamented friend the late Frederic Dixon, Esq., F.G.S., of the remains
figured in T. X. The portions of the lower jaw demonstrate, by the slender pro-
portions of the mandibular rami (figs. 1, 5), the extent of the symphysis, the uniform
level of the alveolar series, and the nearly equal distance of the sockets of the com-
paratively small, slender, and equal-sized teeth, the former existence in England,
during the early tertiary periods, of a Crocodilian with the maxillary and dental
characters of the genus Gavialis. These characters are, however, participated in by
some of the extinct Croccdilians of the secondary strata (see T. XI, fig. 2°); but in
them they coexist with a different type of vertebra from that of the recent and known
tertiary Crocodilian genera: it became necessary, therefore, to ascertain what form of
vertebra might be so associated with the fossil Gavial-like jaws and teeth in the
Bracklesham Hocene deposits, as to justify the conclusion that such vertebre had
belonged to the same species as the jaws. Now, the only Crocodilian vertebrz that
have yet been found at Bracklesham, so far as I can ascertain, present the proccelian
type of articular surfaces of the body (T. X), like that in Mr. Dixon’s collection
fig. 8. This vertebra answers to the fifth cervical vertebra in the existing Crocodilians,
and accords in its proportions with that in the Gangetic Gavial. There are a few
indications of specific distinction; the parapophysis (y) or lower transverse process
articulating with the head of the rib, is relatively shorter antero-posteriorly. The
broad, rough, neurapophysial sutures (~) meet upon the middle of the upper part of
the centrum; the elsewhere intervening narrow neural tract sinks deeper into the
centrum than in the modern Gavial, but is perforated, as in that species, by the two
approximated vertical vascular fissures. The hypapophysis (4s) or process from the
inferior surface of the centrum, has been broken off in the fossil, but it accords in its
place and extent of origin with that in the fifth and following cervical vertebre of the
Gavial. Assuming the fossil proccelian vertebree from Bracklesham, and the above-
described vertebra in particular, to have belonged to the same individual or species as
the portions of fossil jaw (figs. 1, 5), then these mandibular and dental fossils must be
referred to the genus Gavialis, or to the long-, slender-, and subcylindrical-snouted
Crocodia with proccelian vertebre.

This genus is now represented by one or two species peculiar to the great rivers
of India, more especially the Ganges; and the fossil differs from both the Gavialis

CROCODILIA. 47

gangeticus, Auct., and from the (perhaps nominal) Gavials tenuwrostris, Cuv., in the
form and relative size of the teeth. The crown (figs. 6, 7) is less slender in the fossil
than in the existing Gavials, and less compressed, its transverse section being nearly
circular. There are two opposite principal ridges, but they are less marked than in
the existing Gavials; and are placed more obliquely to the axis of the jaw, i. e., the
internal ridge is more forward, and the external one more backward, when the tooth
is in its place in the jaw. In the modern Gavial, the opposite ridges, besides being
more trenchant, are nearly in the same transverse line. The other longitudinal ridges
on the enamel of the fossil teeth, are more numerous, more prominent, and better
defined, than in the existing Gavials: the intermediate tracts of enamel present the
same fine wrinkles in the fossil as in the existing Gavials’ teeth.

The two chief portions of jaw (fig. 1, and figs. 4, 5) belong to two individuals of
different ages: indicated by the difference in the breadth and depth of the ramus:
both specimens being from the corresponding part of the jaw, viz. where it forms the
long symphysis characteristic of the Gavials. The specimen (figs. 4, 5) includes a
larger proportion of the jaw than the fragment delineated in fig. 1.

On comparing the latter fragment of the fossil lower jaw with a specimen of a
lower jaw of the Gavialis gangeticus of the same breadth across the symphysial part,
at the intervals of the sockets, which breadth is 3 centimeters (1 inch 3 lines), I find
that the longitudinal extent of 10 centimeters (near 4 inches) of a ramus of the fossil
jaw includes five sockets; but in the recent Gavial the same extent of jaw includes
seven sockets, showing that the teeth are fewer as well as larger in the fossil Gavial, in
proportion to the breadth of the jaws.

The second portion of the jaw (fig. 2) is from the part where the rami diverge
posteriorly from the symphysis, and near the posterior termination of the dentary
series. Here the teeth become shorter in proportion to their thickness, and somewhat
closer placed together: there is a shallow depression (c) in each interspace of the
teeth, for the reception of the crowns of the opposite teeth when the mouth is shut.
These depressions are longer, deeper, and better defined in the fossil than in the recent
Gavial of the same size.

The fragments of jaw and teeth of the fossil Gavial of Bracklesham show examples
of young teeth penetrating the base of the old ones, according to the law of succession
and shedding of the teeth, which characterises the existing Crocodilia: fig. 2 shows the
apex of one of the successional teeth at d; and fig. 3 d the hollow base of the same
incompletely formed tooth seen from below.

Besides the fossil jaws, teeth, and vertebre of the extinct Gavial, a nearly entire
femur (fig. 9) of a Crocodilian has been discovered in the Eocene deposits at
Bracklesham, which in its proportions, agrees with that bone in the Gavial of the
Ganges. Cuvier, in his comparison of the bones of the Gavial with those of the
Alligators and true Crocodiles, merely observes, ‘‘ La forme des os du Gavial ressemble

A8 FOSSIL REPTILIA OF THE LONDON CLAY.

aussi prodigieusement a celle des os du Crocodile, seulement les apophyses épineuses
des vertebres sont plus carrées.”*

With regard to the femur, this bone is more slender in proportion to its length in
the Gangetic Gavial, than in the Crocodilus biporcatus or the Alhgator lucius, and the
anterior convex bend of the shaft commences nearer the head of the bone; and in these
characters the fossil femur from Bracklesham corresponds with the modern Gavial, and
differs from the Crocodiles and Alligators, and also from the Crocodilus Hastingsie, of
which species specimens of the fossil femur have been kindly submitted to me by the
Marchioness of Hastings and Alexander Pytts Falconer, Esq. The fossil femur of the
Gavial from Bracklesham (fig. 9) may therefore be referred, with the utmost probability,
to the same species as the portions of jaw, teeth, and vertebree above described ; and
as these clearly demonstrate a species distinct from any known Gavial, I propose to
call the extinct species of the Eocene deposits at Bracklesham, Gavialis Dizont, after
my esteemed friend, by whose scientific and zealous investigations so much valuable
additional knowledge has been obtained respecting the fossils of that rich, but, previously
to his researches, little known locality.

The tooth represented of the natural size in fig. 10, T. X, was also discovered at
Bracklesham, and forms part of the collection of G. Coombe, Esq. It resembles, in its
proportions and obtuse extremity, the teeth of the Crocodiles rather than those of the
Gavials, and at first sight reminded me of those of the Goniopholis or amphiccelian
Crocodile of the Wealden period. On comparing it closely with similar-sized teeth of
that species, the enamel ridges were more numerous and decided in the Goniophols ;
and the delicate reticular surface in the interspaces of the more widely separated and
feebler longitudinal ridges in the Bracklesham tooth was wanting in the Gonopholis.
The minute superficial characters of the enamel of the large and strong Crocodilian
tooth from Bracklesham, closely agree with those of the Gavialis Dixoni. It is just
possible that this may be a posterior tooth of a very large individual of that Gavial, as
the teeth become at that part of the jaw shorter in proportion to their thickness in the
modern Gavials. If it should not belong to that Gavial, it must be referred toa
Crocodile distinct from those species of the secondary strata, or those existing Crocodiles
which have teeth of a similar form; since they present a different superficial pattern of
markings on the enamel.

On reviewing the information which we have derived from the study of the fossil
remains of the proccelian Crocodilia, that have been discovered in the Eocene deposits
of England, the great degree of climatal and geographical change, which this part of
Europe must have undergone since the period when every known generic form of that
group of reptiles flourished here, must be forcibly impressed upon the mind.

At the present day the conditions of earth, air, water, and warmth, which are

* Ossemens Fossiles, 4to, tom. v, pt. ii, p. 108.

CROCODILIA. A9

indispensable to the existence and propagation of these most gigantic of living Saurians,
concur only in the tropical or warmer temperate latitudes of the globe. Crocodiles,
Gavials, and Alligators now require, in order to put forth in full vigour the powers of
their cold-blooded constitution, the stimulus of a large amount of solar heat, with ample
verge of watery space for the evolutions which they practise in the capture and disposal
of their prey. Marshes with lakes, extensive estuaries, large rivers, such as the Gambia
and Niger that traverse the pestilential tracts of Africa, or those that inundate the
country through which they run, either periodically, as the Nile for example, or with
less regularity, like the Ganges ; or which bear a broader current of tepid water along
boundless forests and savannahs, like those ploughed in ever-varying channels by the
force of the mighty Amazon or Oronooko ;—such form the theatres of the destructive
existence of the carnivorous and predacious Crocodilian reptiles. And what, then,
must have been the extent and configuration of the eocene continent which was drained
by the rivers that deposited the masses of clay and sand, accumulated in some parts of
the London and Hampshire basins to the height of one thousand feet, and forming the
graveyard of countless Crocodiles and Gavials* Whither trended that great stream,
once the haunt of Alligators and the resort of tapir-like quadrupeds, the sandy bed of
which is now exposed on the upheaved face of Hordwell Cliff?

Had any of the human kind existed and traversed the land where now the base of
Britain rises from the ocean, he might have witnessed the Gavial cleaving the waters of its
native river with the velocity of an arrow, and ever and anon rearing its long and slender
snout above the waves, and making the banks re-echo with the loud and sharp snappings
of its formidably-armed jaws. He might have watched the deadly struggle between the
Crocodile and Palzothere, and have been himself warned by the hoarse and deep bellow-
ings of the Alligator from the dangerous vicinity of its retreat. Our fossil evidences
supply us with ample materials for this most strange picture of the animal life of ancient
Britain, and what adds to the singularity and interest of the restored ‘ tableau vivant,’
is the fact that it could not now be presented in any part of the world. The same
forms of Crocodilian Reptile, it is true, still exist, but the habitats of the Gavial and the
Alligator are wide asunder, thousands of miles of land and ocean intervening: one is
peculiar to the tropical rivers of continental Asia, the other is restricted to the warmer
latitudes of North and South America; both forms are excluded from Africa, in the
rivers of which continent true Crocodiles alone are found. Not one representative of
the Crocodilian order naturally exists in any part of Europe; yet every ferm of the
order once flourished in close proximity to each other in a territory which now forms
part of England.

50 FOSSIL REPTILIA OF THE LONDON CLAY.

Order—LACERTILIA.

PLEURODONT LizaRD(?) Tab. XIV, figs. 43, 44.

Although members of the present order, with the modern proccelian type of
vertebree, existed in England during the Wealden and Chalk periods, and the greater
part of the actual class of Reptiles, m all parts of the world, is composed of the same
order, yet but one solitary example of true Lacertian from the formations of the Hocene
tertiary period has hitherto come under my observation—a fact which has often excited
my surprise. Future researches may bring to light farther and better evidence of the
class.

Among the fossils obtained by Mr. Colchester from the Eocene sand, underlying
the Red Crag at Kyson, or Kingston, in Suffolk, the existence of a Lizard, about the
size of the Iguana, is indicated by a part of a lower jaw, armed with close-set, slender,
subcylindrical, antero-posteriorly compressed teeth, attached to shallow alveoli, and
with their bases protected by an external parapet of bone. The fragment of jaw is
traversed by a longitudinal groove on the inside (fig. 44), and is perforated, as in
most modern Lizards, and as in some Fishes, by numerous vascular foramina along
the outside (fig. 43). The teeth are hollow at their base.

PART I1f.

OrvER OPHIDIA.

SERPENTS.

Prior to the publication of my Memoir on the Pa/gophis in the ‘ Geological Trans-
actions, * and my ‘ Report on British Fossil Reptiles, + the sole notice of any fossil
belonging to the order of Serpents was contained in the following passage from the
Appendix to the concluding volume of the second edition of Baron Cuvier’s great and
comprehensive work, the ‘ Recherches sur les Ossemens Fossiles.. After alluding to
the scarcity of the fossil remains of birds, the immortal author of that work proceeds
to say: “The bones of Serpents are still rarer, if it be possible. I have seen no
specimens of them, save the vertebre from the osseous breccia of Cette, of which I
have spoken in the article on those breccia, and a single one from the fresh-water
deposits of the Isle of Sheppy.’t

We may perhaps gather the reason for the silence of Cuvier respecting the
relations of that vertebra and of the fossil vertebree of Serpents in general to each
other, and to those of the existing species, from his brief notice of the Ophidian fossils
from the breccia of Cette; where, after stating in general terms their resemblance in
form and figure to the vertebree of the common harmless snake (Coluber natrix), he
proceeds to remark, ‘‘ but it may well be conceived, that in a genus where the osteo-
logy of the species has so much similitude, it is not in isolated vertebrae that one can
discover specific characters.Ӥ If, however, this discouraging conclusion of the great
comparative anatomist should be countenanced by the results of a rigorous comparison
of the vertebra of the different species of Co/uher, as that genus may be restricted by
modern naturalists, it is by no means borne out by such comparison of the vertebree of
the species of the wider Linnean genus Co/wécr, and gives place to a very different
estimate of the value of vertebral characters, when these are studied in species of the
different Linnean genera of the ‘ Amphibia Serpentes’ in the ‘Systema Nature.’

Baron Cuvier having, conformably with his convictions, deemed it unnecessary to
give figures or to describe the vertebree of Serpents, recent or fossil, in his ‘Ossemens

* Vol. vi, 2d series (1839), p. 209, pl. xxi.

+ Report of the British Association for the Advancement of Science, for 1841.

t ‘Les os de serpens sont encore plus rares, s’il est possible. Je n’en ai vu que des vertebres des
bréches osseuses de Cette, dont j’ai parlé a l'article de ces bréches, et ume scule des terrains d’eau douce de
Vile de Sheppy.” (Tom. vy, pt. ii, p. 526, 1824.)

§ ‘‘ Mais on sent bien que, dans un genre ou J’osteologie des espéces a tant de ressemblance, ce n’est
pas dans les vertébres isolees que l’on peut trouver les caractéres spécifiques.”’ (Op. cit., tom. iv, p. 180.)

52 FOSSIL REPTILIA OF THE LONDON CLAY.

Fossiles,’ I am compelled to premise such observations on the anatomical construction
of this part of the skeleton of those Reptiles as will render intelligible my description
of the fossil ophidian vertebrze, and vindicate the grounds on which some of these are
referred to distinct species, and others to genera of which we have no evidence of the
actual existence in living Nature.

I have selected as the type of an ophidian vertebra that of a large, terrestrial, con-
stricting Serpent (Python Seb@), an African species, which makes the nearest approach
in size to some of the fossil ophidian vertebre from British tertiary strata. The
vertebra figured in T. XIII, figs. 1-4, is from about the middle of the back of a
specimen which was twenty feet in length. In the Pythons, as in other known
Ophidia, all the autogenous elements, except the pleurapophyses (p/, figs. 2’, 3),
coalesce with one another in the vertebre of the trunk; and the pleurapophyses
(T. XIV, fig. 42, pl) also become anchylosed to the diapophyses (ib. d@) in those of the
tail. There is no trace of suture between the neural arch (T. XIII, figs. 1-4”) and
centrum (c). The outer substance of the vertebra is compact, with a smooth or
polished surface. The vertebrze are proceelian, the cup (fig. 2 ¢) being deep, with its
rim sharply defined and most produced at the sides; the cavity lookmg not directly
forwards, but a little downwards, from the greater prominence of the upper over
the lower border: the well-turned prominent ball (fig. 3 c) terminates the back part of
the centrum rather more obliquely, its aspect being backwards and upwards. The
hypapophysis (4) is developed in different degrees from different vertebre, but
throughout the greater part of the trunk presents the form and proportions shown
in figs. 1,44. A vascular canal perforates the under surface of the centrum (fig. 4),
and there are sometimes two or even three smaller foramina. A large, vertically
oblong, but short diapophysis (¢7) extends from the fore part of the side of the centrum
obliquely upwards and backwards. It is covered by the articular surface for the rib,
which is convex lengthwise, and convex vertically at its upper half, but slightly
concave at its lower half. The base of the neural arch swells outward from its
confluence with the centrum, and developes from each angle a transversely elongated
zygapophysis ; that from the anterior angle (z) lookmg upwards, that (2) from the
posterior angle downwards, both surfaces being flat, and almost horizontal. A thick
rounded ridge connects the anterior with the posterior zygapophysis on each side,
extending along the base of the neural arch. The neural canal (fig. 2, ~) is narrow,
with a subtriedral area, and with a narrow longitudinal ridge on each side. The
neural spine (zs) is of moderate height, which scarcely equals its antero-posterior
extent ; it is compressed and truncate. A wedge-shaped process—the ‘ zygosphene’*
(zs, fig. 2)—is developed from the fore part of the base of the spine; the lower apex
of the wedge being, as it were, cut off, and its sloping sides presenting two smooth,
flat, articular surfaces. This wedge is received into a cavity—the ‘zygantrum’t

* Ziyor, a yoke, opnv, a wedge. + Zivyov, and avrpov, a cavity.

OPHIDIA. 53

(fig. 3, z@)—excavated in the posterior expansion of the neural arch, and having two
smooth articular surfaces to which the zygosphenal surfaces are adapted.

Thus the vertebrz of Serpents articulate with each other by eight joints in addition
to those of the cup and ball on the centrum; and interlock by parts reciprocally
receiving and entering one another, like the joints called tenon-and-mortise in carpentry.

This is the most conspicuous, but is not the peculiar characteristic of an ophidian
vertebra; the zygosphene (zs) and zygantrum (za) being developed in certain Lacertians,
e. g. the genus Jguana (T. XIII, figs. 34, 35), but here the articular diapophysis (fig. 33, d)
is much smaller, and forms a simple, convex, sessile tubercle; the hypapophysis is
wanting: the zygosphene (fig. 34, zs) is deeply notched anteriorly, and the zygantra
(fig. 35, za, za) are shallow, and separated frem each other behind: a thick rounded
eminence extends backwards from the diapophysis to the ball on the back part of the
centrum (fig. 36) ; and that ball is a transverse ellipse (fig. 35), not hemispheroid, as
in the Ophidia.

With regard to the specific distinctions which may be deduced from the characters
of the vertebrz of Serpents, it is requisite first to determine the extent to which those
characters vary in the vertebral column of the same species.

The atlas and axis are modified in the same degree as in the Crocodilia, with the
addition of the entire suppression of their pleurapophyses. The atlas (T. XIV, figs. 38, 39)
has two hypapophyses, one behind the other, as we shall find to be the case in other
vertebree of one of the great fossil Serpents. The normal hypapophysis (A, fig. 39),
answering to that marked ca, ex in the woodcut, fig. 8, p. 85, is autogenous and
wedge-shaped, as usual in the Reptilia; and is articulated on each side to a small
portion of the neurapophysis (z) ; it also presents a concave articular surface anteriorly
(fig. 38,4) for the lower part of the basioccipital tubercle, and a similar surface
behind for the detached central part of the body of the atlas (fig. 40, ca), which is here
confluent with that of the axis (fig. 40, cz), forming the so-called odontoid process of
that vertebra; its Ophidian peculiarity being the development of an exogenous
hypapophysis (/’) from its under and back part, like the posterior hypapophysis of the
succeeding vertebre.

The base of each neurapophysis of the atlas (fig. 38, ~) has an antero-internal
articular surface for the exoccipital tubercle, and a postero-internal surface for the
upper and lateral parts of the odontoid (ca), besides the small median inferior facet for
the detached hypapophysis (/): they thus rest on both the separated parts of their
proper centrum. The neurapophyses expand and arch over the neural canal, but
meet without coalescing. There is no neural spine. Each neurapophysis developes
from its upper and hinder border a short zygapophysis (z): and from its side a still
shorter diapophysis (d).

The axis or second vertebra of the trunk with the partially coalesced body of the
atlas or ‘ odontoid,’ is represented at fig. 40, T. XIV. :

The odontoid presents a convex tubercle anteriorly, which fills up the articular

54 FOSSIL REPTILIA OF THE LONDON CLAY.

cavity in the atlas for the occipital tubercle: below this is the surface for the detached
hypapophysial part of the atlas (fig. 39, 4) and above and behind it are the two
surfaces for the atlantal neurapophyses ; the whole posterior surface of the odontoid
is anchylosed to the proper centrum of the axis.

The neural arch of the axis developes a short ribless diapophysis from each side of
its base ; a short zygapophysis (z), from each side of its anterior border; a thick sub-
bifid zygapophysis (z’) from each side of the posterior border, and a moderately long
retroverted spine (ms) from its upper part. The centrum terminates in a ball behind,
and below this sends downwards and backwards a long hypapophysis (Az).

In the skeleton of an African Constrictor (Python regius, Dum.), which measured
15 feet 6 inches in length, there are 348 vertebrae, of which the 279 following the atlas
and axis support simple moveable ribs ; of these vertebrze about 70 anterior ones have
long hypapophyses, as in fig. 4, 4, T. XIV, which in the rest subside to the obtuse ridge
and tubercle, as in fig. 1, 4, T. XIII: the caudal vertebra have not the ribs moveably
articulated ; they are 67 in number; of these vertebre 56 have bifurcate hypapophyses
as in fig. 42, 4, T. XIV; the six anterior caudals have bifurcate ribs (ib., fig. 41, p/), in
the rest they are simple (ib., fig. 42, /), and lengthen out the diapophyses (ib. d,) to
which they are anchylosed.

The ribs or the trunk-vertebree, like those of the tail, are ‘ pleurapophyses’ or ‘ verte-
bral ribs ;' there are no ‘heemapophyses’ or sternal ribs; the exogenous hypapophyses
(1, fig. 42) take the place of the hemapophyses in the tail. The pleurapophyses of
the trunk (T. XIII, figs. 2’, 3’), are long, slender, curved, subcompressed, expanded at
the proximal end, which presents an articular surface chiefly concave, and adapted to
the diapophysial tubercle (d, figs. 2, 3,) above described ; there is a rough depression
on the fore part of the expansion for the insertion of a ligament, and a tuberosity
projects from the upper and back part; the distal end of the rib is truncate, with a
terminal pit; a medullary cavity extends through a great part of the length of the rib,
as shown in fig. 3”.

There is asmall cavity in the substance of each neurapophysis, which communicates
by a smaller foramen with the zygantrum. A vascular cavity in the centrum com-
municates with the neural canal.

In the skeleton of a Tiger-boa (Python tagris), in the Museum of the Royal College
of Surgeons, measuring eleven feet in length, and having 291 vertebre, the 253
following the axis support simple moveable pleurapophyses, articulated to concavo-
convex sessile diapophyses, and constitute the dorsal, abdominal, or trunk-vertebre ;
70 of the anterior of these vertebree have long hypapophyses, as in fig. 4, 4, T. XIV,
they then begin to shorten, and subside to the ridge and tubercle, as shown in fig. 1
and 4, T. XIII, in the rest of the trunk-vertebree. The first caudal vertebra has free
pleurapophyses, but they are short and bifurcate, the upper prong being the shortest ;
in the second caudal the left bifurcate rib is free, but the right is anchylosed to the
diapophyses ; the prongs are of equal length in this and the two following vertebre.

OPHIDIA. 55

In the fifth caudal the outer prong is again shorter, and in the sixth it is a mere
tubercle; at this part of the tail the hypapophyses begin to lengthen, bifurcate, and
progressively increase in length to the sixteenth caudal, and thence gradually diminish
and subside; yet the general configuration of the neural arch, the contour and degree
of production of its posterior border, and the shape of the zygosphene, remain almost
unaltered throughout.

Ina true Boa constrictor, with 305 vertebree, 71 at the anterior part of the trunk have
long hypapophyses ; and of the 60 caudal vertebre, 44 have bifid hypapophyses. The
first caudal is characterised by the sudden shortening of its ribs, and by a short
process from the middle of their outer surface: this process is longer and nearer the
head in the next rib; and in the third caudal vertebra the rib seems to bifurcate from
its proximal end, which has become anchylosed to the diapophysis. Beyond the eighth
caudal the outer costal prong or process disappears, and the anchylosed rib represents
a long deflected diapophysis to within three or four vertebree from the end of the tail.
The last imperforate obtuse bone of the tail is obviously a coalescence of three vertebre.

In the Rattlesnake (Crotalus, T. XIII, figs. 9-12) the hypapophyses (/) continue to
be developed singly, and of equal length with the neural spines (ws), throughout the
trunk; anda any single vertebra might be distinguished from an anterior trunk-
vertebra of a Boa or Python by the following characters: the diapophysis (7) developes
a small, circumscribed, articular tubercle from its upper convexity, anda short process
(d’) from its under part, extending downwards and forwards below the level of the
centrum (c) ; the anterior zygapophysis (2) seems to be supported by a similar process
(d7”) from the upper end of the diapophysis, the point of which projects a little beyond
the end of the zygapophysis (fig. 10); the zygapophyses are less produced outwards
than in the Python; the zygantra (za, fig. 11) are more distinct excavations.

In the Cobra di Capello (Naa, T. XIII, figs. 13-16), the diapophysis presents the
saine well-marked tubercle (7) upon its upper part, but its lower end (d’) is much less
produced than in the Rattlesnake; the process of bone (¢’) underpropping the
zygapophysis projects proportionally further beyond the articular surface (z): the
neural spine (zs) is much lower, and beyond the anterior third of the trunk the
hypapophysis (4) subsides into a ridge, with its point produced backwards beneath
the articular ball of the centrum; the zygantra (za, fig. 15) are distinct cavities.

In the Coluber elaphus (T. XIII, figs. 17-20) the trunk-vertebrz are distinguished by
the great extent to which the part of the diapophysis (d”, fig. 18) which underprops
the zygapophysis (z) is produced beyond the articular surface, the lower end of the
diapophysis (d’) is less produced; the hypapophysis, beyond the anterior fourth part
of the vertebral column, is reduced to a straight ridge, (fig. 20, 2), extending along the
middle of the under surface of the centrum, and not produced posteriorly: a groove
separates the ridge on each side from the diapophysis and the posterior ball of the
centrum. Both the cup and ball and the articular part of the diapophysis are relatively

56 FOSSIL REPTILIA OF THE LONDON CLAY.

smaller than in the Naja; the neural spine (zs, fig. 17) is lower in proportion to its
antero-posterior extent. The pleurapophysis (y/) is shown articulated to the tubercle
in figs. 19 and 20.

The vertebree of the common harmless Snake, Coluber natriz, differ only in size
from those of the larger continental species above described.

In an African Hryzv (T. XIII, figs. 21-24) the diapophysis (d) does not extend beyond
the articular surface of the anterior zygapophysis (z), but is exclusively devoted to
forming a low, subconvex, articular tubercle, which has a longitudinal depression an-
teriorly ; the posterior margin of the neurapophysis (fig. 21, 2) forms an angle above the
zygantrum, which angle, though slight, is more marked than in any of the foregoing
Ophidians; the hinder end of the hypapophysial ridge (/) is slightly produced; the
zygapophyses (2, 2’, fig. 24) are less extended outwards than in the Pythons.

In a Sea-snake (Hydrophis bicolor, T. XIII, figs. 25-28) I find the height of the neural
spine (fig. 25, zs) greater in proportion to its antero-posterior extent than in any of the
foregoing Ophidians. The diapophysis (d) sends a point (@”) outwards a little beyond
the articular surface of the anterior zygapophysis (z); a very small hypapophysis (4)
projects below the articular ball of the centrum, and a low ridge is continued forwards
from it (fig. 28); the posterior border of the neurapophysis (fig. 25, 2) forms no angle,
but is moderately convex, as in all the foregoing Ophidians, excepting the Lryz.

With this indication of the kind and extent of the vertebral characters of the
different species of Serpent which I have been able to study in reference to the fossils
to be described, I proceed to the comparisons by which the following extinct genera
and species have been established.

Genus—PALEOPHIS.

PaLHoPHIs TypHaHus, Owen. Tab. XIII, figs. 5-8, Tab. XIV, figs. 1-3, 7-9, 16, 17,
26, 27, 28. (Fig. 6, 10-12) ?

Amongst the numerous vertebre of this species of Serpent which have come under
my examination, a few, of small size, have shown the hypapophysis long and com-
pressed, as in the specimen in Mr. Bowerbank’s collection, figured in T. XIV, figs. 1-3, 4,
indicating that the vertebre at the anterior part of the trunk had that character, as in
the large existing Serpents; whilst all the larger vertebrae, with the hypapophysis
perfect, manifest shorter proportions of that process, as in the typical example,
apparently from the middle of the abdomen (T. XIII, figs. 5-8, 4) ; whence I infer that
the Paleophis resembled the Python, Boa, Coluber, and Hydrus, in having different
proportions of the hypapophysis at different parts of the vertebral column. Had
every fossil vertebra shown a long hypapophysis like that in T. XIV, fig. 1, we might have
suspected that the species had been of the venomous family, like the Rattlesnake.

The veritable Ophidian nature of the fossils in question is demonstrated, not only

OPHIDIA. 57

by the superadded zygosphenal (2s, fig. 6) and zygantral (za, fig. 7) articulations, but
by the solidity of the zygosphene, by the size and form of the centrum, by those of its
articular cup (c, fig. 6) and ball (c¢, fig. 7), and of its hypapophysis (#); and also by
the size and prominence of the diapophysis (Z). The largest vertebre (e. g. T. XIII,
figs. 5-8, and T. XIV, figs. 16, 26, 27,28) probably from about the middle of the body,
as compared with the vertebre from the same part of the skeleton of a Python Seba,
twenty feet in length, are longer in proportion to their breadth, and the cup and ball
of the centrum are larger; the hypapophysis (4) is more produced, and there is a
second smaller hypapophysis close to the anterior part of the under surface of the
centrum, which in most of the large vertebree is connected by a ridge with the hinder
and normal hypapophysis; but in a few vertebrz is not so connected. ‘The articular
cup and ball are less obliquely placed upon the extremities of the centrum, being
nearly vertical (compare fig. 5 and fig. 1, c’). The rim of the cup is sharply defined,
and is more produced from between the bases of the diapophyses; a deeper and
narrower chink intervening than in the Python. The transverse diameter of the cup
(c, fig. 6) is greater than that of the zygosphene (ib., zs)—a proportion which I have
not found in the vertebrez of any existing genus of Serpent, in which the base of the
zygosphene always equals at least the parallel diameter of the articular cup. The
articular part of the diapophysis is more produced outwards and less extended
vertically in Palgopkis than in Python, and it is uniformly convex; a ridge is
continued from its upper end obliquely forwards to, but not beyond, the apex of the
anterior zygapophysis (z), forming the angle between the lateral and anterior surfaces,
whilst the horizontal articular facet forms the third surface of that three-sided conical
process. In the Python the non-articular part of the same zygapophysis is convex,
and the process is much more extended outwardly ; the proportions of the zygapophysis
in the Pa/gophis more resemble those in the Coluber and Hydrus, but differ from these,
as also from WVaja and Crotalus, in the non-extension of the diapophysial point beyond
the articular surface.

A ridge or horizontal rising of the bone extends from the anterior to the posterior
zygapophysis, but is more or less blunted or subsides midway, and is by no means so
produced outwards as in Python; in this respect more resembling that in Coluber and
Hydrus. Below the middle of this ridge, on a level with the upper surface of the
centrum, there is a short, nearly parallel rising in Pal@ophis (fig. 5). The zygosphene
(fig. 6, zs) is slightly excavated anteriorly, and shows no trace of the tubercle which
characterises the middle of that surface in the Python (fig. 2); it is also broader in
proportion to its height. But perhaps the most characteristic feature of the vertebra
of the Palzophis is the peculiar production of the posterior border of the neurapophysis
into an angle (x, fig. 5) directed upwards, outwards, and backwards, and this is
common to all the species; there is no trace of this process in the Hydrus (fig. 25),
and the nearest approach to it which I have hitherto met with among existing

8

58 FOSSIL REPTILIA OF THE LONDON CLAY.

Serpeuts, is that low, tuberous angle at the corresponding part of the vertebra of the
Erye (fig. 21). The posterior zygapophysis resembles, of course, the anterior one in
its much less extent, especially transversely, as compared with that in the Python, and
the posterior border of the neurapophysis (fig. 5, 2’, z) rises from its apex vertically,
or a little inclined outwards and backwards, giving a squarish form to the surface of
the neural arch in which the zygantra (ea, fig. 7) are excavated ; these cavities, in
proportion to the articular ball beneath, are smaller and less deep than in the Python,
or any other existing genus of Serpent. The sloping sides of the neural arch above
the zygapophysial ridge are more concave than in Python, and so resemble those parts
in Coluber and fHydrus. The latter genus (fig. 25) and Crotalus (fig. 9) most resemble
Paleophis in the proportions of the neural spine (zs) ; this part, however, in Pale@ophis
differs from that of Hydrus in having its base coextensive with the supporting arch,
springing up from the fore part of the zygosphene, whilst this part entirely projects
forwards, clear of the base of the spine in Hydrus, as in Python, Coluber, and Naja;
but in Crotalus the base of the spine has the same antero-posterior extent as in
Paleophis, and it comes very near to the fore part of the zygosphene in Lryx. The
neural spine has been more or less fractured in every specimen of the brittle crumbling
vertebree of the Palgophis Typhaus from the Bracklesham Clay; only one specimen,
which I carefully worked out in relief from a mass of inatrix, after imparting some of
its original tenacity to the substance of the bone, affords a true idea of the peculiar
character of these Ophidian vertebrze, which is afforded by the great height of the
neural spine (see T. XIV, fig. 27, zs); but even here, although the fore part of the spine
equals in vertical extent that of the rest of the vertebra beneath it, I am not sure that its
entire extent is preserved, the part having been obliquely broken away behind this
point before the specimen came into my hands. Some vertebree of another species of
Palgophis from Sheppy, show this elevated spine to be a generic characteristic of the
fossil vertebree.

Inches. Lines.
The ent-e vertical extent of the vertebra, fig. 27, from the end
of the hypapophysis to the summit of the neural spine is 2 5
The length of the same vertebra is : 1 2
The length of a larger vertebra of the same species 1 4
The length of the smallest free vertebra . 0 5

I have specified this last vertebra as being ‘ free,’ because in Mr. Dixon’s collection,
by far the richest in the remains of the great Palgophis of Bracklesham, there are two
smaller vertebrae anchylosed together by both their bodies and neural arches (T. XIV,
figs. 32, 33, 34), which, therefore, are not ‘atlas and axis,’ but from their compressed
form I should judge rather to Have come from the opposite end of the vertebral
column: they have not formed, however, the very extremity of the tail, like the
terminal anchylosed vertebre in the Boa constrictor, or those supporting the rattle

OPHIDIA. 59

in the Crotalus; for the ball of the centrum, the posterior zygapophyses, and the
zygantral articulations, are present on the back part of the second of these anchylosed
vertebre. But before further pursuing the description of this remarkable specimen, I
shall premise a brief notice of the vertebree which have presented other modifications.

In the series of Palzeophidian vertebree from Bracklesham, which I have had the
opportunity of comparing, a few, as has been already remarked, appear to have
come from the fore part of the body by the length of the hypapophysis, as compared
with the size of the vertebree, which is small; a character that adds to the likelihood
of their having come from that extremity of the series. Fig. 1, T. XIV, shows one of
these vertebre in which the hypapophysis (4) is entire; it is shorter and much more
compressed than that process is in the anterior trunk-vertebre of the Python, fig. 4,
and its base extends forwards, as a sharp ridge, to between the diapophyses (fig. 3),
where like them, it has been mutilated by fracture. The zygapophyses are small, and
there is no ridge continued from the anterior to the posterior one; the neurapophysis
presents the characteristic angular production (fig. 1, ~), and the neural spine (ib., xs)
is coextensive with the supporting arch.

The second form of vertebra is characterised by a single and moderately-developed
hypapophysis, the base of which is confined to the hinder half of the under surface of
the centrum, leaving the fore part of that surface concave where it expands between
the bases of the diapophyses. Ihave received twelve such vertebree from Bracklesham,
varying in size between the two extremes given in figs. 5,6, and 10,11, 12,T. XIV. The
hypapophysis (/) which is best preserved in the vertebra fig. 10, is shorter but thicker
than in fig. |; the articular cup and ball are relatively smaller; the zygosphene (zs)
is larger, and its surfaces larger and more vertical (fig. 5); the neural spine has a less
antero-posterior extent. These may be vertebrze from the hinder part of the abdomen,
near the beginning of the tail. Some of them have a minute ridge at the middle of
the anterior inferior concavity (fig. 9).

A third modification of vertebra shows the same limited extent of the base of the
posterior hypapophysis, but a second shorter hypapophysis is constantly developed
from the middle of the space between the bases of the diapophyses. I have examined
twenty of such vertebra ranging in size between the extremes given in figs. 14, 15,
and 17, T. XIV. As compared with fig. 5, the articular cup of fig. 14 is larger, the
zygosphene less, and of a different shape, concave anteriorly and not straight above, but
forming an obtuse angle there. A ridge is continued from the posterior to the.anterior
zygapophysis (fig. 13).

This ridge is more strongly developed in the larger vertebree with the same
modification of the under surface (figs. 20, 21). The articular ball of the diapophysis
would seem not to have descended so low down as in the typical vertebre referred to
Pal. Typheus. 'The neural spine does not extend to the fore part of the zygosphene ;
there is a short but well-defined space above zygosphene in front of the spine.

60 FOSSIL REPTILIA OF THE LONDON CLAY.

Figs. 18, 19, 20, 21, give views of two of the best-preserved vertebree of the present
form, which I have attributed toa distinct species under the name of Paleophis porcatus.

The fourth modification of the Paleophidian vertebre from Bracklesham is the
most common, and is characterised by the coextension of the base of the hypapophysis
with the under surface of the centrum, or by the whole of the middle of that under
surface forming a ridge: both ends of the ridge being produced, the posterior one the
most, and forming the normal hypapophysis. These vertebre are usually of large
size; I have examined upwards of thirty, ranging between the extremes given in
figs. 25 and 26, T. XIV; and itis from this series that I have selected the type vertebra
of the genus Palgophis, T. XIII, figs. 5-8.

The ridge between the anterior and posterior zygapophyses in these vertebre is
absent (T. XIII, fig. 5) or interrupted (T. XIV, figs. 27, 28). There is no well-defined
space above the zygosphene anterior to the base of the neural spine. These vertebre I
regard as typical of the species Paleophis Typheus: they are rather longer in
proportion to their breadth than those of the Palgophis porcatus.

To this category belongs the vertebra with the unusually well-preserved neural spine
(fig. 27), and likewise the two vertebrae which are preserved in their natural connexion,
showing the reciprocal interlocking of their complex articular processes (fig. 28).

The fifth form of the vertebre from Bracklesham is characterised by the com-
pression of the centrum and the convergence of its almost flattened sides to the ridge
on the inferior surface, from which a single hypapophysis is developed. I have examined
not more than four such vertebre, including the two which are anchylosed together,
those (figs. 32-34, T. XIV) being the smallest in size, and the vertebra (figs. 29-31,
T. XIV) the largest. The ridge between the anterior and posterior zygapophyses is
suppressed ; the neural arch gently swells out as it descends from the base of the
neural spine, and from between the zygapophyses it bends in to coalesce with the
converging sides of the centrum. ‘This vertebra has not that character of a caudal
vertebra, which is manifested in the Python and most modern Op/idia by the transverse
pair of hypapophyses ; it shows plainly the base of a single median hypapophysis from
near the posterior surface of the centrum (fig. 34). The diapophyses of fig. 29 are
broken away, together with the anterior concave end of the centrum; had they been
entire, we might have derived from them evidence of the more constant character of
the caudal vertebree of Serpents, which is derived from the coalescence of a short and
straight pleurapophysis with the diapophysis, lengthening out that transverse process,
as in fig. 42. The zygosphene and zygantra are developed, as, indeed, they continue
to be to near the end of the tail in modern Serpents; and the produced angle of the
posterior border of the neurapophysis is as characteristic of the small compressed
vertebree of the Palgophis (fig. 29) as of the larger specimens.

The two anchylosed vertebrze belonging to the compressed series have been already
alluded to. The base of the neural spine is limited to the posterior half of the neural
arch in both (fig. 33). The hindmost of the two vertebree is the longest, measuring

OPHIDIA. 61

five lines, the length of the two being nine lines. In each, the sides of the centrum
are nearly plane, and converge at an acute angle to a ridge, which forms the under
surface ; a very small hypapophysis was continued from the back end of the ridge.
This process is broken away from each vertebra, as are also the diapophyses, which
are indicated by their rough fractured base; they are situated near the lower part of
the side of the centrum, like the long diapophyses of the posterior caudal vertebre of
the Pythons; had they been preserved, their proportions would have determined
whether the anchylosed vertebre were caudal or not.

In the skeleton of a Tiger-boa (Python tigris) in the museum of the Royal College
of Surgeons, anchylosis of the 148th to the 149th vertebra has taken place; and the
166th and 167th vertebrz have been more completely and abnormally fused together,
so as to appear like a single vertebra on the left side, and a double one on the right
side, where there are two diapophyses and two ribs. The compressed form, however,
and diminutive size of the two anchylosed vertebre of Palgophis, strongly indicate
them to be from near the end of the tail, in which case it must be concluded that that
part was compressed, as in the smaller modern Hydrophides, and that the present
extinct Ophidian was a Sea-serpent of at least twenty feet in length.

All the vertebrze with the characters specified in the description of the large
specimens from the trunk, and referable to the Palgophis Typheus, have been obtained
from the Eocene clay at Bracklesham, Sussex: they form part of the collections of
the late Frederic Dixon, Esq., F.G.S., of Worthing; of James 8. Bowerbank, Esq.,
F.R.S.; and of George Augustus Combe, Esq., of Preston, near Arundel, to whom I
have been indebted for some beautiful examples, including the two vertebre in natural
conjunction (T. XIV, fig. 28), and the vertebra with the best preserved neural spine
(ib., fig. 27.)

PALZOPHIS PORCATUS, Owen. Tab. XIV, figs. 13-15, 18, 20, 21.

On comparing together eighteen Paleophidian vertebre of different sizes from
Bracklesham, the smallest of the dimensions represented in figs. 14, 15, and thence
gradually increasing to the size of the specimen fig. 20, I find the following differences :
in fig. 14, e. g. the articular cup and ball at the ends of the centrum are larger in
proportion to the length of the centrum, as compared with the next-sized vertebra,
fig. 5: the under surface of fig. 15 is convex transversely between the diapophyses
and sends down a short median ridge; in fig. 6 it is concave at the same part, and
without the median ridge; but both vertebree have the median process or ‘ hypapo-
physis’ at the back part of the under surface. In fig. 14 the fore part of the zygosphene
is concave, in fig. 5 it is flat; in fig. 5 the upper border is straight, in fig. 14 it forms
an open angle; the space between the zygosphene and zygapophysis is greater in
fig. 5 than in fig. 14.

62 FOSSIL REPTILIA OF THE LONDON CLAY.

Twelve vertebre of progressively increasing size repeat the characters of the
vertebree (fig. 6); i.e. they have the fore part of the under surface between the
diapophyses excavated, and have only one inferior spine, viz. the hypapophysis
developed from the hind part of the under surface; they have also the zygosphenal
articulations nearly vertical, and raised high above those of the zygapophyses (fig. 8).
A vertebra (figs. 22, 23, 24) of the same size as the largest of these twelve differs from
them, and repeats the general characters of the small vertebra (fig. 14): it has the
anterior as well as the posterior hypapophysis ; larger terminal cup-and-ball surfaces in
proportion to its size; smaller intervals between the zygosphenal and zygapophysial
articulations (fig. 24) ; less lofty posterior aliform extensions of the neural arch, and the
base of the neural spine extending nearly to the fore part of that arch. These vertebre,
and especially the larger specimens (figs. 18, 20) have a strong external ridge extending
from the anterior to the posterior zygapophyses on each side of the neural arch. On
comparing one of these vertebrae with another of the ordinary character and of the
same size, the following further differences presented themselves : in the ridged vertebrae,
which are provisionally referred for the convenience of description and comparison to a
distinct species, with the name of Palgophis porcatus, the articular ball is broader in
proportion to its he'ght (compare fig. 23 with fig. 27); the anterior zygapophyses are
more produced outwards and less produced forwards, so that they do not extend
beyond the border of the articular cup, so far as in the non-ridged vertebre of
Paleophis Typheus ; the fore part of the zygosphene in the ridged vertebre is broader,
and less excavated. The breadth of the base of the neurapophysis is greater in the
ridged vertebree than in the unridged ones, in proportion to its length. The articular
surfaces of the zygapophyses are smaller in the ridged than in the unridged vertebre.

Figs. 13, 18, 20, 22, T. XIV, show the ridged character of the sides of the neural
arch in Palgophis porcatus, and fig. 19 shows the consequent superior breadth of the
base of that arch in relation to the length of the vertebra as compared with fig. 8,
T. XIII, a corresponding vertebra of the Paleophis Typheus. Fig. 14 in the same Plate
shows the striking difference in the proportions of the same part of the vertebra in the
Python tigris.

Such are the observed differences which seemed worthy of mention in the series of
Paleeophidian vertebree from the Eocene deposits at Bracklesham which I have had
the opportunity of comparing. The nature of the differences may be interpreted in
different ways: with regard to the small vertebrae, for example, those with a single
spine from the posterior part of the under surface (figs. 1, 2, 3, T. XIII) may be small
cervical vertebree of the same species as that to which the large vertebrze with the two
inferior spines belong; and the small vertebrze with two inferior spines (figs. 14, 15)
may have belonged to a smaller and younger individual of the same species, and have
come from a more posterior part of the vertebral column of such individual. The
anterior vertebre of both Pythons and Boas, for example, are distinguished by an

OPHIDIA. 63

inferior spine, the remaining vertebree to the tail being merely ridged beneath: but I
have not met with such modifications in the trunk-vertebre of the same existing
Serpent, as those that have been pointed out in the vertebre (figs. 5, 6, and figs. 14,
15); and in no specimen of Python or Boa, have I found the vertebree presenting such
differences of character as those indicated in the larger fossil Paleeophidian vertebree
which I have described as ‘ ridged’ and ‘ not ridged.’ Leaving therefore the question
of the nature of the differences in the smaller vertebree (figs. 1 and 14) open, and as
possibly depending upon difference of age and position in the series, I believe the
characters of the ridged vertebra to be those of a distinct species of Palgophis.

Masses of mutilated vertebra and ribs, irregularly cemented tvgether by their
matrix, are occasionally though rarely discovered in the Eocene clay at Bracklesham.
The specimens of such aggregates which I have as yet seen have not exhibited any
verte bre sufficiently complete to yield more than the means of determining the generic
relations. That of which a small portion is figured in T. XVI, fig. 4, is the most
instructive, since it shows the form and structure of the ribs. The proximal half of
the pleurapophysis (p/) equals in size the corresponding part in the Python regius of
twenty feet; it shows the same fine cancellous structure of the articular end, and a
similar medullary cavity, with thin compact walls, forming the body of the vertebra.
The more slender distal portion of another rib is preserved, with the medullary cavity
exposed at its fractured parts.

PALZOPHIS TOLIAPICUS, Owen. Tab. XV and XVI.

Transactions of the Geological Society of London, vol. vi, part ii, p. 209.
Report on British Fossil Reptiles, in the Report of the British Association, 1841, p. 180.

The fossil Ophidian vertebrze which have been discovered in the London clay at
Sheppy are, for the most part, smaller than those from Bracklesham; their common
dimensions equalling those of a Boa constrictor of from ten to twelve feet in length.
They all repeat, however, the generic modifications characteristic of Paleophis; the
hinder margin of the neurapophyses (T. XV, fig. 5) is produced into a pointed or
angular plate; the articular prominence for the rib (ib., fig. 3, 7) is wholly convex; the
zygapophyses are short, and no diapophysial point extends beyond the anterior ones ;_ the
height of the neural spine (T. XV, fig. 1, and T. XVI, fig. 2, zs) exceeds its antero-posterior
extent. The veritable Ophidian character of the Reptile to which these fossil vertebre
belonged, is not only shown by their individual structure, but is well illustrated by the
number of them in natural articulation which have occasionally been found cemented
together in the petrified clay.

One of these Ophidiolites from the clay of Sheppy, in Mr. Bowerbank’s collection,
exhibits a portion of the vertebral column of the Palgophis suddenly bent upon itself,
and indicating the usual lateral flexibility of the spine: in another specimen, including
about thirty vertebre, the vertebree have been partially dislocated and are bent in a
semicircle, Tab. XVI.

64 FOSSIL REPTILIA OF THE LONDON CLAY.

As compared with either of the species of Paleophis from Bracklesham, the
vertebree from Sheppy have the centrum proportionally longer and more slender, with
a smaller terminal cup and ball. In vertebree from Sheppy and Bracklesham in which
those articulations were of equal size, the length of the neural arch at and including
the zygapophyses, was two centimetres in the Pal/gophis tolapicus, and one centimetre,
seven millimetres in the Puleophis Typheus.

The hypapophysial ridge is more constant and better marked; it is produced at
both extremities, and most so at the hinder one, but here in a less degree than in the
Paleophis Typheus, or Pal. porcatus, and the ridge is not interrupted between the two
hypapophyses, as in most of the large vertebree of the Palgophis porcatus. On the
other hand, the rising of the bone continued from the anterior to the posterior
zygapophysis does subside midway more completely than in the Paleophis Typheus; and
the ridge, which in that species extends to the apex of the produced posterior border of
the neurapophysis along the outside of that aliform production, is less developed in
the Paleophis tolapicus: the neural arch is less suddenly compressed above, or
inclines more gradually to the base of the spine; this spine, also, although its base is
extended to near the anterior border of the zygosphene, appears to be higher in
proportion to its antero-posterior extent than in the Palgophis Typheus. 'The diapo-
physis is less produced outwards and downwards than in the Palgophis Typheus or
Paleophis porcatus. In a group of thirty vertebrz of this species cemented together
by the indurated clay from Sheppy, in the Hunterian Collection, and which, in the
original MS. Catalogue of that part of John Hunter’s Collection, were called
‘vertebree of a Crocodile,’ T. XV, fig. 1, several of the long and slender subcylindrical
ribs are also preserved, in the fractured parts of which the medullary cavity is shown.
The articular surface at the proximal end presents the uniform concavity suited to the
convexity of the diapophysis. I have seen no evidence of the process from the upper
and back part of the proximal end of the rib which is present in the Python.

The finest and most strikingly Ophidian example of the great fossil snake of
Sheppy has been obtained from that locality by Mr. Bowerbank since the publication
of the Memoir in which his earlier specimens of the Paleophis toliapicus were deter-
mined and described. It consists of a series of thirty vertebre, from about the
middle of the abdomen, bent into an oval form upon their dorsal aspect, and measuring
twenty inches in length (T. XVI, figs. 1, 2).

As the strong and complex articulations of these vertebre in Serpents opposes
any inflection of the column except from side to side, their unnatural bend in the fossil
is attended with just the amount of mutual dislocation that was requisite to admit of
it; but beyond this amount of dislocation, which chiefly affects the terminal ball and
socket-joints, the vertebre have been preserved in their natural juxtaposition and
succession. The dead body of this eocene serpent has apparently sunk or
been washed into the great stream or estuary, where it has been driven about to

OPHIDIA. 6

eal |

and fro, and variously contorted as it was swept along by the current; the portion
here preserved has been by some external influences obstructed and bent upon itself
in its present unnatural curve, as it finally sank in the sediment in that state of
decomposition when the ligaments were ready to give way to the strain upon them;
but the tough integuments, which have longer resisted dissolution, have served to
retain the partially-dislocated vertebree together until they became fixed in the
matrix in the position in which they are now fossilized. We have in this condition
very good evidence of that long and slender form of body which would admit of such
an extent of inflection from external pressure in a direction contrary to that which the
natural articulations of the vertebrz would allow; but since in Serpents those articu-
lations are so strong, when fresh, as to offer considerable obstacles to any vertical
inflection upwards or downwards, we may infer that the body of the Palgophis, of
which the example in question formed a part, must have floated long enough to have
undergone that degree of internal decomposition, which allowed it easily to yield to
external pressure in any direction.

The characteristically long and comparatively slender spine is well preserved in the
vertebree at ms, fig. 2; and the equally characteristic angular production of the
hinder border of the neural arch is shown in some other of the vertebre. In many
vertebre the ribs are preserved just in that degree of juxtaposition in which they
would remain after yielding to the pressure and movements of the overlying and
accumulating sediment upon the integument of the body. Fig. 3 shows a portion of
the coil, in which a few of the ribs offer to our view the concave articular surface (p/),
which was articulated with the diapophysial tubercle (¢): in fractured portions of the ribs
their medullary cavity is shown. The hypapophysis which terminates the thick and
low inferior ridge of the vertebrz of the present species offers that small degree of
development characteristic of the middle and posterior part of the long abdominal region.

In T. XVI, fig. 1 shows this remarkable chain of vertebrze from the right side; fig. 2
the middle portion of the same chain from the left side; and fig. 3 the under surface
of the vertebrze with the juxtaposed ends of the ribs.

In T. XV, fig. 1 shows a group of the vertebrae of Paleophis toliapicus, in some of
which the long and slender spine (ws) characteristic of the genus is well preserved.
In fig. 3 of the same plate, the position and form of the diapophysial tubercle (¢, @) are
shown. The character of the under surface of the vertebrze is shown in fig. 4, and the
angular aliform production of the neural arch is shown in fig. 5.

One of these characteristic examples of the Palzophis toliapicus is preserved in the
Hunterian Museum’, the others in that of James 8. Bowerbank, Esq., F.R.S. In the
Museum of Mr. Saull, F.G.S., a few vertebrae, and a fragment of the skull of probably
the same species of Palgophis, likewise from Sheppy, are preserved.

* This was figured, by the permission of the President and Council of the College, in illustration of my
original Memoir on the Genus Palgophis in the Geological Transactions.

9

66 FOSSIL REPTILIA OF THE LONDON CLAY.

On a general review of these numerous and rich accessions to our previously
scanty evidence of extinct Serpents, I may sum up by stating that the generic character
of Paleophis is chiefly manifested in the length of the neural spine, in the pointed aliform
productions of the back part of the neurapophyses, in the uniform convexity of the
diapophysial tubercles, and the minor transverse production of the zygapophyses.

The Palgophis tolapicus is distinguished by its longer vertebrae in proportion to
their breadth, by its sessile diapophyses, and by the carinate character of the lower
part of the centrum in the vertebre of the abdomen.

The Palgophis Typhaeus is distinguished by its shorter and broader vertebre, by its
pedunculate diapophysis, and by the anterior and posterior hypapophyses of the
vertebree of the abdomen ; its neural arch is narrower, and its sides not longitudinally
ridged.

The Pale@ophis porcatus is characterised by the longitudinal ridges connecting the
anterior with the posterior zygapophyses, by its broader and squarer neural arch; but
it has the two hemal spines below like the other large species from Bracklesham.

PALZOPHIS (?) LONGUS, Owen. Tab. XIV, figs. 35, 36, 37, 45, 46.

Vertebrze of a serpent agreeing in character with those of the London clay at Sheppy,
but smaller, have been obtained by Mr. Colchester, from the sand of the Eocene
formation underlying the Red Crag at Kyson or Kingston in Suffolk. In these, as in
most of the trunk vertebre of Paleophis Typhaeus, the hypapophysis is a small sub-
compressed tubercle at the under and back part of the body of the vertebra ; but there
is no repetition of a smaller process at the fore part; and no ridge is continued
‘backward from the hypapophysis, as in the Palgophis tohapicus. The tubercle for the
rib is single; in Vaja it is almost divided into two, the upper being convex, the lower
moiety concave; in the Python the upper half of the tubercle is convex, and the lower
half concave, but the two facets are not marked off. In the fossil serpent from
Kingston, as in the Pal@ophis from Sheppy and Bracklesham, the costal tubercle is
simply convex. The chief characteristic of the Ophidian vertebre from Kingston is
the length and slenderness of their bodies, in which respect they exceed those of the
Paleophis toliapicus, and resemble some of the existing tree-snakes (Dendrophis) with
elongated vertebree. The origin of the neural spine is limited to the posterior half of
the arch (fig. 36); but the mutilation of the neurai arch in the specimens I have yet
had the opportunity of examining, prevents a prosecution of the comparison with any
adequate advantage.

OPHIDIA.

Genus—PALERYX.

The vertebre of this extinct genus of Serpent (T. XIII, figs. 29-32, and figs. 37-38)
differ from those of Palgophis, in the absence of the pointed aliform production of the
hinder border of the neurapophyses, that border (fig. 29, z) descending from the neural
spine to the posterior zygapophyses, with a convex curve as in most modern Serpents.
The neural spine (zs) is low, the antero-posterior extent of its truncated summit
exceeding the height. There is no point of bone extending outwards beyond the
articular surface of the anterior zygapophysis (z), as in Coluber, Vipera, Naja, Crotalus,
and Hydrus ; in this character Paleryr resembles Eryx, Python, Boa, and Pal@ophis.
The middle and posterior trunk-vertebre of Paleryz differ from those of Python and
Boa, and resemble those of Hryz in having a sharp and well-developed hypapophysial
ridge (/) coextensive with the under surface of the centrum, and deepest at its posterior
half; but the border here is gently convex, not angular as in Hryz; and the posterior
border of the neurapophysis is less produced than in Hryz; the articular cup and ball
are relatively larger, especially transversely ; the cup is a full transverse ellipse, not
circular as in Hryz ; in this respect it resembles that of Python and Pal@ophis.

PALERYX RHOMBIFER. Tab. XIII, figs. 29-32.

In the vertebre of this species the hypapophysial ridge (4) is sharp and well
produced; the neural spine (zs) is rhomboid, not rounded off anteriorly; the zygo-
sphene (zs, fig. 30) has the same relative vertical extent as in the Python. The
diapophysial tubercle (7) is less elongated vertically than in Python and Boa,
presenting proportions like those of the vertebra of the Eryx (fig. 22, d) ; the zygapo-
physes (zz’) are more pointed at their terminations. The figures 29-32, Pl. 2 represent
the largest of the trunk-vertebree upon which has been founded the genus and species
above defined: they indicate a land Serpent of about four feet in length. They were
obtained from the Eocene sand at Hordwell by Alex. Pytts Falconer, Esq., of Christ-
church, Hants., to whose liberality I am indebted for the specimen figured.

PALERYX DEPRESSUS, Jwen. Tab. XIII, figs. 37 and 38.

The smaller Ophidian vertebree, indicative of the above species, agree in their
generic characters with the foregoing; that is to say, in the shape and development
of the hinder border of the neural arch in the relations of the diapophysis to the anterior
zygapophysis, in the shape and size of the articular cup and ball of the centrum, and
in the shape of the diapophysial tubercle for the rib. But the whole vertebra is more
depressed ; the hypapophysial ridge is relatively thicker and less produced; the
zygosphene has much less vertical thickness, and there is a corresponding modification

68 FOSSIL REPTILIA OF THE LONDON CLAY.

of the zygantra; the neural spine is relatively lower and of a different shape, having
its anterior angle rounded off, and its posterior one more produced backwards.

As I have failed to discover modifications of the kind and degree above described
in the dorsal or free rib-bearing vertebrae of the same species in any of the existing
genera of Serpents, I am left to interpret such characters as indicative of a distinct
species, probably of the extinct genus of Eocene Serpent above defined. The
specimens of the vertebre of the present species, which indicate a serpent of between
two and three feet in length, were obtained from the Eocene sand at Hordwell Cliff,
by Searles Wood, Esq., F.G.S., in whose museym they are preserved.

A few bones of serpents have been found in the superficial stalagmite, and in clefts
of caves, in peat bogs, and the like localities, which bring their occurrence and
deposition within the period of human history. None of these Ophidian remains,
however, have offered any differences in size or other character from the corresponding
parts of the skeleton of our common harmless snake (Coluber natriz). As yet no
Ophidian fossils have been found in British fresh-water formations of the pre-adamitic
or pleistocene period, from which formations the remains of the Mammoth, tichorrhine
Rhinoceros, great Hippopotamus, and other extinct species of existing genera of
Mammaha have been so abundantly obtained. Between the newest and the oldest
deposits of the tertiary period in Geology, there is a great gap in England, the middle
or miocene formations being very incompletely represented by some confused and
dubious parts of the crag of fluviomarine origin in which teeth of a Mastodon have
been found.

The deposits in which the remains of the large serpents of the genus Paleophis
occur so abundantly, carry back the date of their existence to a period much more
remote from that at which human history commences. Yet, as the strange and
gigantic Reptiles that have been restored, and, as it were, called again to life, from
times vastly more ancient, realise in some measure the fabulous dragons of medizval
romance; so the locality on our shore of the English channel in which the Eocene
serpents have been found in most abundance and of largest size, recalls to mind, by a
similar coincidence, the passage cited by an accomplished and popular historian, in
his masterly sketch of the rise and progress of the English nation. “There was one
province of our Island in which, as Procopius had been told, the ground was covered
with serpents, and the air was such that no man could inhale it and live. To this
desolate region the spirits of the departed were ferried over from the land of the
Franks at midnight.” (Macaulay’s History of England, vol. i, p. 5.)

C AND J. ADLARD, PRINTERS, BARTHOLOMEW CLOSE.

i
‘ A
; feyy i ; ; i
a y BECK
i] : i i
¢
i f 4
; ; ar
eae
i
ity ee i Bi ; ; ie

CE ee

A

ox

ws

9)

TAB. XXIX.

CHELONIA.

Side view of the fore part of the skull of the Platemys Bowerbanku, nat. size.
Upper view of the same fossil.

. Side view of a fractured tympanic bone of a large Turtle (Chelone), from Brackle-

sham, showing the long and slender ossicle or ‘ columella’ (16) 2 stu ; nat. size.

Extremity of the same tympanie bone, to which the ‘membrana tympani’ was
attached.

Proximal end of the femur of a very large Chelonian from the Isle of Sheppy.
nat. size.

. Left femur of a Turtle (Chelone mydas) which weighed 150 lbs., nat. size.

T. XXIX.

Day Se dh le he en

Vrxleben

St

TAB. I.

Crocodilus Suchus, half the nat. size.
Fig.
1. Upper view of the skull.

2. Under view of ditto. Both ends of the bony palate have suffered fracture and loss.
apparently in the process of mummifying the body of the Crocodile.

iss
N

oe
cone t tlh ape

Wwe le

ie

ca

R

Day k Sen tthe lie Carers

J. Erxicben

[From an Egyptian Munn)

AGES

Crocodilus

TAB. II.

Kg.

1. Upper view of the skull of the Crocodilus tohapicus, one third the nat. size.

2. Under view of the hinder portion of the skull of the Crocodilus champsoides, on
which Dr. Buckland founded his species of ‘ Crocodile with a short and broad
snout, called Crocodilus Spencert ; half the nat. size.

Upper view of the cranial platform of a young Crocodilus Hastingsia@, nat. size.

ve

mdioen

4

From Nat. on Stone by Fh:

TAB. II 2.

Crocodilus toliapicus, one third the nat. size.

Fig.
1. Oblique side view of the skull.

2. Under view of tus skull.

WaeA

TAB. II.

Crocodilus champsoides, half the nat. size.

. Upper view of the skull.

Under view of the fore part of ditto.

. Side view of ditto.

The left half of the back part of the skull of ditto, nat. size.
The point of an anterior young tooth coming into place, nat. size.
Half of the crown of one of the large conical teeth, nat. size.

One of the shorter and more obtuse posterior teeth, nat. size.

>)

oo

CANADA BR wWhD —

TAB. IV.

Vertebre of the Crocodilus toliapicus, nat. size.

. Fore part of a mutilated seventh cervical vertebra.
. Under pa-t of ditto.
. Side view of a mutilated eighth cervical vertebra.

Front view of the same vertebra.

. Under view of the centrum of a lumbar vertebra.

Under view of the centrum of the fourth or fifth dorsal vertebra.

. Side view of the first caudal vertebra.
. Side view of a middle caudal vertebra.

Under view of the same vertebra.

lve

~

TAB. V.

OF

CONS oR OW

Side view of the fourth cervical vertebra (mutilated) of the Crocodilus tolapicus.
Under view of the same, showing the shape of the Aypapophysis h.

Side view of the sixth cervical vertebra (mutilated) of ditto.

Side view of the first dorsal vertebra (mutilated) of the Crocodilus champsoides.
Front view of the anterior sacral vertebra (mutilated) of the Crocodilus tohapicus.
. Side view of the same vertebra.

Side view of the third cervical vertebra (mutilated) of the Crocodilus champsoides.
. Under view of the same vertebra.

Side view of a posterior dorsal vertebra (mutilated) of the Crocodilus champsoides.
Side view of the centrum of the first caudal vertebra of ditto.

. An anterior tooth of the Crocodi/us champsoides.

. A posterior tooth of the Crocodilus tohapicus.

All the figures are of the natural size.

TAB. VI.
CrocopiLia—Crocodilus Hastingsie, half the nat. size.
Fig.
1. Upper view of the skull.

2. Back view of ditto.

~-.

3. Ectopterygoids (25) and pterygoids (24), with the posterior aperture of the nostril.

TAB. Vil.

CrocopiLia—Crocodilus Hastingsia, half the nat. size.

Fig.
1. Side view of the skull.

2. Under view of the cranial and facial parts of ditto.

Ie WUE

«ggopassnninnoneonstieis HERE a ieeg
ee eh, ~

2. Rg
i i
3 ‘ : oe - rem pts

i
sy.

TAB. VIII.

. Under view of the fore part of the upper jaw of the Crocodilus Hastingsia,

nat. size.

. The same view of the A/hgator Hantoniensis, nat. size.

A GNI,

3 R z ae AR,
cere ee rs,
3 z

Ds

La te
¥
,

ig

hens op

—_

MYA OTE be DME. 4X

!, Crocodilus Hastingsia. 2 Alligator Hantonrensis.

TAB. IX.

Crocodilus Hastingsie, rat. size

. Side view of the fourth cervical vertebra.

]
2. Back view of the same vertebra.

. Side view of a similar vertebra, m2vus the pleurapophyses.

3
4. Under view of the third cervical vertebra.
5. Back view of a lumbar vertebra.

6

. Front view of the first sacral vertebra.

7. Side view of the first caudal vertebra.

I

Day 2 der Un the Min

TAB. X.

Gavialis Dizon, nat. size.

=
ee
oR

1. A fragment of the symphysis of the lower jaw.

2. A fragment of one of the rami from the back part of the symphysis, showing the
depressions cc, in the interspaces of the alveoli.

. The fractured under part of a fragment of the same (d) jaw, exposing the hollow
base of a young tooth.

4. A fragment of a ramus forming the long symphysis of the under jaw of a younger
individual of the same species; upper surface.

. Under side of the same fragment.

. A portion of a tooth of the same species of Gavial.

. A tooth with the germ of its successor, which has entered its base, of the same
Gavial.

ey)

“SIOo Gn

8. The centrum of a cervical vertebra of the same Gavial.
9. The femur of the same Gavial.
0. The crown of a tooth of a large Crocodilian from Bracklesham.

Lav AS

ot

TAB. XI.

CROCODILIA.
Fig.
1. Side view of the skeleton of a Gavial.
1 a. Upper view of the skull of ditto.
2. Side view of the skeleton of a Teleosaur.

2a. Upper view of the skull of ditto.

On the scale of one inch to a foot.

Wap Pht he LIE tea raed

RENEE

Gy hates tal eee

BAIS Sim goatee te Joatbiatectie St i ae
ROK, =< Ss. = x Cee ee

pce ot he
. "hea,

| Vadern Gavial cf the Ganges— 2. Ancient Gavial, er Teleasaur. of Ue Whitby las

aa) ; ;
pe
re ae

TAB. XII.

. Upper surface of the symphysial end of the right ramus of the lower jaw of

Crocodilus biporcatus.

. The same part of Crocodilus lastinysia.

The same part of d/hgator niger.

The hinder part of the right ramus of the lower jaw of Crocodilus hiporcatus.

5. The same part of Crocodilus Hastingsia.

. The same part of Alhgator niger.

Figures |, 2, 4, 5, two-thirds, 3 and 6 one half the nat. size.

TAB. XU.

Oprips.-Reeent and Fossil Ophidian Vertebra ; nat. size.

Kie.
I—4. Vertebra from the middle of the body of a Python Seba, twenty fect in length.
2, 3. A rib of the same Python.
3°. Section of the proximal end of a rib of the same Python, showing the medullary
cavity.
5—8. Vertebree from the middle of the body of the Paleophis typhaus.
9—12. Corresponding vertebree of a rattle-snake. (Crota/us.)
13—16. Corresponding vertebra of a hooded-snake. (qa.)
17—20. Corresponding vertebrae of a harmless snake. (Coluber elaphus.)
21—24. Corresponding vertebree of an African Liye.
25—28. Corresponding vertebree of a sea-snake. (//ydrus bicolor.)
29—32. Corresponding vertebrae of Paleryx rhombifer.

33—36. Corresponding vertebree of an Iguana.
37—38 Corresponding vertebra of Pa/erye depressus.

OE,

TAB. XIV.

Oputp1a.—Fossil and Recent Ophidian Vertebra; nat. size.

Pig.
1—3. A cervical or anterior trunk vertebra of a Palaeophis.
4. A corresponding vertebra of a Python tigris.
5—b. A small vertebra of a Palaeophis with one hypapophysis.
7—9Q. A larger vertcbra of the same species.
10—12. A similar vertebra of the same species.
13—15. A small vertebra of Palgophis porcatus.
\6—17. A large vertebra of Palseophis, longer in proportion to its breadth than
18—20. A type vertebra of Paleophis porcatus.
21. A vertebra of Palcophis porcatus.
22—24. A middle trunk vertebra of the same species of Pal@ophis, as figs. 1—3.
25. A small vertebra of a Pa/@ophis, with an inferior ridge.
26. A large vertebra of do. do.
27. Two views of a vertebra of Palaophis typhaus, with the major part of the long
ncural spme preserved.
28. Two vertebrae of Palcophis typhaus, in natural articulation.
29—31. A vertebra of a Palcophis, of the compressed kind.
32-—34. Two similar but smaller vertebrae of the same kind anchylosed, perhaps from
the tail.
35—37. A trunk vertebra of the Palaophis longus.
38. Front view of the atlas vertcbra of the Python Scbe.
39. Side view of the same vertebra.
40. Side view of the axis vertebra of the same Python.
41. Front view of an anterior caudal vertebra of a Python tigris.
42. Front view of a middle caudal vertebra of the same Python.
43—44. Two views of a portion of the lower jaw of a lizard or sauroid fish.
45—46. The centrum of a vertebra of the Pal@ophis longus.
With the exception of figs. 35—37 and 45—46, which are from Kingston in
Suffolk, all the specimens of Pa/eophis figured in this plate are from Bracklesham,

Sussex.

xIV.

’

sk

on Pb WwW Ww

r= 5

TAB. XV.

Opuipia.—Paleophis toliapicus ; at. size.

. A group of vertebrae, some of which show the long neural spine entire.

. A group of vertebree and ribs.

. Five vertebree in natural articulation with the diapophysis well preserved.
. Under surface of four partially dislocated vertebre.

. Side view of the same vertebre ; natural size.

He XY

lay & om, Ld

bev

gs lis
ree

TAB. XVI.

OPHIDIA.

Fig.
1. Left side view of a chain of thirty trunk vertebre of the Palgophis tolhapicus,
from Sheppy.
2. Right side view of the same vertebre.

3. Under view of five vertebree of the same chain, with the articular ends of some of
of the ribs.

4. Portion of a group of Paleeophis vertebree from Bracklesham, showing the size
and structure of the ribs; natural size

el lth.

JS Dinkel, del

PALMONTOGRAPHICAL SOCIETY.

INSTITUTED MDCCCXLVII.

i

A MONOGRAPH

Cio) ces

moll ist FOssiizi CORALS.

BY
H. MILNE EDWARDS,

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

AND

JULES HAIME.

Lieve 1ezN asd le
INTRODUCTION ; CORALS FROM THE TERTIARY AND CRETACEOUS FORMATIONS.

LONDON:
PRINTED FOR THE PALZONTOGRAPHICAL SOCIETY.

1850.

Reprinted with the permission of the Palaeontographical Society, London

JOHNSON REPRINT CORPORATION JOHNSON REPRINT COMPANY LTD.
111 Fifth Avenue, New York, N. Y. 10003 Berkely Square House, London, W. 1

First reprinting, 1966, Johnson Reprint Corporation
Printed in the United States of America

A MONOGRAPH

OF

THE BRITISH FOSSIL CORALS.

INTRODUCTION.

we

Naturatists often designate under the general name of Cora/, not only the stony
substance of a vivid red which is found on the coast of Barbary, and has been long used
for ornamental purposes, but also a vast number of other marine productions, which have
a calcareous structure, and are considered as appertaining to Zoophytes, more or less
analogous to the Polypi that form the Js¢s zodilis of Linneus, or real Mediterranean Coral.
The remains of the minute plant-like animals which abound in most tropical seas, and con-
stitute in some parts of the globe extensive reefs, or even large clusters of islands, have thus
been very properly called Cora/s. But the same appellation has been erroneously given to
the lapidified teguments of many beings which differ most essentially from all Zoophytes,
and belong some to the great Mollusca tribe, some to the family of Sponges, and others to
the Vegetable kingdom. In all Natural classifications it is necessary to separate that which
is fundamentally different, and to unite that which is in reality similar. Zoologists must,
therefore, be more reserved in the use of this expression, and cannot, without impropriety,
continue to comprehend under the same name all the natural productions which are com-

' Yn writing this Monograph in English, a language with which I am not so familiar as I could wish,
I much fear that the incorrectness of the phraseology will often strike the reader. I preferred, however,
not having recourse to a translator, for the meaning of an author is often misrepresented by those who lend
him their pen, and I thought that in a work of this kind accuracy of description would be preferable
to elegance of style. Before commencing the task I have undertaken, I must also beg leave to express
publicly my grateful feelings for the kind and liberal manner in which Sir H. De la Beche, Mr. Stokes,
Mr. J. 8. Bowerbank, Professor John Phillips, Mr. Frederick Edwards, Mr. Searles Wood, Mr. Dixon,
Mr. Pratt,Mr. Sharpe, Dr. Battersby, Mr. F. W. Fletcher, Mr. J. Gray, and the Council of the Geological
Society of London, have communicated to me the paleontological treasures belonging to their respective
collections. —H. Mitnz Epwarps.

]

il BRITISH FOSSIL CORALS.

monly thus blended together. For us the word Coral, or Corallum, must be synonymous
with Polypidom,* and signify the hard or ossified parts of the body of a Polyp.

In treating of the ‘“ Fossil Corals of Great Britain,’ we must, therefore, exclude from
our investigation the various organic remains which bear a certain resemblance to
Polypidoms, but which do not in reality belong to beings of the same structure, and we
must circumscribe our researches within the boundaries of the group of Zoophytes, which,
in a Natural arrangement of the Animal Kingdom, is represented by the Cuass or Ponyrt.?

These Zoophytes are closely allied to Medusz, and in the actual state of science there is
some uncertainty respecting the natural limits which separate these two groups; but the
mode of organization common to both is so characteristic, that the most superficial
anatomical investigation will always enable the zoologist to distinguish a Polyp or an
Acaleph from the Bryozoa and the Spongide, which, till lately, have been erroneously
considered as belonging to the class of Corals. Polypi have a radiate structure ; a pro-
tractile mouth, surrounded by non-ciliate tentacula; a large and well-organized digestive
cavity ; but have no anus. In Spongide no appearance of tentacula or of a stomach is,
ever met with; and in Bryozoa an intestinal canal, much resembling that of ordinary
Mollusca, is always provided with two distinct openings, a mouth and an anus, the first of
which is encircled by ciliated tentacula. The structure of the digestive organs is, therefore,
characteristic in all these animals, and in most instances the radiate form of the tegumentary
system will alone suffice to render the diagnosis of Polypi an easy task. But when the
Polypidom is reduced to its most simple condition, it sometimes bears great resemblance
to the calcareous or horny covering of certain Bryozoa, or to the reticulate skeleton of some
of the Spongide ; and the Polypidom being the only part of these animals which is found
in the fossil state, it is sometimes hard for the paleontologist to decide whether the
organic remains that assume this form are in reality Corals, or whether they do not belong
to one of the other above-mentioned Zoological divisions.

Polypidoms may present two very distinct forms. Some, belonging to aggregate Polypi,
are developed on the basal surface of these Zoophytes, and constitute a sort. of stem in the

1 In translating the French expression Polypier by the word Polypidom, which has of late been adopted
by some of the most eminent English zoophytologists, we deem it necessry to guard the reader against the
erroneous ideas which the etymology of that name might lead to. Till of late the nature of Corals was in
general misunderstood ; they were supposed to be produced by a yiastic exudation moulded round the body
of the Polyp, and serving as a dwelling for these singular beings, hut not forming a part of their organism.
Such is far from being the case ; the corallum is a part of the animal, in the same way as the coating of the
armadillo or the shell of the lobster belong to the structure of these beings. The words ‘“ Polypidom,”
Polypier, &c., might therefore be objected to, if their meaning was not generally known, and had not
become independent of their etymology.

* The class of Polypi, reduced tg its natural limits, corresponds to the dnthozoa of M. Ehrenberg, and
to the sub-class of Radiated Zoophytes of Mr. Johnston. In the excellent work recently published by
Mr. Dana, the same group is designated by the name of Zoophytes, which is usually employed in a much
wider acceptation, and had long ago been given by Cuvier to the great division of radiate animals, com-
prising Echinoderma and Acalephz, as well as Polypi, ete.

INTRODUCTION. il

centre of the ramified mass produced bv the multiplication of these plant-like animals. The
dendroid red Coral of the Mediterranean Sea and the horny skeleton of Gorgonia are thus
inclosed m the axis of cylindrical branches, formed by the thick coriaceous tegumentary
tissue belonging to the whole community of aggregate Polypi, and studded, as it were, by
the radiate protractile heads of the many individual Zoophytes thus united. Other
Corals, appertaining either to simple or to compound Polypi, are, on the contrary, pro-
duced by the ossification of this tegumentary tissue itself, and mstead of forming a sort of
stem, constitute a sheath, or an assemblage of calcareous tubes, each of which belong to an
individual Zoophyte, correspond to tne lower part of its digestive cavity, and serve as a
kind of cell or lodge into which tke anterior portion of the animal’s body recedes when in
a contracted state.

The basal or stalk-like Corals are in general well characterised by their dendroid form,
compact tissue, and concentric layers. At first sight they may bear a slight resemblance
to certam Bryozoa that have attained a very advanced age ;! but even then the remains of
some non-obliterated cells will always enable an attentive observer to recognise the latter,
and the absence of all trace of any such cavities can easily be ascertained, by grinding down
or fracturing the stem of the above-mentioned Zoophytes. In some few instances these
basal Polypidoms are more like the reticulated skeleton of certain foliaceous Spongide ; but
the concentric lamellz of their stem contrasting with the fibrous structure of the tissue of
the Sponge, will still render them recognisable.

Dermal Corals are in general characterised by features of a more striking aspect,
and it is only when these Polypidoms are reduced to their most simple and degraded form,
that they can be mistaken for the tegumentary skeleton of some of the lowest Bryozoa, or
the reticulate, stony tissue of some highly-organized Spongidee. In all well-developed
Corals of this kind, the central cavity or visceral chamber is more or less completely
divided by a certain number of vertical plates, which project from its walls towards its axis,
and produce that radiate structure which is so remarkable in the Astrean tribe. In most
Bryozoa the mouth, or cephalic aperture of the tegumentary cell, is provided with a horny
operculum,” but no such organ ever exists in a true Coral; and, on the other hand, the
radiate septa which we have just alluded to as bemg conspicuous in most Polypidoms,
never exist in the cells of Bryozoa. The absence of an operculum, or of vertical septa,
will not, however, enable the observer to decide whether the coral-lke organic remains
submitted to his investigation belong to the one or to the other of the two great zoological
divisions, for it is a well-known fact that, in many of the inferior forms among recent
Bryozoa, the tegumentary skeleton is reduced to a simple non-operculated tubular sheath,
and that im certam Polypi (the Tubipora for example), no longitudinal septa are to be
found; and the Polypidom is equally reduced to a calcareous tube, tapering and closed
at its base, open and more or less enlarged at its upper end.

1 The Millepora truncata of Ellis and Solander, for example.
* See “Recherches sur les Eschares,’’ Annales des Sciences Naturelles, 2™ serie, t. vi, pl. i.

iV BRITISH FOSSIL CORALS.

In cases of this kind the distinction between the Polypi and the Bryozoa is always
rendered easy by the most superficial examination of the soft parts of the animal; but it is
sometimes a matter of great difficulty for the palzeontologist, who is necessarily deprived of
all such resources, and can only be guided by the peculiarities observable in the ossified
tissues.

In general, the distinction between Corals and Spongide is also very easy, for the
lamellar structure, so prevalent among the former, is never met with in the latter; but m
some Polypidoms (certain Milleporid for example), the vertical plates disappear, and the
mural tissue becomes extremely porous, irregular, and abundant, so as to resemble much
the reticulated mass formed by the stony skeleton of some Spongidz, where the oscula
aud aquiferous canals are on the contrary more regular than usual. In cases of this kind
it may be necessary to seek for distinctive characters in the internal structure of the
Zoophyte ; and, independently of the benefit to be obtained by the microscopical investi-
gation of the tissue itself, it will sometimes be found useful to examine the form of the
tubular cavities which pervade the mass, and correspond either to the visceral chambers of
the Polypi, or to the great aquiferous ducts of the Spongide; for in the first instance
they are always simple, whereas in the latter they are more or less ramified.

§ IL

The external forms of Corals vary considerably, but are in general more dependent on
the mode of aggregation of the different individuals produced by a common parent than
on the mode of organization peculiar to the animals to which these tegumentary skeletons
belong. Characters derived from these forms can therefore be but of little avail for the
natural arrangement of Polypi; and the classification of these Zoophytes, like that of the
higher animals, must be founded on the principal moditications observable in their struc-
ture. It would lead us too far from the special object of this Monograph, if we were to
enter on the investigation of the anatomical facts which alone can furnish satisfactory
elements for such a classification ; but in order to facilitate the study of the Corals about
to be described, it may be useful for us to revert to a few of the leading points in the
structure of Polypi, and to define some of the expressions which we shall often have to
employ.’

The Scierencuyma, or hardened tissue of Polypi, by which Corals are formed, is
always a portion of the tegumentary system of these Zoophytes, but, as we have already
stated, it may be produced in two very different ways. In some cases it is the result of
a sort of ossification of the chorion or principal tunic of the Polypi; in others it grows on

1 For more ample details on this subject we must refer to our “‘ Memoir on the Structure and Develop-
ment of Corals,” published in the Annales des Sciences Naturelles, 3™° série, t. ix.

INTRODUCTION. v

certain parts of the surface of that membrane in a manner somewhat similar to that in which
calciferous epidermis covers the skin of Crustacea and Mollusca. ‘This epidermic scleren-
chyma constitutes the tissue which Mr. Dana has designated by the name of “ foot-secretion, ”
and is the only anatomical element employed by nature in the formation of the common
red Coral, and the horny tubes of Sertularie ; but in most Polypidoms it is of secondary
importance, and the structure is essentially made up with the dermic sclerenchyma, or
ossified chorion. The calcification of this tegumentary tissue always commences in the
centre of the inferior part of the Polyp, and, spreading gradually, rises as the animal
grows, so as to inclose the lower part of the gastric cavity, and to constitute a sort of cup
or cell, which is sometimes broad and shallow, sometimes long and tubular.

In general the fundamental part of these Corals corresponds to the parietes of the
great gastric or visceral cavity of the Polyp, and forms what may be called the wails of the
Polypidom. The basal disc, the spreading cup, or the columnar sheath so produced, very
seldom remains in this simple condition, and in general soon gives rise to a certain number
of laminate processes, which converge towards the axis of the body, and divide the central
cavity into so many radiating /oculi. These vertical laminz, to which we shall exclusively
apply the name of septa, cover the upper surface of the wall when this spreads out in the
form of a disc (asin Fungiz); but in general they are more or less completely inclosed in
the cup-shaped or tubular cell produced by the growth of this wall around the visceral
cavity, which pervades the body of the Polyp from top to bottom. In some Corals the
septa remain free all along their inner edge ; in other species they adhere to a sort of central
style or plate, which rises from the bottom of the same cavity, and which M. Ehrenberg
has proposed calling the columella. The loculi, or interseptal spaces, are then completely
separated ; and in many Polypidoms, where there is no true columella, the same result is
produced by a greater development of the septa, which become united by means of
regular trabicule branching off from their inner edge, and forming a spurious columella,
the structure of which is usually loose and spongy.

Other lamellar or styliform processes, quite distinct from the septa and the columella,
are in some Corals interposed between these organs, and form around the central style a
sort of circular palisade, somewhat like the staminee which im most flowers surround the
pistil. These additional elements of the Polypidom have been designated by the name of
pali, and form sometimes one, sometimes two or three, circular rows or coronets.

In most Corals other lamellar or spiniform processes extend from the walls outward, and
constitute the parts which we propose calling the cost@ of the Polypidom. In general they
correspond exactly to the septa; and in many cases they seem to be mere prolongations of
these organs through the sort of sheath formed by the walls. Sometimes, indeed, the walls
themselves are no longer composed of a distinct, independent, calcified lamina, and are
made up by a slight thickening and cementing of the septa along the line corresponding to
the boundaries of the gastric cavity and the inner margin of the cost.

The cavity thus circumscribed by the walls of the corallum, and subdivided by the

VI BRITISH FOSSIL CORALS.

septa, the pali, and the columella, is always closed at its bottom and open at its upper
extremity, where it usually presents the appearance of a sort of radiated cup, and constitutes
the calice. In some species, this central cavity, or visceral chamber, remains com-
pletely pervious from one extremity of the corallum to the other; and the membranous
appendices containing the reproductive organs, and situated in the loculi, extend to its
basis, without encountering any obstacle; but in other species a certain number of trans-
verse trabicule or synapticule extend from one septum to another at various heights, and
fill up, more or less completely, the inferior part of the loculi. In other cases, horizontal
or oblique lamimee occupy the same position, and subdivide the loculi into a series of small,
irregular cells ; and sometimes these partitions are developed to such an extent that no
direct communication is preserved between the lower and the upper parts of the visceral
chamber, so that the calice, instead of resembling a deep tubular cup, is reduced to the
form of a shallow basin. In general, these transversal laminz, to which the name of
dissepiments has been given, grow from the sides of the septa in an irregular manner, and
do not unite so as to constitute complete horizontal tabulze, extending from wall to wall;
but in some Corals, where the septal apparatus is even rudimentary, the bottom of the
visceral chamber is incessantly raised by the formation of new floors or ¢abule, which
extend horizontally through the centre of the Polypidom, and constitute, under the calices,
a vertical series of secondary chambers.

Intercostal dissepiments are frequently met with on the outside of the walls of the
corallum and in compound Polypidoms, where the costz are highly developed, a thick
cellular mass is thus formed, and often assumes the appearance of a cenenchyma, or common
tissuc. In other instances, the calcified derm continues to extend exteriorly without con-
stituting distinct costee, and forms a dense or a reticulate tissue, which, in certain aggregate
Corals, is nowhere referable to any individual Polyp, and produces a sort of intermediate
mass or true ccenenchyma.

It is also to be remarked, that the exterior surface of most Corals is covered by a layer
of epithelic sclerenchyma, which is sometimes thick and spongy, but in general thin and
dense, and then constitutes a species of coating, which may be called the epitheca.

‘These different constitutive parts of the Polypidom furnish the principal characters
employed in the classification of Corals ; but the mode of multiplication of the Polypi must
also be attended to in the methodical arrangement of these Zoophytes. In some species,
the young are only produced by the ova, and each corallum is formed by the skeleton of a
single individual; but in most, reproduction also takes place by fissiparity or by gemmation,
and in those cases the young usually remain adherent to the body of their parent, and
thus produce compound Polypidoms. The manner in which the different mdividual
Polypidoms, or corallites thus united, are grouped together, varies very much, and furnishes
also useful zoological characters. It is equally necessary not to neglect studying the
changes which take place in the structure of Polypidoms by the progress of age. Corals,
when young, are in general much less complicated than in the adult state, and the manner

INTRODUCTION. vil

in which the multiplication of their constituent parts is effected is often a subject of great
interest for classifiers as well as for physiologists.

The natural affinities of recent Corals can, in general, be easily recognised by means of
facts obtained from these different sources; but the study of fossil Polypidoms presents
greater difficulties, and the palzontologist must also direct his attention to the modifica-
tions which may have taken place after the death of the Zoophyte, and have been
produced by the slow, but long-continued action of solvent or lapidescent fluids.
Changes of this kind sometimes efface the most important features of these organic
remains, for it often haynens that the different parts of a corallum are not modified with
an equal degree of facility, and the complete destruction of certain organs in specimens,
where other parts are well preserved, may give rise to most delusive »ppearances. Even
generic divisions have thus been established by some paleontologists, on accidental changes
due to fossilization alone, and it is indeed often very difficult to avoid errors of this kmd in
the distinction of species, when the observer is not able to compare a sufficient number
of specimens.

§ Ill.

This Monograph being intended principally for the use of Geologists, we have
thought it advisable not to follow the Zoological classification of Corals in describing
the species belonging to the Fossil Fauna of Great Britam, but to distribute them im
reference to the different Formations in which they are found. We must, however, not
lose sight of the Natural arrangement of these Zoophytes, and before entering on the
specific history of the organic remains which we have to study, it is necessary that we
should make known to the reader the system of classification which we have adopted for
Polypi in general. The following Synopsis will suffice for that purpose, and will serve as
a sort of framework illustrative of the divers Zoological divisions to which we shall often
have to revert as we proceed in the descriptive part of our work.

Vili BRITISH FOSSIL CORALS.

CLASSIFICATION OF POLYPI.

Sus-k1Incpom ZOOPHYTA; Srcrion RADIATA.

CLASS ‘

POLYPI.

Animals of the sub-kingdom of Zooruyra, and of the section of Rapiata,’ organized
for a sedentary mode of life, having no locomotive organs, and being provided with a circle of
retractile tentaculee around the mouth, and acentral gastric cavity, not communicating with
an anus, and contaiming the reproductive organs when these exist ; in general fissiparous,
or multiplying by buds as well as by ovules.

The systems adopted by Cuvier, Lamarck, Lamouroux, and their contemporaries, for the
subdivision of the class of Polypi, were founded on external characters of very little value,
and were quite artificial. In a Memoir, published about twenty years ago,” a first attempt
was made to establish this classification on anatomical facts, and the Zoophytes presenting
the above-mentioned structure were distributed in two groups, characterised by the pre-
scence or the absence of mternal ovaria, and a membranaceous tube leading from the mouth
to the great gastric cavity. Subsequent observations have confirmed these views, and
Mr. Dana, whose recent work’ is one of the most valuable contributions which America has
vet made to Natural History, divides in a similar manner the class of Polypi into two
secondary groups. We shall continue adopting this classification here; but the name of
Actinoidea, which Mr. Dana applies to the first of the two sub-classes thus established,
havig been previously employed by other zoologists m a much narrower acceptation, we
have thought it advisable not to make use of it here, and we propose substituting for it that
of Corallaria. The second group comprises the Sertularian Polypi (Milne Edw.), and may
be designated by the name of //ydraria.

i The sub-kingdom of Zoophytes may be divided into two natural groups: the one comprising all the
true Radiate animals (Echinoderma, Acalephee, and Polypi); the other containing the spheroidal or amor-
phous Zoophytes (such as Spongidee and certain Infusoria). The first may retain the name of Radiata ;
the sccond has been designated by that of Sareodaria (Milne Edwards, Cours élémentaire de Zoologie).

2 Recherches sur les Animaux sans Vertebres, faites aux tiles Chausay, par MM. Audouin et Milne
Edwards (Annales des Sciences Naturelles, premiere serie, t. xv, p. 18, Septembre, 1828).

5 United States Exploring Expedition; Zoophytes. Philadelphia, 1846.

INTRODUCTION. 1x

Sus-cuass lL.

CORALLARIA.

Actinoidea, Dana. Op. cit., p. 16, 1846.

Polypi possessing distinct internal reproductive organs, and having the gastric or
visceral cavity surrounded by vertical, radiating, membranaceous lamellz:

in this division of the class of Polypi, the Corallum is in general calcareous, and may be
either tubular, cyathoid, discoidal, or basal; but never assumes the form of cylindrical,
tubular, horny sprigs, bearing simple bell-shaped cells, for the reception of the contracted
tentacula, as we usually find in the sub-class of Hydraria.

Corallaria present three principal structural modifications, and must therefore be
subdivided into three corresponding groups or orders: Zoantharia, Alcyonaria, and
Podactinaria.

Orper |.
ZOANTHARIA.

Zoanthaires (Zoantha), Bainville. Manuel d’Actinologie, p. 308, 1834.

Zoanthaires (Zoantharia), Milne Edwards. Elém. de Zoologie, p. 1045, 1835; Annot. de Lamarck,
Anim. sans Verteéb., tom. ii, p. 106, 1836.

Zoophyta helianthoidea, Johnston ; in Mag. of Zool. and Bot., vol. i, p. 448, 1837; Hist. of British
Zoophytes, p. 207, 1838.

Zoantharia, J. E. Gray. Synop. Brit. Mus., 1842.

Actinaria, Dana. United States Exploring Expedition, Zoophytes, p. 112, 1846.

Anthozoa helianthoidea, Johnston. Hist. of Brit. Zooph., 2d ed., vol. i, p. 181, 1847.

Polypi with conical, tubular, simple or arborescent, but not bipinnate, tentacula, and
with numerous perigastric membranaceous lamine, containing the reproductive organs.

Zoantharia are in general. coralligenous, and almost all the known fossil Polypidoms
belong to this natural group of Zoophytes.

These Corals are very seldom essentially composed of epidermic tissues, nor do they
scarcely ever constitute basal stems, as is usually the case in Alcyonaria. They are almost
always formed of calcified dermic sclerenchyma, and inclose, more or less completely, the
inferior portion of the great visceral or gastric cavity of the Polyp. Each individual has in
general the form of a deep cup or a tubular sheath, the cavity of which is subdivided into a
circle of loculi, by vertical septa affecting a radiate disposition. No trace of any such septa
is ever met within Corals belonging to other animals of the same class, and although these
parts are sometimes rudimentary in Zoantharia, the starlike appearance of the calice pro-

2

x BRITISH FOSSIL CORALS.

duced by their existence must be considered as one of the most. striking features of this
zoological division. The septa are developed successively, as the Polyp grows, and in general
six of these vertical lamine constitute the primary or fundamental cyclum. Shortly after-
wards a second circle, equally composed of six septa, appears, and the twelve loculi situated
between these secondary septa and the primary ones are next subdivided by a third row or
cyclum of twelve younger septa. The number of the septa often augments still more, and is
sometimes carried very high ; but in general the primary septa continue to be more developed
than ‘the others, and thus divide the whole of the radiate structure into six distinct groups or
systems. In some instances, however, the secondary, or even the tertiary, septa grow so rapidly,
that they soon exactly resemble those of the first cyclum, and in such cases the number of the
systems is apparently much greater.’ Sometimes the number of the primary septa is, on
the contrary, reduced to four, or perhaps even to two, but never reaches eight, as would be
the case if the Polypi of this order had ever eight tentacula and eight perigastric lamelle,
a structure which is always met with in the order of Alcyonaria. It is also to be noted,
that the septa vary considerably im their structure, and thus furnish most important
characters, not only for the distinction of species and genera, but even for the formation of
higher zoological divisions in this order of Polypi.

Zoantharia may be divided into two principal groups, characterised by the structure of
the parietes of their body. One of these sections comprises the species in which the
dermal tissue remains soft and flexible; the other contains those the teguments of which
assume an osseous structure and constitute a calcareous Polypidom.

The ScrerencnyMatous ZoantHaria are the only Zoophytes of this order which we
shall have to mention in the sequel of this work ; it would, therefore, be superfluous for us
to treat of the classification of Malacodermous Zoantharia; but it is necessary that we
should give a detailed account of the methodical arrangement of the first of these groups.
Little is known concerning the anatomical modifications of the soft parts im the different
representatives of this zoological form; but the structure of the Polypidom offers great
variety, and furnishes, to an attentive observer, data which appear sufficient for the natural
classification of Sclerenchymatous Zoantharia. ‘The principal characters which we have
made use of for that purpose, are derived from the dense or porous structure of the
sclerenchyma ; the predominance of the septal apparatus, the mural tissue or the tabular
system in the formation of the corallum; the existence or the absence of dissepiments
uniting the septa and subdividing the loculi, and the mode of development of the Polypi.
Five principal divisions may be thus established in this section, and may be designated by
the following appellations: Zoantharia aporosa, Zoantharia perforata, Zoantharia tabulata,
Zoantharia rugosa, and Zoantharia cauliculata.

1 The laws by which the development of the septal apparatus appears to be regulated, have been laid
down in our memoir on the Structure of Corals, ‘published in the Annales des Sciences Naturelles, 3™°

série, tom. ix, 1848.

INTRODUCTION. x1

Sub-order 1.
ZOANTHARIA APOROSA.

Corallum composed essentially of lamellar dermic sclerenchyma, with the septal ap-
paratus highly developed, completely lamellar, and primitively composed of six elements ;
no tabule.

The foliaceous or lamellar structure of the calcified tissue, which furnishes one of the
principal characters of these Corals, is always recognisable in the exterior part of the septa ;
these organs are never composed of irregular trabiculz, as is the case in Porites, or even
perforated, excepting near their inner margin. ‘The walls are also very seldom porous, and
usually constitute an uninterrupted theca, so as to admit of no communication between the
visceral chamber and the exterior, except by the calice. The septa form the most im-
portant part of the Polypidom ; they augment more or less in number as the Polyp rises,
but in general remain unequally developed, and are disposed in groups corresponding to
the six primitive radii, or to a multiple of that number, but never present a quaternary
arrangement, as is often the case in Cyathophyllide. The visceral chamber remains open
from top to bottom, or is only subdivided by synapticule, or by irregular dissepiments,
which extend from one septum to another without joing together, so as to form a series
of distinct tabulz or discoid floors ; a mode of structure which is on the contrary prevalent,
and very remarkable in most of the Corals belonging to our third and fourth sections.

The Zoantharia aporosa are the most lamelliferous and stelliform of all the Corallaria ;
they are very numerous, and belong to four principal families: the Zurdznolide, the
Oculinide, the Astreide, and the Fungide ; but some few of them cannot find a proper
place in any of these natural divisions, and appear to constitute a certain number of satellite
or transitional minor groups, which partake of some of the characters of two or more of the
above-mentioned principal forms, without possessing any structural peculiarity of sufficient
importance to make us consider them as the representatives of a special type; these groups
are therefore not of the same zoological value as the preceding, and in order to point out
their aberrant nature, we shall designate them by names indicative at once of their principal
affinities and their dependent character: Pseudastreide and Pseudoturbinolide for example.

Family I.
TURBINOLID A.

Milne Edwards and Jules Haime, Recherches sur les Polypiers; Annales des Sciences Naturelles,
3™° série, tom. ix, p. 211, 1848.

Corallum in general simple, never fissiparous, and multiplying by lateral gemmation in
compound species. Interseptal loculi extending from the top to the bottom of the visceral

Xl BRITISH FOSSIL CORALS.

chamber, and contaiming neither dissepiments, as in the Astreide, nor synapticule, as in
the Fungide. Walls thin, lamellar, and imperforated. Septa highly developed, simple,
compact, in general regularly granulated on each side, and never denticulated or lobulated at
their apex. Cost in general well marked and straight. No ccenenchyma in the compound
Polypidoms.

First Tribe—CY ATHININ AS.
Milne Edwards and J. Haime, loc. cit., p. 289, 1848.

Calicule presenting one or more rows of pali, placed between the columella and the septa.

§ 1. A single coronet of pali.
1. Genus CYATHINA.

Caryophyllia, Stokes. Zool. Journ., vol. iii, p. 486, 1828.
Cyathina, Ehrenberg. Corall. des Rothen Meeres, p. 76, 1834; Milne Edwards and J. Haime, op. cit., p. 285.

Corallum simple, never gemmiparous, subturbimate and adherent. Calice circular or
nearly so, with a broad but not very deep central fossula. Columella fasciculate, composed
of a certain number (3 to 20) of vertical, narrow, and twisted lamellar processes, and termi-
nated by a convex, crispate surface. Padi broad, entire, free in aconsiderable part of their
length, and equally developed. Sepéa straight, broad, exsert, and forming six systems,
which are in general unequally developed, and become in appearance much more numerous.

Joste straight, slightly prominent near the calice, more or less obsolete lower down,
delicately granulated, and never armed with tubercles, crests, or spines.

Typical species, Cyathina cyathus, Khrenb., loc. cit. ; Milne Edwards and J. Haime, Ann. des Sc. Nat.,

3™° série, tom. ix, tab. iv, fig. 1.
5) fo}

2. Genus C@NOCYATHUS.
Milne Edwards and J. Haime, Ann. des Sc. Nat., 3™° serie, tom. ix, p. 297, 1848.

Corallum composite and adherent ; the corallites sub-cylindrical, rather tall, segregate
(united near their basis, but free in the greatest part of their length), and not grouped in
rows. Calice circular; fossula not very deep. Columella composed of a few twisted,
lamellar, vertical processes. Padi entire, equidistant from the centre, and similar in size.
Septa rather broad, not projecting much above the walls, and forming four cycla, the
last of which is incomplete in one of the six systems. Coste distinct near the calice
only, straight, flat, broad, and delicately granulated.

These Corals have great affinity to Cyathina, from which they differ principally by

their gemmiparous mode of multiplication, and the permanent union of the young to the
parent.

Typ. sp., Cenocyathus cylindricus, Milne Edw. and J. Haime, loc. cit., tab. ix, fig. 8.

INTRODUCTION.

3. Genus ACANTHOCYATHUS.
Milne Edw. and J. Haime, loc. cit., p. 292, 1848.

Corallum simple, free, subturbinate, slightly compressed, and subpedicellate.
Calice more or less oval. Columella and pali as in Cyathina. Septa broad, exsert, and
forming five cycla; systems unequally developed, so as to form sixteen groups. Coste
partly armed with crests or spines.

Typ. sp., Acanthocyathus Gray, Milne Edw. and J. Haime, loc. cit., tab. ix, fig. 2.

4. Genus BATHYCYATHUS.
Milne Edw. and J. Haime, loc. cit., p. 294, 1848.

Corallum simple, adherent by a broad basis, tall, subturbinate, and slightly com-
pressed. Calice subelliptical, with a broad and very deep fossula. Colwmella small and
crispate. Pal narrow, feeble, entire, and closely united to the septa. Sepéa exsert, thin,
closely set, and forming apparently twelve equally developed systems ; five cycla, the last of
which is more developed than the penultimate one, the septa of which are closely approxi-
mated towards the wall, or even cemented to those of the primary, secondary, and ternary
cycla. Coste very narrow, straight, unarmed, delicately granulated, and distinct down to
the basis of the corallum.

Typ. sp., Bathycyathus chilensis, Milne Edw. and J. Haime, loc. cit., tab. ix, fig. 5.

5. Genus BRACHYCYATHUS.
Milne Edw. and J. Haime, loc. cit., p. 295, 1848.

Corallum simple, extremely short, widening very rapidly, and becoming free in the
adult state. Calice circular, and very slightly excavated. Colwmella very thick, fasciculate,
and terminated by circular papilla. Pali very broad, entire. Septa exsert, narrow, and
forming four cycla; the systems equally developed, and apparently twelve in number.
Coste unarmed.

Typ. sp., Brachycyathus Orbignyanus, Milne Edw. and J. Haime, loc. cit., tab. ix, fig. 6.

6. Genus DiscocyATHUs.

Milne Edw. and J. Haime, loc. cit., p. 296, 1848.

Corallum simple, free, and discoidal. Calice circular and slightly convex. Columella
formed by a single vertical lamina; its apex smooth and undivided. Pai free and
corresponding to the septa of the antepenultimate cyclum. Septa very exsert, broad, and
striated laterally near their apex. Mail horizontal, and covered with an epitheca presenting
some concentric striz.

Typ. sp., Discocyathus Eudesii, Milne Edw. and J. Haime, loc. cit., tab. ix, fig. 7.

XIV BRITISH FOSSIL CORALS.

7. Genus CycLocyatuus.

Corallum simple, discoidal, and having the same characters as the preceding genus,
except that the columella is fasciculate and papillous.
Typ. sp., Cyclocyathus Fittonii, nob.

§ 2. Pali of divers orders, formmg two or more coronets.

8. Genus TRocHocyaTnus.
Milne Edw. and J. Haime, los. cit., p. 300, 1848.

Corallum simple, pediculate or sub-pediculate, but free in the adult state. Calice with
a broad but not very deep fossula. Coldumedla well developed, and composed of prismatic or
twisted processes disposed fascicularly or in a single row. Padi well developed, entire, free
on both edges, and differmg in breadth according to the coronet to which they belong.
Septa very exsert, broad, thick near the wall, striated laterally, and forming from four to
six cycla. Coste often armed.

Typ. sp.. Trochocyathus mitratus, nob. (T. mitratus et T. plicatus, Milne Edw. and J. Haime, loc. cit.,
p- 303); Turtinolia mitrata, Goldfuss, op. cit., pl. xv, fig. 5; Turbinolia plicata, Michelotti, Specim.
Zooph. dil., tab. ii, fig. 9.

9. Genus LEPTOCYATHUS.

Corallum presenting most of the characters of the preceding genus, from which it differs
by its subdiscoid form, and its not showing any trace of adherence.

Typ. sp., Leptocyathus elegans, nob.

10. Genus TuucocyaTHUs.
Milne Edw. and J. Haime, loc. cit., p. 317, 1848.

Corallum simple, very short, and adherent, at least when young. Cadice circular, with
the fossula shallow. Cv/uwmella very large, fasciculate, formed by a great number of prismatic
processes, and terminated by a flat papillous apex. Pal thick, narrow, short, and entire,
those corresponding to the penultimate cyclum of septa the most uweveloped. Septa not
exsert, thick, closely set, and almost equally developed; systems equally developed. Wail
covered by a complete epitheca, slightly striated transversely, and constituting around the
calice a small projecting ring.

Typ. sp., Thecocyathus tintinnabulum, Milne Edw. and J. Haime, loc. cit., p. 317; Cyathophyllum
tintinnabulum, Goldfuss, Petref. Germ., tab. xvi, fig. 6

11. Genus PARACYATHUS.
Milne Edw. and J. Haime, loc. cit., p. 318, 1848.

Corallum simple, subturbinate, and having a broad adherent basis. Calice with a large
but not very deep fossula. Colwmella very broa:|, terminated by a papillous surface, and

INTRODUCTION. XV

formed by processes that appear to arise from the lower part of the inner edge of the
septa. Pali in general lobulated at their apex, narrow, tall, and appearing also to proceed
from the inferior part of the margin of the septa, their size diminishing as they approach
nearer to the columella. Septa nearly equal, very slightly exsert, and closely set, their lateral
surface strongly granulated, and presenting sometimes traces of imperfect dissepiments ;
four or five cycla; systems equally developed. Coste nearly equal, straight, closely set,
projecting very little, and delicately granulated.

Typ. sp., Paracyathus procumbens, Milne Edw. and J. Haime, loc. cit., tab. x, fig. 6.

12. Genus HETEROCYATHUS.
Milne Edw. and J. Haime, loc. cit., p. 323, 1848.

Corallum simple, sub-cylindrical, extremely short, and adherent by a basis at least as
broad as the calice, but appearing free, because in the adult state it imbeds in its tissue the
small shell to which it is fixed. Ca/ice circular, or nearly so, with a broad, deep fossula ;
Columella small, and composed of very slender vertical styli. Pal broad, thin, and denti-.
culate. Septa very exsert, broad, thick, and covered with conicai granulations arranged in
radiate series ; four or five cycla, the last of which is more developed than the penultimate
one, and composed of septa that diverge from the older septa as they advance towards the
centre of the visceral chamber. Coste straight, thick, closely set, and strongly granulated.

Typ. sp., Heterocyathus equicostatus, Milne Edw. and J. Haime, loc. cit., tab. x, fig. 8.

13. Genus DELTOCYATHUS.
Milne Edw. and J. Haime, loc. cit., p. 325, 1848.

Corallum short, conical, free, and presenting no trace indicating its having been
adherent when young. Calice circular, and almost flat. Colwmell/a multipartite. Pali highly
developed, and very unequal, those of the penultimate circle the largest, and turned towards
those of the antepenultimate row, so as to form with them a series of delta. Septa slightly
exsert. Coste straight, unequal, distinct down to the basis of the corallum, and strongly
granulated, so as to assume a moniliform appearance.

Typ. sp., Deltocyathus italicus, Milne Edw. and J. Haime, op. cit., tab. x, fig. 11; Stephanophyllia
ttalica, Michelin, Icon. Zooph., tab. vui, fig. 3.

14. Genus TROPIDOCYATHUS.
Milne Edw. and J. Haime, loc. cit., p. 326, 1848.

Corallum simple, free, presenting no trace of former adherence, compressed, and having
at its basis a large, thick, transverse, vertical crest, or two projecting lobes, resembling
wings, or the fins of sepia. Cadice elliptic and arched, its small axis being much higher
than its long axis; fossula not very deep. Co/wmella oblong and multipartite. Pali entire ;
those corresponding to the penultimate cyclum of septa taller and broader than the others.
Septa exsert ; the six systems equally developed. Coste well marked, especially at the
upper part of the wall, and covered with small granulations.

Typ. sp., Tropidocyathus Lessonii, Milne Edw. and J. Haime, loc. cit.; Flabellum Lessonii, Michelin,
in Guerin’s Mag. de Zool., 1843, tab. vi.

Xv1 BRITISH FOSSIL CORALS.

15. Genus PLACOCYATHUS.
Milne Edw. and J. Haime, loc. cit., p. 327, 1848.

Corallum simple, pedicellate, and slightly compressed. Colwmella lamellar, with its
apical margin straight. Padi thin, resembling lobes of the septa, and corresponding only to
the septa of the penultimate and antipenultimate cycla; those facing the latter more
developed than the others (a disposition which forms an exception to the common rule).
Septa numerous, thin, broad, and slightly exsert. Coste distinct from the top to the
bottom of the walls, but projecting very little, nearly equal, and appearing to bifurcate
towards the upper part of the corallum.

Typ. sp., Placocyathus apertus, Milne Edw. and J. Haime, loc. cit., tab. x, fig. 10.

Second Tribe—TURBINOLIN 2.
Milne Edw. and J. Haime, loc. cit., p. 235, 1848.

Corallum destitute of pali; the septa extending to the columella, or meeting in the
centre of the visceral chamber.

§ 1. Wall naked, or having only an incomplete epitheca.
16. Genus TURBINOLIA.

Turbinolia (in parte), Lamarck, An. sans Vert., vol. ii, p. 359, 1816 ; Turbinolia (in parte), Ehrenberg,
op. cit., p. 53, 1834; Dana, op. cit., p. 374; Turbinolia, Milne Edw. and J. Haime, loc. cit., p. 235, 1848.

Corallum simple, conical, straight, and presenting no trace of adherence. Calice circular.
Columella styliform. Septa exsert, those of the last cyclum bent toward the neighbouring
ones and united to them. Cos¢@ lamellar, straight, entire, and very projecting ; the inter-
costal grooves presenting a double series of small dimples, resembling pores.

Typ. sp., Turbinolia sulcata, Lamarck, Hist. Anim. sans Vert., vol. ii, p. 231; Cuvier and Brongniart,
Géographie Minéral. des Envir. de Paris, tab. i, fig. 3.

17. Genus SPHENOTROCHUS.
Milne Edw. and J. Haime, loc. cit., p. 240, 1848.

Corallum simple, presenting no trace of adherence, straight, and cuneiform. Catice
elliptical. Colwmedia lamellar, and occupying the great axis of the calice; its upper margin
flexuous and bilobate. Seta broad, slightly exsert, and forming three cycla; apparently
twelve systems in the adult. Coste broad, not very prominent, in general crispate, or
represented by series of papillous tubercles.

Typ.sp., Sphenotrochus crispus, Milne Edw. and J. Haime, loc. cit., p. 241 ; Turbinolia crispa, Lamarck,
op. cit., vol. ii, p. 231; Milne Edwards, Atlas du Régne Animal de Cuvier, Zooph., pl. Ixxxil, fig. 4.

INTRODUCTION. XV

18. Genus PLATYTROCHUS.

Milne Edw. and J. Haime, loc. cit., p. 246, 1848.

Corallum simple, straight, cuneiform, and presenting no trace of adherence. Calice
elliptical. Columella fasciculate, and terminated by papilla. Septa exsert, very broad,
nearly equal, and very strongly granulated ; three cycla; systems equally developed. Coste
of two sorts, those that occupy the middle of each side of the corallum enlarging as they
ascend ; the lateral ones larger and much broader at their bases than near the calice, so as
to render the lateral edges of the corallum almost parallel.

Typ. sp., Platytrochus Stokesii, Milne Edw. and J. Haime, loc. cit. tab. vii, fig. 7; Turbinolia Stokesii,
Lea, Contrib. to Geol., tab. vi, fig. 207.

19. Genus CERATOTROCHUS.

Milne Edw. and J. Haime, loc. cit., p. 248, 1848.

Corallum simple, subpedicellate, free in the adult state, and recurved towards its basis.
Calice circular, or nearly so. Coluwmella fasciculate, and highly developed. Septa straight,
broad, and exsert. Costz partly armed with spines, crests, or small lobular processes.

Typ. sp., Ceratotrochus multiserialis, Milne Edw. and J. Haime, loc. cit., tab. vii, fig. 5; Turbinolia
multiserialis, Michelotti, Spec. Zool. tab. ii, fig. 7.

20. Genus DiscotTrocuts.
Milne Edw. and J. Haime, loc. cit., p. 251, 1848.

Corallum simple, discoidal, and presenting no trace of adherence. Ca/ice circular, and
almost flat. Columella fasciculate, and terminated by papillz equal in size. Septa straight,
very broad, and projecting but little laterally. MWal/ horizontal. Coste straight and simple.

Typ. sp., Discotrochus Orbignyanus, Milne Edw. and J. Haime, loc. cit., tab. vii, fig. 6.

21. Genus DESMOPHYLLUM.
Ehrenberg, op. cit., p. 75, 1834.

Corallum simple, and adherent by a broad basis. Calice with a very deep fossula. No
columella (a character which distinguishes this group from all the preceding Turbinoline).
Septa broad, very exsert, free almost all along their inner edge, and grouped in fascicule ;
those of the last cyclum taller than those of the penultimate cyclum, and cemented
exteriorly to the older septa. Coste distinct near the calice, but obsolete on the lower part
of the wall, where there are only a few granulations.

Typ. sp., Desmophyllum crista-galli, Milne Edw. and J. Haime, loc. cit., tab. vii, fig. 10.

3

xvill BRITISH FOSSIL CORALS.

§ 2. Wall completely covered by a pellicular epitheca.
22. Genus FLABELLUM.
Lesson, Illustr. de Zoologie, 1831; Phyllodes, Philippi, Neues Jahrbuch fiir Miner. Geol. 1841.

Corallum simple, compressed, and in general free in the adult state. Calice usually
elliptic, very strongly arched in the direction of its long axis; fossula narrow, and very
deep. Columella spurious, and formed by marginal trabicule of the septa; very little
developed, or even quite rudimentary. Septa in general very numerous, appertaining in
reality to six primitive systems, but forming in appearance a much greater number of
systems ; not projecting above the margin of the wall, and presenting laterally regular rows
of well-developed granulations. Walls completely covered with a thin, slightly-striated
epitheca, and in general armed laterally with long spiniform processes, corresponding with
the direction of the long axis of the calice. No radiciform appendices.

Typ. sp., Flabellum pavoninum, Lesson, op. cit., pl. xiv.

23. Genus PLACOTROCHUS.
Milne Edw. and J. Haime, loc. cit., p. 282, 1848.

Corallum resembling much those of the preceding genus, but having a lamellar
columella.

Typ. sp., Placotrochus levis, Milne Edw. and J. Haime, loc. cit., tab. viii, fig. 15.

24. Genus BLASTOTROCHUS.

Milne Edw. and J. Haime, p. 282, 1848.

Corallum resembling those of the genus Flabellum, but gemmiparous ; the young pro-
duced by buds placed along the lateral edges of the corallum, and becoming free by the
progress of their development.

Typ. sp., Blastotrochus nutriz, Milne Edw. and J. Haime, loc. cit., tab. viii, fig. 14.

25. Genus RHIZOTROCHUS.
Milne Edw. and J. Haime, loc. cit., p. 281, 1848.

Corallum simple, trochoid, and adherent by means of cylindrical radiciform appendices,
which proceed from the wall, at different heights, and descend to embrace the extraneous
body on which the Zoophyte lives. Calzce almost oval, with a very narrow and very deep
fossula. No columella. Septa extending to the middle of the visceral chamber, where they
unite without presenting any trabiculz.

Typ. sp., Rhizotrochus typus, Milne Edw. ana J. Haime, loc. cit., tab. 8, fig. 16.

INTRODUCTION. XIX

Aberrant Group.
PSEUDOTURBINOLID 4.

Corallum simple, with the loculi open and devoid of synapticulz or dissepiments, as in
Turbinolide, but having the septa represented by groups of three vertical lamine, not
adhering together, excepting near their external margin, where they are united by a common
costa; a mode of structure, which is quite anormal in the whole order of Zoantharia.

Genus DASMIA.
Milne Edw. and J. Haime, op. cit., p. 328, 1848.
Corallum subturbinate, and appearing not to be free. Septa strongly granulated. Coste
thick, equal, not numerous, and separated by deep grooves.

Typ. sp., Dasmia Sowerbyi, Milne Edw. and J. Haime, loc. cit., tab. vu, fig. 8.

Family II.
OCULINID.

Corallum composite, produced by gemmation, and presenting in general an abundant,
compact coenenchyma or common tissue, the surface of which is smooth, delicately striate
near the calices, or slightly granular, but never echinulate. Valls of the corallites com-
plete (that is to say, presenting no perforations), not distinct from the ccenenchyma, and
increasing by their internal surface, so as to mvade progressively the inferior part of the
visceral cavity, and to fill it up more or less completely in old age. Loculi imperfectly
divided by a few dissepiments ; no synapticule. Szpta entire, or having their upper edge

slightly divided.
§ 1. Septa of various sizes, forming distinct cycla.

1. Genus OcULINA.

(Pars) Lamarck, Hist. des An. sans Vert., t. ii, p. 283, 1816; Milne Edw. and J. Haime, Comptes
rend. de l’Ac. des Sc., t. xxix, p. 68, 1849.

Corallum in general arborescent ; gemmation irregular or affecting a spiral disposition ;
ccenenchyma highly developed ; its surface smooth, excepting near the calices, where it
presents slight radiating strie. Corallites with the calice very deep; a columella well
developed, papillose at its apex, and becoming compact towards its basis. Pa/i corresponding
to all the septa, excepting those of the last cyclum. Scepfa almost entire, slightly exsert,
and very unequally developed.

Typ. sp., Oculina virginea, Lamarck, An. sans Vert., p. 289; Madrepora virginea, Ellis and Sol., tab. xxxvi.

2. Genus TRYMHELIA.

Milne Edw. and J. Haime, Comptes rend. de l Académie des Sciences, t. xxix, p. 68, 1849.

Corallum arborescent, differmg from Oculina by the non-existence of a colwmellu, ond the
great development of the yal, which are cemented together, so as to form a verticai tube.

Typ. sp., Zrymhelia eburnea, Milne Edw. and J. Haime, loc. cit., p. 68.

XX BRITISH FOSSIL CORALS.

3. Genus CYATHELIA.
Milne Edw. and J. Haime, loc. cit., p. 68, 1849.

Corallum arborescent ; gemmation terminal and regularly opposite. Corallites free to a
considerable distance from the calice, which are grouped in a way similar to that of flowers
constituting a dichotomous cyme. Colwmella large and papillose. Pali well developed.
Septa entire, exsert, and strongly granulated.

Typ. sp., Cyathelia axillaris, nob. ; Madrepora axillaris, Ellis and Solander, tab. xiii, fig. 5.

4. Genus ASTRHELIA.
Milne Edw. and J. Haime, loc. cit., p. 68, 1849.
Corallum in general arborescent, and resembling Oculina by its form and its mode of

gemmation, but differing from the three preceding genera by the non-existence of pali.
Calice with a deep central fossula. Colwmella septal ; edges of the septa denticulated.

Typ. sp., Astrhelia palmata, nob.; Madrepora palmata, Goldfuss, tab. xxv, fig, 6.

5. Genus SYNHELIA.
Milne Edw. and J. Haime, loc. cit., p. 68, 1849.

Corallum arborescent, with thick branches ; gemmation irregular. Calices very shallow,
their border scarcely projecting above the surface of the ccenenchyma, and united by com-
mon striz. Columella compact, styliform, and terminated by a small tubercle. Septa
scarcely exsert.

Typ. sp., Synhelia gibbosa, nob. ; Lithodendron gibbosum, Goldfuss, op. cit., tab. xxxvii, fig. 9.

6. Genus ACRHELIA.
Milne Edw. and J. Haime, op. cit., p. 69, 1849.

Corallum arborescent, or forming a ramified cluster ; gemmation pretty regularly spiral.
Surface of the coenenchyma smooth, excepting in the immediate vicinity of the calices, where
slight traces of radiating cost are perceptible. Scpta extremely exsert, lanceolate, and
entire ; the principal ones uniting towards the lower part of their mer edge, without there
being either a columella or pali m the centre of the visceral chamber.

Typ. sp., Acrhelia Seba, Milne Edw. and J. Haime, loc. cit., p. 69; Seba, Thes., vol. iii, tab. cxvii, fig. 5.

7. Genus LOPHELIA.
Milne Ecw. and J. Haime, loc. cit., p. 69, 1549.

Corallum arborescent, segregate, with coalescent branches; no true coenenchyma, but
walls very thick ; gemmation irregularly alternate and subterminal. Ca/zces with a reverted
lamellar border. Sepéa entire, exsert, and uniting at the bottom of the visceral chamber as
in the preceding genus. No colunclla nor palt.

Typ. sp., Lophelia prolifera, nob.; Ellis and Sol., tab. xxxii, fig. 2; Oculina prolifera, Lamarck, An.
sans Vert., vol. ii, p. 286.

INTRODUCTION. XX1

8. Genus AMPHELIA.

Milne Edw. and J. Haime, loc. cit., p. 69, 1849.

Corallum arborescent, with coalescent branches, and well-developed ccenenchyma in
aged parts; gemmation subterminal, regularly alternate. Calice deep. Columella
rudimentary. Septa slightly exsert, entire, and small. No distinct coste; the surface of
the corallum smooth or very delicately striated.

Typ. sp., Amphelia oculata, nob. ; Madrepora oculata, Esper, tab. xii.

9. Genus DIPLHELIA.

Corallum resembling Amphelia, but having a large columella and denticulated septa.
Typ. sp., Diplhelia raristella, nob. Oculina raristella, Defrance, Dict. des Sc. Nat., vol. xxxv, p. 356.

10. Genus ENALLHELIA.

D’Orbigny MSS.; Milne Edw. and J. Haime, loc. cit., p. 69, 1849.

Differs from Amphelia by the shallowness of the calices, a greater development of the
septa, and the existence of long costal striz.

Typ. sp., Enallhelia compressa, D’Orbigny ; Lithodendron compressum, Goldfuss, op. cit., tab. xxxvii,
fig. 11.

§ 2. Septa equally developed, and forming apparently a single cyclum.

ll. Genus AXHELIA.

Milne Edw. and J. Haime, Compt. rend., t. xxix, p. 69, 1849.

Corallum arborescent, with coalescent branches, and a well-developed ccenenchyma, the
surface of which is entirely covered with sub-granulose strie. Calices very shallow.
Columella compact, very thick, and terminated by a rounded tubercle. No pali. Septa
exsert, entire.

Typ. sp., Avhelia myriaster, nob. ; Oculina myriaster, Valenciennes MSS., Catal. of the Museum of
Nat. Hist. of Paris.

12. Genus CRYPTHELIA.
Milne Edw. and J. Haime, loc. cit., p. 69, 1849.

Corallum arborescent, flabellate, and unifacial, all the corallites opening on one of the
surfaces of the flabellum ; surface of the branches quite smooth. Calices very prominent,
pediculate, explanate, and folded in two. No culwmella nor palit.

Typ. sp., Crypthelia pudica, Milne Edw. and J. Haime, loc. cit., p. 69.

XX BRITISH FOSSIL CORALS.

13. Genus ENDHELIA.
Milne Edw. and J. Haime, loc. cit., p. 69, 1849.

Corallum of the same general form as in the preceding genus, but with the corallites
alternate on the branches, which are thick and coalescent. Cadices immersed ; their border
not projecting, but armed with a tongue-shaped process. No columella nor pali.

Typ. sp., Endhelia Japonica, Milne Edw. and J. Haime, loc. cit., p. 69 (Mus. of Leyden).

14. Genus STYLASTER.

Gray, Zool. Miscel., p. 36, 1831; Allopora, Ehrenb., Cor. Roth. Meeres, p. 147, 1834; Dana, op. cit.,
p- 693, 1846; Milne Edw. and J. Haime, loc. cit., p. 69, 1849.

Corallum arborescent and subflabellate ; ccenenchyma highly developed, smooth, and
presenting certain excrescences or tubercles, the nature of which is problematic. Calices
rare and not projecting much. Neither colwmella nor palt.

Typ. sp., Stylaster rosea, Gray, loc. cit.; Oculina rosea, Lamarck, op. cit., t. xi, p. 287; Esper.,

tab. xxxvi.

Transitional Group.
PSEUDOCULINID A.

Corallum composite, with a highly developed, spongy, or cellulose, echinulate, dermic,
coenenchyma. Costal apparatus rudimentary. Walls imperforate, and never invading the
visceral cavity. Septal apparatus well developed; dissepiments few im number.

This small group participates of the characters belonging to the Oculimide and the
Astreide, but differs essentially from both. It does not, however, present any important
structural peculiarity, and does not appear to be derived from a special zoological type.

1. Genus MapRAcis.
Milne Edw. and J. Haime, Comptes rend. de I’ Acad. des Sc., t. xxix, p. 70, 1849.

Corallum arborescent ; coenenchyma almost compact, and highly echinulated. Calices
unarmed. Columella styliform. Septa exsert and equally developed.

Typ. sp., Madracis asperula, Milne Edw. and J. Haime, loc. cit., p. 70; Dentipora asperula, Gray,
MSS. British Museum.

2. Genus STYLOPHORA.

Schweigger, Beobacht. auf Natur., t. v, 1819; Sideropora and Stylopora, Blainville, Manuel d’ Actinologie,
p- 348, 1830; Sideropora, Milne Edw. and J. Haime, loc. cit., p. 70, 1849.

Corallum arborescent ; ccenenchyma sub-compact, with a granulated surface. Calices
armed with a labial process near the upper part of their margin. Columella styliform.

Typ. sp., Stylophora pistillaris, Schweigger, loc. cit.; Madrepora pistillaris, Esper., tab. 1x.

INTRODUCTION. XX1L

3. Genus DENDRACIS.
Milne Edw. and J. Haime, Comp. rend., t. xxix, p. 70, 1849.

Corallum arborescent ; coenenchyma almost compact, with its surface granulated. Calices
sub-mammiform. No columella. Septa not exsert, or only very slightly so ; nearly equal.

Typ. sp., Dendracis Gervilli, nob. ; Madrepora Gervillii, Defrance, Dict. des Sc. Nat., vol. xxviii, p. 8;
Michelin, Icon., Zooph., pl. xlix, fig. 8.

4. Genus AR#ACIS.
Milne Edw. and J. Haime, loc. cit., p. 70, 1849.

Corallum massive; ccenenchyma spongy, with its surface echinulate. Calices with a
thin projecting margin. No columella. Septa wnequally developed, entire.

Typ. sp. Areacis spheroidalis, nob.; Astrea spheroidalis, Michelin, pl. xliv, fig. 9.

Family If.
ASTREID A.

Dana, Exploring Expedition, Zooph., p. 194, 1846.

Corallum composite or simple, circumscribed by imperforated walls, and often
increasing by fissiparity. Corallites becoming tall by the progress of their growth; each
individual or series of individuals well defined, and separated from the others by perfect
walls. Ccenenchyma not existing, or being formed either by the development of the coste
and their dissepiments, or by the epithecal tissue alone, and not forming a compact mass
as m the Oculinidee. The visceral chamber never obliterated inferiorly by the growth of
the walls, but subdivided and more or less completely closed up by the interseptal dis-
sepiments, which are in general very abundant ; never any synapticule like those of the

Fungide.

First Tribe—EUS MILIN AE.

Septa completely developed and entire (that is to say, with their apical margin neither
lobate nor denticulate). Coste always unarmed. Colwmella often compact, or even styliform.

Section 1—EUSMILINAL PROPRLA#.

Corallum simple or composite, and in that case formed by distinct corallites, affecting
an arborescent disposition, fasciculate, or presenting a lmear arrangement ; free laterally,
at least in a great part of their length, and never having their calices blended together.
Reproduction usually fissiparous in the compound species.

XX1V BRITISH FOSSIL CORALS.

1. Genus CYLICOSMILIA.
Milne Edw. and J. Haime, Ann. des Sc. Nat., 3™° serie, t. x, p. 232, 1848.

Corallum simple, adherent. Colwmella well developed, and of a spongy structure.
Septa thin, slightly exsert, covered laterally with small granulations, and closely set.
Dissepiments very abundant. Vail thin, with a rudimentary epitheca. Coste simple, not
ramified, and distinct down to the basis of the corallum.

Typ. sp., Cylicosmilia altavillinsis, Milne Edw. and J. Haime, loc. cit., p. 233 ; Caryophyllia altavillinsis,
Defrance; Michelin, Icon. Zooph., tab. Ixxiv, fig. 2.

2. Genus PLACOSMILIA.
Milne Edw. and J. Haime, loc. cit., p. 233, 1848.

Corallum simple, compressed, free, and subpediculate. Calice more or less elliptical.
Columella lamellar. Septa numerous, closely set, slightly exsert, and not much granulated ;
systems apparently very numerous. Dzsseprments abundant. Wall naked, or with a rudi-
mentary epitheca. Coste simple, not ramified, and distinct from the basis of the corallum.

Typ. sp., Placosmilia cymbula, Milne Edw. and J. Haime, loc. cit., p. 234; Turbinolia cymbula, Michelin,
Icon., pl. lxvii, fig. 1.

3. Genus TROCHOSMILIA.
Milne Edw. and J. Haime, loc. cit., p. 236, 1848.

Corallum simple, subpedicellate or adherent. Cadice nearly horizontal. No columella.
Septa meeting in the centre of the visceral chamber, numerous, and closely set; systems
apparently very numerous. Dissepiments abundant. Wall naked, or with a rudimentary
epitheca. Coste simple, granulated, delicate, usually distinct from the basis, and never
ramified.

Typ. sp., Trochosmilia Fawasii, Milne Edw. and J. Haime, loc. cit., tab. v, fig. 6.

4. Genus PARASMILIA.
Milne Edw. and J. Haime, loc. cit., p. 243, 1848.

Corallum simple, adherent or pedicellate, tall, subturbimate, and presenting in general
indications of an intermittent growth. Calice nearly circular; fossula not very deep. Columella
spongy. Septa exsert, very granular laterally, and arched at their apex. Dissepiments not
abundant, and existing only in the inferior part of the loculi. Wall naked, or with a rudi-
mentary epitheca. Coste straight, simple, not ramified, somewhat granulated, and m
general projecting more near the calice than in the lower part of the coral.

Typ. sp., Parasmilia centralis, Milne Edw. and J. Haime, loc. cit.; Madrepora centralis, Mantell,
Geol. of Sussex, tab. xvi, figs. 2, 4.

INTRODUCTION. XXV

5. Genus C@LOSMILIA.

Differs from Parasmilia by not having any rudiments of a columella.

Typ. sp., Calosmilia poculum ; Parasmilia poculum, Milne Edw. and J. Haime, loc. cit., tab. v, fig. 5.

6. Genus LOPHOSMILIA.
Milne Edw. and J. Haime, loc. cit., p. 246, 1848.

Corallum simple, subturbinate, adherent. Calice almost circular. Colwmella lamellar,
small. Septa very exsert, unequal; their apical margin highly arched, and their sides
granular; the six systems equally developed. Vail naked. Coste simple, and but
slightly marked ; growth not intermittent.

Typ. sp., Lophosmilia rotundifolia, Milne Edw. and J. Haime, loc. cit., tab. v, fig. 3.

7. Genus DIPLOCTENIUM.
Goldfuss, Petref. Germ., p. 50, 1826-30.

Corallum simple, extremely compressed, flabelliform, free, but retaimimg a thick
peduncle. Calice representing a very long ellipse, arched so strongly that the extremities
of its long axis descend much below the level of its small axis; fossula very narrow, very
long, and shallow. No columella. Septa extremely numerous, nearly equal, thin, very closely
set, and slightly exsert. Dissepiments simple and numerous. Walls naked. Coste
extremely numerous, narrow, crowded, nearly equal, distinct from the basis, and dichoto-
mosing, or even dividing into three branches as they rise.

Typ. sp., Diploctenium lunatum, Michelin, Icon. Zooph., tab. lxy, fig. 8 ; Madrepora lunata, Bruguictre,
Journ. d’Hist. Nat., vol. i, tab. xxiv, figs. 5, 6.

8. Genus MONTLIVALTIA.
Lamouroux, Exposit. Méthod. des Genres de Polypiers, p. 78; Milne Edw. and J. Haime, loc. cit., p. 250.

Corallum simple, adherent, or sub-pedicellate. No columella. Septa exsert, in general
numerous and crowded, very broad, and forming apparently twelve or more cycla. Wail
covered by a highly-developed membraniform epitheca ; growth not intermittent.

Typ. sp., Montlivaltia caryophyllata, Lamouroux, op. cit., tab. lxxix, figs. 8, 9, 10; Michelin, op. cit.,
tab. liv, fig. 2.

9. Genus PEPLOSMILIA.

Corallum resembling Montlivaitia, but having a large, lamelliform columella.

Typ. sp., Peplosmilia Austenii, nob.
4.

X\VI BRITISH FOSSIL CORALS.

10. Genus AXOSMILIA.
Milne Edw. and J. Haime, loc. cit., p. 261, 1848.

Coralluim simple, free m the adult state, tall, turbinate. Ca/ice circular ; fossula large and
deep. Columella styliform, lavge, and slightly con:pressed. Sepa neither exsert nor crowded,
deheately granulated, and all, excepting those of the youngest cyclum, cemented to the
columella ; loculi deep. Walls entirely covered by a membraniform epitheca, presenting
strong transverse folds, and extendiny to the edge of the calice.

Typ. sp., Awxosmilia extinctorium, Milne Edw. and J. Haime, loc. cit.; Caryophyllia extinctorium,
Michelin, op. cit., tab. ix, fig. 3°.

Ll. Genus Kusmivia.
Milne Edw. and J. Haime, loc. cit., p. 262, 1848.

Coralluin composite, cespitose, with dichotomous or trichotomous branches, and a stem
that does not thicken much by the progress of age. Corallites multiplying by fissiparity,
hecoming rapidly segregate, and not remaining disposed in series at their calicular extremity.
Calices vather regular in form, but im general nearly circular ; fossula deep. Colwmella of a
loose lamello-spongiate texture. Sepfa exsert, broad, thin, straight, not crowded, with their
apex strongly arched, and their surface almost smooth. Dzssepiments well formed, but not
very abundant. Walls naked or covered inferiorly by a slight pellicular epitheca. Coste
indistinct towards the basis of the corallites, but becoming sub-cristiform near the calice.

Typ. sp., Lusmilia fastigiata, Milne Eew. and J. Haime, loc. cit., tab. v, fig. 1; Madrepora fastigiata,
Pallas, Eleuch. Zooph., p. 301.

12. Genus APLOSMILIA.
D’Ovbigny MSS.
Corallum composite, and having the characters of Eusmilia, excepting that the columella
is. lamellar.

Typ. sp., Aplosmilia aspera, D’Orbigny MS8S.; Lobophyllia aspera, Michelin, Icon., tab. xx, fig. 4 ;
Eusmilia (2) aspera, Milne Edw. and J. Haime, loc. cit., p. 266.

13. Genus LeprosM1itA.
Milne Edw. and J. Haime, loc. cit., ». 267, 184°.

Coralltm composite, cespitose, fissiparous, and presenting the same general disposition as
in the preceding genus. No columella. Septa extremely thin, crowded, broad, very slightly
exsert, with their apex slightly arched, and their lateral surfaces sub-glabrous. Déssepo-
ments very abundant. Walls very thin, plain towards the basis, and costulate near the
calices.

Typ. sp., Leptosmilia ramosa, Milne Edw. and J. Haime, loc. cit., tab. vi, fig. 1.

14. Genus THECOSMILIA.
Milne Edw. and J. Haime, loc. cit., p. 270, 1848.

Corallum composite, cespitose, fissiparous, and affecting the same general disposition as

INTRODUCTION. XXVII

in the two preceding genera. No columella. Septa closely set, not remarkably thin, slightly
exsert, and granulate. Walls covered with a strong epitheca, reaching almost to the margin
of the calices.

Typ. sp., Thecosmilia trichotoma, Milne Edw. and J. Haime, loc. cit.; Lnthodendron trichotomum,
Goldfuss, Petref. Germ., tab. xiii, fig. 6.

15. Genus BARYSMILIA.
Milne Edw. and J. Haime, loc. cit., p. 273, 1848.

Corallum composite, increasing by fissiparity, and forming a very thick stem, on the
apex of which the corallites become distinct, and are disposed in transverse series. Co/umella
rudimentary or not existing. Septa closely set. Walls very thick, naked, and covered with
delicate costal lines, which are nearly equal and granulate.

Typ. sp., Barysmilia Cordieri, Milne Edw. and J. Haime, loc. cit., tab. v, fig. 4

16. Genus DENDROSMILIA.
Milne Edw. and J. Haime, loc. cit., p. 274, 1848.

Corallum composite, somewhat arborescent, and increasing by lateral gemmation.
Corallites with large septa, and a spongious columella.
Typ. sp., Dendrosmilia Duvaliana, Milne Edw. and J. Haime, loc. cit., p. 274.

17. Genus STYLOSMILIA.
Milne Edw. and J. Haime, loc. cit., p. 275, 1848.

Corallum composite, fasciculate, and increasing by lateral gemmation. Corallites tall,
with a small number of thick septa, and a styliform columella. Walls thick, with obsolete
cost@.

Typ. sp., Stylosmilia Michelinii, Milne Edw. and J. Haime, loc. cit., p. 275, pl. vi, figs. 2, 2°

18. Genus PLACOPHYLLIA.
D’Orbigny MSS.

Corallum composite, segregate, and increasing by gemmation, which is almost basal.
Corallites cylindrical and low. Columella well developed. Septa probably entire. Walls
completely covered with a membraniform epitheca, presenting thick transverse folds.

Typ. sp., Placophyllia dianthus, D’Orbigny MSS. ; Lithodendron dianthus, Goldfuss, Petref. Germ.,
tab. xii, fig. 8.

Section I1—EUSMILINA’ CONFLUENTES.

Corallum composite, and presenting no separation between the corallites, united m
rows, so as to assume a meandriform disposition ; multiplication essentially fisstparous.

XXVIl BRITISH FOSSIL CORALS.

19. Genus CTENOPHYLLIA.
Dana, Zoophytes, p. 169, 1846.

Corallum pedunculate, but increasing very little by its basis, and terminated by a large
oval, almost flat, calicular surface ; the different series of corallites intimately united together
by means of their common walls, and without there being im general any coenenchyma ; the
gyri or calicular grooves very long, and the mural ridges thin. Colwmella lamellar, and
almost uninterrupted from one end of the gyrus to the other. In general, some traces of
pali. Septa rather closely set, shghtly exsert, and delicately granulated. Dossepiments very
abundant, arched, and oblique; sometimes simple, but in general producing a vesicular
mass. The common epitheca rudimentary, and covering only the inferior part of the common
exterior walls, in the upper part of which are coste, nearly equal, and more or less cristiform
near the margin of the calicular surface.

Typ. sp., Ctenophyllia meandrites, Milne Edw. and J. Haime, Ann. des Sc. Nat., 3™° série, vol. x, p. 277 ;
Meandrina pectinata, Lamarck ; Madrep. meandrites, Solander and Ellis, Zooph., tab. xlviui, fig. 1.

20. Genus DENDROGYRA.
Ehrenberg, Corall. des Roth. Meeres, p. 100, 1834.

Corallum composite, having the form of a thick, massive, vertical column, in which the
corallites. are placed perpendicularly to the axis, and constitute very tortuous gyri,
completely united by their walls ; mural ridges broad, flat, and compact ; grooves shallow.
Columella highly developed, and formed by a series of very compact, enlarged processes.
Septa very thick and closely set. Dissepiments large, but not crowded.

Typ. sp., Dendrogyra cylindrus, Ehrenb., op. cit. ; Milne Edw. and J. Haime, Ann. des Sc. Nat., 3™ série,
Zool., t. x, pl. vi, fig. 9.
21. Genus RuIPiDOGYRA.

Milne Edw. and J. Haime, loc. cit., p. 281, 1848.

Corallum composed of a single series of corallites and constituting a flabelliform or
tall tortuous mass, the lateral walls of which are always free from top to bottom. Colwmella
lamellar, but almost rudimentary. Septa exsert and crowded. Dissepiments abundant.
Coste delicate, in general subcristate near the margin of the calice. No epitheca, or only
a rudimentary one.

Typ. sp., Rhipidogyra flabellum, Milne Edw. and J. Haime, loc. cit.; Lobophyllia flabellum, Michelin,
Icon., tab. xviii, fig. 1.
22. Genus PACHYGYRA.

Milne Edw. and J. Haime, loc. cit., p. 468, 184.

Corallum adherent by a very thick peduncle; gyri with a narrow calicular groove,
and united by a very broad mass of dense ccenenchyma. Colwmella lamellar. Septa
crowded. Coste delicate and granulated ; little or no epitheca.

Typ. sp., Pachygyra labyrinthica, Milne Edw. and J. Haime, loc. cit. ; Lobop. lia labyrinthica, Michelin,
Icon., pl. Ixvi, fig. 3.

INTRODUCTION. XX1X

23. Genus PLEROGYRA.
Milne Edw. and J. Haime, loc. cit., p. 284, 1848.

Corallum composed of long, thick, slightly ramified gyri, united laterally by their lower
part, and free only near the calicular margin. No columella. Septa exsert, and broad ;
interseptal loculi very broad, and almost entirely filled up with large vesicular dissepiments,
constituting a cellular mass. Walls presenting some costa/ strize near the calicular margin,
but covered in all the other parts by a vesicular structure, which becomes highly developed
between the gyri.

Typ. sp., Plerogyra laxa, Milne Edw. and J. Haime, loc. cit., tab. vi, fig. 8.

Section II] —EUSMILINA) AGGREGAT A.

Corallum composite and massive, in which the corallites are not arranged in series,
and although remaining quite distinct, are united together by their walls, by a costal
ccenenchyma, or by mural annular expansions.

This group corresponds to the division of the Astrecne aggregate of the second tribe of
this family, and constitutes with these the great genus Astrea of most authors.

24. Genus STYLINA.
Lamarck, Hist. des Anim. sans Vert., t. ii, p. 220, 1816; Fuscicularia, Lamarck, Extrait du Cours, 1812.

Corallum glomerate, astreiform. Corallites very tall, united by means of the costal
system and its dissepiments, and having the appearance of small truncate cones at their
upper end. Calices circular, with their margin free; usually distant from each other.
Columella styliform and projecting. Sepfa exsert, arched at their apex; in general not
numerous, and forming as usual six systems. No pali. Walls thick.

Typ. sp., Stylina echinulata, Lamarck, loc. cit.; Milne Edw., Atlas du Regne Animal de Cuvier,
Zooph., pl. Ixxxv, fig. 3.

25. Genus STYLOCE@NIA.
Milne Edw. and J. Haime, An. des Sc. Nat., 3™° série, t. x, p. 298, 1848.

Corallum having the form of a very thick sheet, convex or bent in different ways ;
covered inferiorly by a finely-striated epitheca; and mcreasig by marginal gemmation.
Corallites united by their walls, which are thin and prismatic. Calices polygonal, thew
margins simple, and bearing at their angles small, columnar, grooved processes. Colwmella
styliform, projecting. Sepa very thin, not exsert, nor numerous, and forming six systems.

Typ. sp., Stylocenia emarciata, Milne Edw. and J. Haime, loc. cit., tab. vii, fig. 2; Astrea emarciata,
Lamarck, op. cit., t. li, p. 266.

200K BRITISH FOSSIL CORALS.

26. Genus ASTROC@NIA.
Milne Edw. and J. Haime, loc. cit., p. 296, 1848.

Corallum very dense, and not bearing columnar processes, as in the preceding genus.
Calices polygonal. Columella styliform, not projecting much. No pal. -Septathick ; ap-
parently eight or ten systems, two or four of the secondary septa being as much developed
as the six primary ones. Wadls thick and united, as in Styloccenia.

Typ. sp., Astroceenia Orbignyana, Milne Edw. and J. Haime, loc. cit., p. 297; Astrea formosissima,
Michelin, Icon., pl. Lxxii, fig. 9.

27. Genus STEPHANOCENIA.
Milne Edw. and J. Haime, loc. cit., p. 300, 1848.

Corallum glomerulate ; the corallites united by their walls, which are thick and
compact ; gemmation lateral and marginal. Calices subpolygonal. Colwmedla styliform, and
not projecting much. A coronet of pal, corresponding to the septa of the older cycla.
Septa scarcely exsert, granulated on their sides, and forming six systems, which are in
general equally developed.

Typ. sp., Stephanocenia intersepta, Milne Edw. and J. Haime, loc. cit., tab. vii; fig. 1; Astrea intersepta,
Lamarck, Anim. sans Vert., t. 1, p. 266.

28. Genus PHYLLOCENIA.
Milne Edw. and J. Haime, loc. cit., p. 469, 1848.

Corallum glomerate, astreiform. Corallites united by the coste and the exotheca, which
are highly developed. Calices with a free margin, slightly elevated. No columella, or only
traces of a rudimentary one. No palit. Septa very broad, exsert, and forming six
systems ; gemmation lateral.

Typ. sp., Phylloceenia irradians, Milne Edw. and J. Haime, loc. cit. ; Astrea radiata, Michelin, pl. xii, fig. 4.

29. Genus DicHOCENIA.
Milne Edw. and J. Haime, loc. cit., p. 305, 1848.

Corallum glomerate, astreiform. Cora/lites united by a very: abundant and dense costal
coemnenchyma, the upper surface of which is subgranulate. Cadices circular or elliptical, with
a projecting margin. Colwmella small. Pal corresponding to most of the septa, which are
exsert and granulated. Multiplication fissiparous.

Typ. sp., Dichoceenia porcata, Milne Edw. and J. Haime, loc. cit.; Astrea porcata, Lamarck, Anim. sans

Vert., t. i, p. 260.

INTRODUCTION. XXX1

30. Genus HEevERocentia.
Milne Edw. and J. Haime, loc. cit., p. 308, 1848.

Corallum resembling that of Sarcmula, but differmg from all the preceding genera
by the small number and the unequal development of the septa, which form in appearance
only three systems. In general, one of the three large primary septa is more developed
than the others, and remains sometimes alone in fossil species. Calices circular, with a
projecting free margin. No columella nor pal. Septa exsert ; ccenenchyma abundant, of
a foliate structure, and having a granular surface.

Typ. sp., Heterocenia exievis, Milne Edw. and J. Haime, loc. cit., tab. 1x, fig. 13 ; Lethodendron exigue,
Michelin, Icon., Zooph. tab. Ixxu, fig. 7.

Section [V—HUSMILIN A) IMMERSE.

Corallum composite. Corallites disposed as in the preceding Section, but imbedded in
an epithecal cellular tissue, and not united by costal laminz or mural annular expansions ;
gemmation lateral and basal ; reproduction never fissiparous.

31. Genus SARCINULA.

(In parte) Lamarck, Hist. des Anim. sans Vert., t. li, p. 222, 1816; Anthophyllum, Ehrenb., op. cit.,
p- 89, 1834.

Corallum fasciculate, and almost massive. Corallites tall, free towards their upper end,
which projects more or less above the surface of the cellular exotheca. Walls strong, with
coste but little developed. No columella, or only a rudimentary one. Septa very exsert.
Dissepiments in general simple, and not abundant.

Typ. sp., Sarcinula organum, Lamarck, loc. cit., p. 223; Milne Edw., Atlas du Régne Animal de
Cuvier, Zooph., pl. lxxxv, fig. 1.

Second Tribe—ASTREIN Ad.

Septa having their upper edge lobulated, dentate, or armed with spines, and often
imperfect near their inner edge. Coste also spinulous, dentate or crenulate, but never
forming simple cristz, as is often the case in Eusmilinee. Co/wmella in general spongy,
rarely lamellar, and never styhform. Corallum in general massive.

Section .— ASTREIN AL HIRT As.

Corallum simple or composite, and then formed by perfectly delineated corallites, pro-
duced by fissiparity, or by calicular gemmation.

XXX BRITISH FOSSIL CORALS.

32. Genus CARYOPHYLLIA.

(In parte) Lamarck, Hist. des Anim. sans Vert., t. ii, p. 224, 1816; Milne Edw. and J. Haime, Comptes
rend. de l’Ac. des Sc., t. xxvii, p. 491, 1848.

Corallum simple, and adherent by a broad basis. Calice circular, or almost so.
Columella well developed, spongy, and composed of twisted lamellae, that advance one over
the other. Septa broad, exsert, numerous, close set, and armed with spines, the size of
which augments from the centre of the calice towards its margin. Diéssepiments vesicular
and abundant. Wall presenting coste, formed by a series of spines; epitheca rudimentary.

Typ. sp., Caryophyllia lacera, Milne Edw. and J. Haime, An. des Sc. Nat., 3™° série, t. xi, p. 237;
Madrepora lacera, Esper, Pflanz., tab. xxv, fig. 2.

33. Genus CIRCOPHYLLIA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 491, 1848.

Corallum simple, subturbinate. Colwmella large and papillose. Septa broad, numerous,
exsert, with their calicular edge divided in small obtuse lobes. Dissepiments abundant,
vesicular, and arranged in spiral concentric lines. Coste thin, nearly equal, simple, and
delicately granulated.

Typ. sp., Circophyllia truncata, Milne Edw. and J. Haime, Ann. des Sc. Nat., 3° série, t. x, tab. viii,
fig.3; Anthophyllum truncatum, Goldfuss, Petref., tab. xiii, fig. 9.

34. Genus THECOPHYLLIA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 491, 1848.

Corallum simple, adherent, or sub-pedicellate. Calice circular, or nearly so. No colu-
mella. Septa very broad, in general slightly exsert, numerous, and armed with nearly equal
spiniform teeth. Wall covered with a thick, membraniform epitheca.

Typ. sp., Thecophyllia decipiens, Milne Edw. and J. Haime, Ann., t. xi, p. 241 ;. Anthophyllum decipiens,
Goldfuss, Petref., tab. lxv, fig. 3.

35. Genus LOBOPHYLLIA.
(Pars) Blainville, Dict. des Sc. Nat., t. lx, 1830; Milne Edw. and J. Haime, Ann., t. xi, p. 244.

Corallum composite, tall, and increasing by fissiparity. Coralhtes segregate, or united in
series, which are always simple, and free laterally. Ca/ice with a deep fossula. Columella
spongy. Sepfa numerous, exsert, very granular, and armed with strong marginal teeth, the
most external of which are the largest ; loculi shallow. Wails striated longitudinally, and
armed with spines ; epitheca rudimentary.

§ 1. Lobophyllia cymose. Typ. sp., Lobophyllia angulosa, Blainy. ; Caryophyllia angulosa, Lamarck.—
§ 2. Lobophyllia gyrose. Typ. sp., Lobophyllia multilobata, Milne Edw. and J. Haime, loc. cit., p. 250;
Fungus marinus, Seba, Rer. Nat. Thes., vol. iii, tab. cix, No. 4.

INTRODUCTION. XXX

36. Genus SYMPHYLLIA.
Milne Edw. and J. Haime, Comptes rend. de l’Ac. des Sc., t. xxvii, p. 491, 1848.

Corallum composite, massive, short, and increasing by fissiparity. Coradlites having dis-
tinct calicula, but united in linear series, which are cemented together laterally. The
other characters as in the preceding genus.

Typ. sp., Symphyllia sinuosa, Milne Edw. and J. Haime, Ann. des Sc. Nat., vol. x, tab. viii, fig. 7.

37. Genus MycnroPHyLiia.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 491, 1848.

Corallum massive, composed of corallites intimately united in series by their walls,
which are very thin. Lzterior common walls \obulate, spmmulous, and presenting but rudi-
ments of an epitheca. Calicular grooves, very shallow. No columella, or only rudiments of
one. Septa not numerous, scarcely exsert, strongly dentate, and confluent. Déssepiments
vesicular, large, and abundant ; loculi closed almost to their top.

Typ. sp., Mycetophyllia Lamarckiana, Milne Edw. and J. Haime, Ann. des Sc. Nat., vol. x, tab. viii, fig. 6.

38. Genus HUNOMIA.
Lamouroux, Exposit. Method. des Polypiers, p. 83, 1824.

Corallum cespitose, fissiparous; Corallites segregate, tall, cylindroid. Calices almost
circular. Columellarudimentary. Septa not verynumerous. Walls covered with a complete
membraniform epitheca, strongly striated transversely.

Typ. sp., Hunomia radiata, Lamouroux, op. cit., p. 83 ; Lithodendron Eunomia, Michelin, Icon., pl. xxxiv,
fig. 6; Eunomia levis, Milne Edw. and J. Haime, Ann., t. xi, p. 260; Lithod. leve, Michelin, loc. cit.,
pl. xix, fig. 8.

39. Genus CALAMOPHYLLIA.

Calamites, Guettard, Mém. sur les Sc. et les Arts, vol. ii, p. 404, 1770; Culamophyllia, Blainville, Dict. des
Sc. Nat., t. lx, p. 312, 1830.

Corallum fasciculate, cespitose, and dichotomous. Corallites very long and segregate.
Calices not very deep. Columella rudimentary or not existing. Septa thm, numerous,
crowded, and armed with apical teeth, the size of which increases from the margin towards
the centre of the calice. Dissepiments very oblique and crowded. Waills delicately striated,
devoid of epitheca, but presenting at certain points circular foliaceous expansions.

Typ. sp., Calamophyllia striata, Blainville, Dict. des Sc. Nat., pl. cccxii; Calamite strié, Guettarc,
Mém. sur les Sc., t. ili, pl. xxxiv.

XXXIV BRITISH FOSSIL CORALS.

40. Genus DASYPHYLLIA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 492, 1848.

Corallum fasciculate, cespitose, and dichotomous. Corallites very long and segregate.
Columella spongy. Septa thin, slightly exsert, and armed with apical teeth, the size of which
is much greater near the columella than towards the margin of the calice.

Typ. sp., Dasyphyllia echinulata, Milne Edw. and J. Haime, Ann. des Sc. Nat., 3° série, t. x,
pl. viii, fig. 5.

41. Genus CoLPOPHYLLIA.
Milne Edw. and J. Haime, Comptes rend. de |’Acad. des Sc., t. xxvii, p. 492, 1848.

Corallum sub-glomerate, remarkably light and fragile, composed of series of corallites
cemented together laterally, without their respective walls ceasing to be distinct on the
calicular surface, where they are parallel, very thin, and constitute a double ridge on each
side of the calicular trench. Calices individualized by the direction of their septa. Colwmella
rudimentary, or not existing. Sept extremely thin, broad, and slightly exsert ; their apical
edge armed with small delicate teeth, and emarginate near the middle. Dissepiments
very abundant, and closing up the loculi almost to the margin of the calice, and forming a
vesicular mass. Common exterior walls of the corallum or plate presenting small, lamellar,
nearly equal, denticulate cost ; epitheca rudimentary.

Typ. sp., Colpophyllia gyrosa, Milne Edw. and J. Haime, Ann., t. xi, p. 266; Madrepora gyrosa, Ellis
and Solander, op. cit.; tab. li, fig. 2; Manicina gyrosa, Ehrenberg, op. cit.; Mussa gyrosa, Dana, op.
cit., p. 186.

42. Genus OULOPHYLLIA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 492, 1848.

Corallum composed of a series of corallites, mtimately united by their lateral walls,
which constitute simple ridges between the trenches formed by the aggregate calices.
Columella spongy, and in general not highly developed. Sep¢a thin, slightly exsert, closely
set, and armed with numerous long, sharp, apical teeth, the size of which augments towards
the centre of the corailite. Common exterior walls sometimes covered with a thin epitheca ;
multiplication fissiparous.

Typ. sp., Oulophyllia Stokesiana, Milne Edw. and J. Haiine, Ann. des Sc. Nat., vol. x, tab. viii, fig. 10.

43. Genus LATOMEANDRA.
D’Orbigny MSS. ; Milne Edw. and J. Haime, Ann. des Sc. Nat., 3™° série, t. xi, p. 270, 1849.

Corallum having most of the characters of the preceding genus, but increasing by
calicular gemmation. The gyri in general short, the marginal ones distinct, and not forming
a common rim. No epitheca.

Typ. sp., Latomeandra plicata, Milne Edw. and J. Haime, loc. cit., p. 271; Lithodendron plicatum,
Goldfuss, Petref. Germ., tab. xiii, fig. 5.

INTRODUCTION. XXXV

44. Genus TRIDACOPHYLLIA.
Blainville, Dict. des Sc. Nat., vol. lx, p. 327, 1830.

Corallum short, and composed of corallites, arranged in series intimately united by their
lateral walls, which, instead of forming a simple ridge as in the preceding genera, constitute
very tall, foliaceous expansions, variously twisted, and terminated by asub-crenulate margin ;
the calicular trenches broad, very deep, and winding. Co/wmella quite rudimentary, but the
calicular centres very distinct. Septa projecting very little, thin, nearly equal, and serrate.
Dissepiments abundant, very oblique, convex, and forming long vesicules. Plate or exterior
surface of the common wall of the corallum covered with lamellar costee, which extend from
the basis of the mass, project slightly, and are irregularly denticulate.

Typ.sp., Tridacophyllia lactuca, Blainville, loc. cit. ; Concha fungiformis, Seba, Thes., v. 11, tab. cxxxix,
No. 10; Pavonia lactuca, Lamarck, An. sans Vert., vol. ii, p. 239; Manicina lactuca, Ehrenberg, op. cit., p. 103.

45. Genus ''RACHYPHYLLIA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 492, 1848.

Corallum short, increasing by fissiparity, and composed of very flexuous series of coral-
lites, free laterally. Common walls strongly echinulate. /pitheca rudimentary. Columella
well developed, but of a very loose, spongy texture. Septa numerous, crowded, exsert, and
strongly granulated laterally.

Typ. sp., Trachyphyllia amarantum, Milne Edw. and J. Haime, Ann. des Sc. Nat., 3"° sér., vol. xi,
p-. 275 ; Amarantum saxum, &c. Rumph. Amb. Hort. vi, tab. Ixxxvii, fig. 1.

46. Genus ASPIDISCUS.

Konig, Icon. Foss. Sect., p. 1, 1825; Cyelophyllia, Milne Edw. and J. Haime, Comptes rend. de |’ Acad.
des Sc., vol. xxvii, p. 492, 1848.

Corallum discoidal, with its inferior surface flat, and its upper surface convex. Corallites
arranged in radiating series, separated by thick and simple, crest-like mural ridges, excepting
towards the margin of the calicular surface, where the young individuals spread out so as to
form a broad, continuous, lamello-striate border. Colwmella rudimentary, but the calicules
well individualized. Septa very thin and crowded, but not numerous in each corallite.
common plate covered with a thick epitheca, presenting concentric striz or folds.

Typ. sp., Aspidiscus cristatus, Milne Edw. and J. Haime, Ann. des Sc. Nat., 3™° série, vol. xi, p. 277 ;
Aspidiscus Shawi, Konig, Icon. Foss. Sect., pl. i, fig. 6; Cyclolites cristata, Lamarck, Anim. sans Vert.,
t. li, p. 234.

47. Genus SCAPOPHYLLIA.
Milne Edw. and J. Haime, Comptes rend. de l’Acad. des Sc., t. xxvii, p. 492, 1848.

Corallum columnar, erect, very dense, and formed of corallites arranged in series,
completely united laterally. Colwmella tubercular, somewhat compact. Sepa very thick,
neither closely set nor numerous ; with their sides very echinulate, and the apex denticulate.
Dissepiments simple and distant.

Typ. sp., Scapophyllia cylindrica, Milne Edw. and J. Haime, Ann. des Sc. Nat., 3™° série, vol. x,
tab. viii, fig. 8.

XXXVI BRITISH FOSSIL CORALS.

Section I1.—ASTREINA, CONFLUENTES.

Corallum massive, increasing by fissiparity, and formed by a series of corallites, the
individuality of which is not distinct. The calices, thus united in a common trench, have
their septa arranged in a parallel manner in two lines; and the columella, when existing,
is continuous in the whole length of the series.

‘hese meandriform Corals much resemble the confluent Eusmiline, and in fossils where
the apical teeth of the septa may be worn away, it is often difficult to distinguish them. It
may therefore be useful to mention that, in the confluent Astreine, the gyri are always com-
pletely united laterally, and never more or less segregate, which is sometimes the case with
the confluent Eusmiline ; that the columella, which is generally spongy in the latter, never
presents that loose structure in this section ; and when it is lamellar, the septa are united to
it by an undivided margin in the confluent Eusmiline, and by a series of trabicule or
processes in the confluent Astreinz; lastly, that the sides of the septa are more or less
granulated in all these Astreinze, and are on the contrary almost glabrous in the meandroid
Kusmiline.

48. Genus MEANDRINA.

(Pars) Lamarck, Hist. des Anim. sans Vert., t. ii, p. 244, 1816; Milne Edw. and J. Haime, Comptes rend.,
t. xxvii, p. 493.

Corallum glomerate, adherent by a very broad basis, and having a very dense
structure. Gyri intimately united by their lateral walls, which constitute simple, compact
ridges, with a cristate apex. Calicular trenches very long. Colwmella much developed,
spongy and essential (that is to say, not arising from the septa, and distinct from the
bottom of the visceral chamber). Septa crowded, enlarging near the columella, and not
presenting any appearance of a paliform lobe. Plate or exterior common walls of the
corallum covered with a complete delicate epitheca.

Typ. sp., Meandrina filograna, Lamarck, loc. cit., vol. ii, p. 248; Michelin, Icon., pl. xi, fig. 7.

49. Genus MANICINA.

(In parte) Ehrenberg, Corall. des Roth. Meeres, p. 101, 1534; Dana, op. cit., p. 183, 1846; Milne Edw.
and J. Haime, Ann. des Sc. Nat., vol. xi, p. 285, 1849.

Corallum free or sub-pedicellate, in the adult state ; sub-turbinate when young, but
becoming convex, and massive. Gyri very long, and united by their walls, so as to
form simple ridges, as in the preceding genus ; the apex of the ridge cristate or sulcate.
Calicular trench broad and deep. Columella spongy, and even more developed than
in Meandrina. Septa thin, crowded, strongly granulated, and armed with delicate, equal
teeth ; a well-characterised paliform lobe arismg from the edge of the principal septa
near the columella. Plate or exterior common wall covered with thin and very delicately-
serrated coste ; its inferior part having an incomplete epitheca.

Typ. sp., Manicina areolata, Ehrenberg, loc. cit.; Madrepora areolata, Ellis and Solander, op. cit.,
tab. xlvii, fig. 5.

INTRODUCTION. XXXVH

50. Genus Dipioria.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 493, 1848.

Corallum glomerate, adherent by a broad basis, and of a dense structure. Gyrz long,
very sinuous, and united by highly-developed costa, and not by the walls themselves ;
ridges complex, presenting on each side a mural crest, and in the middle a broad concave
groove or ambulacrum, formed by the costz and their dissepiments. Co/umella spongy,
essential, and well developed. Szpfa strong, exsert, and armed with closely-set teeth,
the largest of which are near the walls.

Typ.sp., Diploria cerebriformis, Milne Edw. and J. Haime, Ann., t. xi, p. 289 ; Meandrina cerebriformis,
Lamarck, op. cit., vol. ii, p. 246; Seba Thes., vol. iii, tab. exii, No. 6.

51. Genus Leprorta.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 493, 1848.

Corallum glomerate, of a light spongy structure. Gyrz very long, and limited by
their walls, which are thin or cellulose, and form simple intercalicular ridges. Columella
lamellar ; its upper edge projecting slightly, and regularly lobated. Seta united to the
columella by means of marginal trabicule ; their upper edge shghtly exsert, and armed
with very small irregular teeth. Plate covered with a thin but complete common epitheca.

Typ. sp., Leptoria tenuis, Milne Edw. and Haime, Ann. des Sc. Nat., 3™° ser., vol. x, tab. viii, fig. 11;
Meandrina cerebriformis, Quoy and Gaimard, Voy. de |’ Astrol., Zooph., pl. xvii, figs. 2,3; Meandrina tenuis,
Dana, op. cit., p. 262; Milne Edw., Atlas du Régne An. de Cuvier, Zooph., pl. Ixxxiv ter, fig. 2.

52. Genus C@ortia.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 493, 1848.

Corallum resembling much the true Meandrina, but differmg from the four preceding
genera by its rudimentary co/wmella, which is not essential, but septal, and formed by
trabicule, springing from the margin of the septa. Gyri long, and united by their walls,
the tissue of which is cellular; ridges simple and continuous. Sepéa delicate, and having
neither a paliform lobe nor a lateral expansion near the columella.

Typ. sp., Caloria labyrinthica, Milne Edw. and J. Haime, Ann. des Sc. Nat., 3™° série, t. xi, p. 194;
Madrepora labyrinthica, Ellis and Solander, op. cit., tab. xlvi, figs. 3, 4.

53. Genus ASTRORIA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 493, 1848.

Corallum having the same structure as in the preceding genus, but formed of very short
gyri, the corallites tending to individualization. This form is intermediate between the
ordinary confluent Astreide (or Meandrinz) and the agglomerated Astreide, such as true
Astrea.

Typ. sp., Astroria dedalea, Milne Edw. and J. Haime, Ann. des Sc. Nat., t. xi, p. 297; Madrepora
dedalea, Ellis and Solander, op. cit., tab. xlvi, figs. 1, 2.

XXXVI BRITISH FOSSIL CORALS.

54. Genus HyDNOPHORA.

Fischer de Waldheim, Descrip. du Mus. Démidoff, vol. iii, p. 295, 1810; Monticularia, Lamarck, Hist. des
Anim. sans Vert., t. 1, p. 248, 1816.

Corallum formed of irregular series of corallites, united by their walls, which are thick,
compact, and constitute ridges, divided longitudinally, so as to represent rows of conical
prominences, or monticule. ‘The calicular trenches are transversal as well as longitudinal,
and there is no columella. Sepfa nearly equal, and rising to the apex of the conical mural
monticulae. General form sometimes massive and sub-globose or gibbous; sometimes
sub-explanate.

Typ. sp., Hydnophora Demidovii, Fischer, Oryct. du Gouv. de Moscou, pl. xxxii.

Section HI.—ASTREIN A) DENDROID A.

Corallum always increasing by lateral gemmation. ‘The corallites segregate, and having ~
an arborescent or fasciculate arrangement. Septa regularly and delicately serrated ; those
of the principal cycla always bearing pall.

55. Genus CLADOCORA.
(In parte) Ehrenberg, Corall. des Roth. Meeres, p. 85, 1834; Caryophyllia, Dana, Zoophytes, p. 378, 1846.

Corallum arborescent, forming branched clumps. Coradlites cylindrical, very long, and
completely free laterally. Calices circular, or almost so. Colwmella papillose. Pali well
developed, and corresponding to all the septa, except those of the last cyclum. Septa
slightly exsert, nearly equal, granulated, and having their apex arched and delicately serrated.
Walls compact, with simple, granulated, or echinulated costa, and an incomplete epitheca,
which often expands into circular, horizontal leaves, extending to the neighbouring
corallites.

Typ. sp., Cladocora cespitosa, Milne Edw. and J. Haime; Medrepora flexuosa, Solander and Ellis, tab.
xxxi, figs. 5, 6.

56. Genus PLEUROCORA.
Milne Edw. and J. Haime, Comptes rend. de |’Ac. des Sc., t. »xvii, p. 494, 1848.

Corallum sub-dendroid. Coradlites cylindrical, very short; united by their basal part,
and free towards their upper end. Columella, pali, and septa much as in the preceding
genus. Walls compact, extremely thick, and never presenting any traces of an epitheca.
Coste distinct from one end of the corallites to the other, and vermiculate.

This genus approximates in some degree to Dendrophyllia and to Oculina.

Typ. sp., Pleurocora explanata, Milne Edw. and J. Haime, Ann. des Sc, Nat., 3™° série, vol. x,
tab. vu, fig. 10.

INTRODUCTION. XXXIX

Section IV.—ASTREINA) AGGREGAT.

Corallum composite, massive, increasing by gemmation or by fissiparity, and in that
case not presenting a linear mode of arrangement of the corallites, which are always com-
pletely united laterally, but remain well defined, and never lose their individuality, as in
the confluent Astreina.

57. Genus ASTREA.

(In parte) Lamarck, Syst. des Anim. sans Vert., p. 371, 1801; Milne Edw. and J. Haime, Comptes rend.,
t. axvii, p. 494, 1848.

Corallum massive, in general convex or sub-globose. Gemmation extra-calicular. Coral-
lites tall. Calicules having a free, exsert, obtuse, circular margin; fossula not very deep.
Columella spongy, and not projecting at the bottom of the calicule. No pali. Septa
complete, exsert, broad, and strongly dentated or lobated ; the largest of their apical teeth
near the columella; their sides strongly granulated. Coste highly developed, and com-
posed of lamellee ; in general perforated, and united by numerous dissepiments.

Typ. sp., Astrea cavernosa, Milne Edw. and J. Haime, loc. cit., vol. x, tab. ix, fig. 1; Madrepora
cavernosa, Esper, Pflanz. Suppl. Mad., tab. xxxvii; Astrea argus, Lamarck, Hist. des Anim. sans Vert.,
t. x1, p. 259.

58. Genus CYPHASTRBA.
Milne Edw. and J. Haime, Comptes rend. de |’Acad. des Sc., vol. xxvii, p. 494, 1848.

Corallum massive, convex, and globose. Gemmation extra-calicular. Coradlites united
by a compact septal ccenenchyma, the surface of which is strongly granulated, or even
echinulated. Cal/icular rims as in the preceding genus. Co/wmella papillose, and well
developed. Sepa lamellar near the wall, but cmibriform towards the columella, where
they are formed by a series of oblique processes, representing a sort of lattice; their
calicular teeth rather larger towards the calice than near the walls.

Typ. sp., Cyphastrea microphtholma, Milne Edw. and J. Haime, Ann. des Sc. Nat., 3™° série, vol. x,
tab. ix, fig. 5; Astrea microphthalma, Lamarck, op. cit.

59. Genus OULASTREA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 495, 1848.

Corallum massive and incrustating. Gemmation extra-calicular. Corallites low.
Calices circular, with a free margin. Columella pepillose, and appearing to be formed by
the inner apical teeth of the septa. No pal. Septa with a crispate, denticulated, apical
margin, and echinulate sides. Coste also echinulate and crispate.

Typ. sp., Oulastrea crispata, Milne Edw. and J. Haime, Ann. des Sc. Nat., 3° série, vol. x, tab. ix,
fig. 4; Astrea crispata, Lamarck, Hist. des Anim. sans Vert., vol. ii, p. 265.

xl BRITISH FOSSIL CORALS.

60. Genus PLESIASTREA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 494, 1848.

Corallum globose; its under surface having the form of a naked costulated plate.
Gemmation extra-calicular. Calices with a free margin, and a fossula rather shallow.
Columella spongy. Pali well developed, and corresponding to all the septa except those
of the last cyclum. Septa exsert, formed by a well-developed lamina, and having a deli-
cately-serrated apex. Coste and their dissepiments in general well developed.

Typ. sp., Plesiastrea Urvillii, Milne Edw. and J. Haime, Ann. des Sc. Nat., vol. x, tab. ix, fig. 2;
Astrea galaxea, Quoy and Gaim., Voyage de l’ Astrolabe, Zooph., pl. xvii, figs. 10-14.

61. Genus LEPTASTREA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 494, 1848.

Corallum very dense and incrusting, and increasing by fissiparity, as well as by
extra-calicular gemmation. Costal coonenchyma quite compact. Calices in general much
crowded together, but preserving their margins distinct. Colwmella papiliose. Sepéa thin,
closely set, exsert, delicately granulated, and having their apical margin almost entire near
the walls, but delicately denticulated towards the columella. Dissepiments not very abun-
dant. Coste rather indistinct.

Typ. sp., Leptastrea Roissyana, Milne Edw. and J. Haime, Ann. Sc. Nat., 3° serie, vol. x, tab. ix, fig. 6.

62. Genus SOLENASTREA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 494, 1848.

Corallum forming in general a convex mass, of a light and cellular structure. Gem-
mation extra-calicular. Corallites long, slender, and united by an exothecal structure,
and not by the coste, which do not meet, and are often rudimentary. Calices circular,
with an exsert margin. Columella spongy, and in general small. Septa very thin; their
margin denticulated. Déssepiments simple, numerous, and closely set.

Typ. sp., Solenastrea Turonensis, nob. ; Astrea Turonensis, Michelin, Icon., pl. Ixxv, figs. 1, 2.

63. Genus PHYMASTREA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 494, 1848.

Corallum forming a convex or a horizontal mass. Corallites prismatical ; surrounded
from top to bottom by a thin epitheca ; very nearly approximated to each other, but not
united by their walls, and cemented together by means of a certain number of large
wart-like processes, so as to leave an empty space between them. Gemmation extra-
calicular. Oalices sub-polygonal, with a free margin. Colwmella spongy, well developed.
Septa large, slightly exsert, and strongly dentated. Wails thick ; no trace of costae.

Typ. sp., Phymastrea Valenciennesii, Milne Edw. and J. Haime, Aun. Sc. Nat., 3™° série, vol. x,
tab. ix, fig. 3.

INTRODUCTION. xli

64. Genus ASTROIDES.
Quoy and Gaimard, Ann. des Sc. Nat., 17° série, vol. x, p. 187, 1827; Astrgitis, Dana, Zooph., p. 405, 1846.

Corallum incrusting, and formed of corallites very unequally approximated; some
almost entirely free, others crowded so as to become polygonal, but always separated by a
more or less developed epitheca. Gemmation extra-calicular. Calices deep. Columella
spongy, large, and projecting very much at the bottom of the fossula, a character which
does not exist in any of the preceding Astreine. Septa not much developed, very thin,
not exsert, irregularly and delicately denticulated. Dissepiments very abundant. Walls
composed of a dense spongy tissue. Epitheca complete.

Typ. sp., Astroides calicularis, Blainville, Dict. des Se. Nat., vol. lx.; Caryophyllia calicularis, Lamarck,
op. cit., vol. ii, p. 226; Milne Edwards, Atlas du Regne Anim. de Cuvier, Zooph., tab. Ixxxiii, fig. 2.

65. Genus PRIONASTREA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 495, 1848.

Corallum forming a convex or gibbose mass, the under surface of which constitutes a
common plate, covered with a thin, complete epitheca. Gemmation sub-marginal. Calices
distinct, polygonal; fossula deep; margins united so as to form a simple crest between
the different corallites. Columella spongy. Septa thin, crowded, delicately granulated on
their sides, and strongly dentated at their apex; the largest of these teeth are those nearest
the columella. Dzssepiments well developed. Walls in general independent towards the
basis of the coral, but uniting to the adjacent ones near the calices, so that the visceral
chambers appear to be separated only by a single simple lamina.

Typ. sp., Prionastrea abdita, Milne Edw. and J. Haime, loc. cit.; Astrea abdita, Lamarck, Hist. des
Anim. sans Vert., t. i, p. 265, 1816; Madrepora aébdita, Soland. and Ellis, t. 50, f. 2.

66. Genus SIDERASTREA.
(In parte) Blainville, Dict. des Sc. Nat., t. lx, p. 335, 1830; Siderina, Dana, Zooph., p. 218, 1846.

Corallum incrusting, forming a convex mass of a very dense tissue. Gemmation
sub-marginal. Corallites united by their walls, which are thin, and sometimes indistinct.
Calices sub-pentagonal, with a deep fossula, and their margins rendered thick by the pro-
longation of the septa. Colwmella papillose, in general not much developed, but having a
tendency to become compact. Sepfa very closely set, thin, and regularly denticulated ;
their lateral surfaces covered with large granulations, which come in contact with those of
the adjoining septa, but are not united to them. Dzssepiments rudimentary.

Typ. sp., Siderastrea galaxea, Blainville, loc. cit.; Madrepora galaxeo. Ellis and Solander, Hist. of
Zooph., tab. xlvii, fig. 7.

67. Genus BARYASTREA.
Milne Edw. and J. Haime, Comptes rend. de l’Acad. des Sc., t. xxvii, p. 495, 1848.

Corallum incrusting ; its tissue very dense and compact. Gemmation marginal or
sub-marginal. Corallites very intimately united by their walls. Culices polygonal and
6

xl BRITISH FOSSIL CORALS.

indistinctly separated by superficial, narrow grooves. Colwmella not much developed at
its apex; but having a tendency to become compact, and to fill up the visceral chamber
towards its basis. Septa very thick, closely set, scarcely granulated, and very feebly
denticulated. Dissepiments little developed.

Typ. sp., Baryastrea solida, Milne Edw. and J. Haime, loc. cit.

68. Genus ACANTHASTREA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 495, 1848.

Corallum forming a slightly convex mass, with its upper surface strongly echinulate, and
its under surface constituting a plate, covered with a complete, thin epitheca. Gemmation
sub-marginal or marginal. Coradlites united by their walls, which are somewhat cellular.
Calices sub-polygonal, with broad, spiniferous, simple, common margins. Columella rudi-
mentary or septal. Sepa exsert, strong, and armed with projecting spiniform teeth, the
largest of which are situated near the walls, mstead of bemg the central ones, as in the
preceding genera. Dissepiments very numerous.

Typ. sp., Acanthastrea spinosa, Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 495.

69. Genus SYNASTREA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 495, 1848.

Corallum pediculate, and increasing in breadth more than in height. _Gemmation
sub-marginal. Coradlites intimately united by their walls. Calices superficial, distinct at
their centre, but not so towards their circumference. Columella very small. Septa con-
fluent, progressing from one calicular centre to another without imterruption, exsert, and
hiding the walls, over which they extend; their calicular margin almost horizontal, and
armed with nearly equal teeth.

Typ. sp., Synastrea Saviynyi, Milne Edw. and J. Haime, Ann. des Sc. Nat., 3™° série, vol. x,
tab. ix, fig. 12.

70. Genus THAMNASTREA.

Thamzasteria (in parte), Le Sauvage, Mém. de la Soc. d’Hist. Nat. de Paris, vol. 1, p. 241, 1822; Tham-
nastrea, ejusd., Ann. des Sc. Nat., 1"° série, vol. xxvi, p. 328.

Corallum having confluent septa, and most of the other characters of Synastrea, but
forming a fasciculus of columns or thick branches, erect, and of a more or less arborescent
aspect.

Typ. sp., Thamnastrea dendroidea, Le Sauvage, Mém. de la Soc. d’ Hist. Nat., vol. i, tab. xiv.

71. Genus GONIASTREA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 495, 1848.

Corallum always increasing by successive fissiparity, and forming a convex or lobulated
mass, of a dense structure. Corallites intimately united from top to bottom by thei walls,

INTRODUCTION. xhit

which thus form simple partitions between the visceral cavities, and are thick and compact.
Calices polygonal ; fossula rather deep. Columella spongy. Septa slightly exsert, their
apex arched and denticulated. Well-characterised, dehticulated pa/z, corresponding to all
the septa, except those of the last cyclum.

Typ. sp., Goniastrea solida, Milne Edw. and J. Haime, Ann., 3” série, t. x, pl. ix, fig. 7; Madrepora
solida, var. 6, Forskal, Descr. Anim. in Itin. Orient., p. 131.

72. Genus APHRASTREA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 495, 1848.

Corallum increasing by fissiparity, and forming a convex mass of a light cellular
structure, presenting on its under surface a complete, common epitheca Calices intimately
united by their margins, which thus assume the appearance of simple partitions. Colwmella
spongy. Pali or paliform lobes of the septa corresponding to all the cycla, except the last.
Septa denticulated, slightly exsert. Dzssepiments vesicular, and highly developed. Wadls
extremely thick, and completely vesicular.

Typ. sp., Aphrastrea deformis, Milne Edw. and J. Haime, Ann. des Sc. Nat., 3™° série, vol. x, tab. ix,
fig. 11; Astrea deformis, Lamarck, Hist. des An. sans Vert., t. xi, p. 264.

73. Genus PARASTREA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 495, 1848.

Corallum increasing by fissiparity, and having the same general form as in the preceding
genus, but differing from it by the mode of union of the corallites, which takes place by
means of the costz and their dissepiments, so that the calices, instead of being separated
only by a common simple margin, have each a distinct margin independent of those
surrounding it. Septa exsert, and armed with teeth, the largest of which are placed near
the centre of the calice, and often assume the appearance of pali. Dissepiments well
developed.

Typ. sp., Parastrea amicorum, Milne Edw. and J. Haime, Ann., 3™ série, vol. x, pl. ix, fig. 9.

Section V.—ASTREIN A REPTANTES.

Corallum increasing by the development of buds on stolons, or on membraniform
basal expansions. The corallites not umited by their sides, excepting accidentally by
means of their walls, and remaining short. Septa feebly denticulated. Dissepiments
almost rudimentary.

74. Genus ANGIA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 496, 1848.

Corallum composed of short, cylindrical corallites, united by a common gemmniferous
basal expansion, and completely free laterally. Ca/ices sub-circular ; fossula broad and

xliv BRITISH FOSSIL CORALS.

deep. Columella papillose, well developed. Septa thin, not exsert; the principal ones
having their upper margin almost entire, the others strongly dentated. Wadlls covered with
a complete epitheca.

Typ. sp., Angta rubeola, Milne Edw. and J. Haime, Ann. des Sc. Nat., 3™° série, vol. x, tab. vii, fig. 6;
Dendrophyllia rubeola, Quoy and Gaimard, Astrolabe, Zooph., tab. xv, figs. 12-15.

75. Genus CRYPTANGIA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 496, 1848.

Corallum composed of agglomerate, cylindro-turbmate corallites, which appear to mul-
tiply by gemmation on a non-persistent, soft stolon, so that they cease to be organically
united when in the adult state, but remain imbedded in an extraneous mass composed of
Cellepora. Calices circular, with a well-formed fossula. Columella papillose, well developed.
Septa thin, not very closely set; the upper edge of all of them dentate. Walls covered
with a complete epitheca.

Typ. sp., Cryptangia Woodii, Milne Edw. and J. Haime, loc. cit.; Cladocora cariosa, Wood, Ann. of
Nat. Hist., vol. xiii, p. 12.

76. Genus RHIZANGIA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 496, 1848.

Corallum increasing by the gemmation of stolons, which sometimes become calicified,
and are persistent. Corallites agglomerate, sub-cylindrical. Calices circular ; fossula shallow.
Columella papillose, and not very distinct from the neighbouring denticulations of the
septa. Septa thin, scarcely exsert, nearly equal, very closely set, with the upper edge
shghtly arched, and armed with small, regular teeth. Walls covered with a complete
epitheca, which extends almost as high as the apex of the septa.

Typ. sp., Rhizangia brevissima, Milne Edw. and J. Haime, Ann. des Sc. Nat., 3™° serie, vol. x, t. vu,
figs. 7, 8; Astrea brevissima, Deshayes, in Ladoucette, Hist. des Hautes Alpes, tab. xiii, fig. 13.

77. Genus ASTRANGIA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 496, 1848.

Corallum incrusting. Corallites very short, produced by gemmation on a thin,
common, basal expansion, the surface of which is granulated. Calices circular, with a
deep fossula. Columella papillose, sub-echinulate and not distinctly delimitated. Septa
thin, exsert, nearly equal, granulate, and armed with teeth much resembling those of the
columella; the tertiary septa bent towards those of the second cycla, and united to them.
Dissepiments in general simple and distant. Walls naked, with broad, delicately-granu-
lated costee.

Typ. sp.. Astrangia Michelinii, Milue Edw. and J. Haime, Aun. des Sc. Nat., 3™° serie, vol. x, t. vii, f. 5.

73. Genus PHYLLANGIA.
Milne Edw. and J. Haime, Comptes rend., t. xxvu, p. 497, 1848.

Corallum differmg from those of the preccding genus by the structure of the sepéa,
the upper edge of which is aimost euitire in the principal cycla, and slightly denticulated in

the others. Columella rudimentary.
Typ. sp., Phyllangia americana, nob.

INTRODUCTION. xlv

79. Genus OULANGIA.
Milne Edw. and J. Haime, Comptes rend., t. xxvii, p. 497, 1848.

Corallum composed of very low, cylindrical corallites, which appear to arise by gem-
mation on a basal incrusting expansion, and having their walls naked and costate, as
in the preceding genus, but with a highly-developed, papillose columella. Septa very
exsert, closely set ; those of the principal cycla having their upper edge almost entire.

Typ. sp., Oulangia Stokesiana, Milne Edw. and J. Haime, Ann. des Sc. Nat., 3™° série, vol. x,
tab. vii, fig. 4.

Aberrant Group.
PSEUDASTREID &.

Corallum composite, thin, and foliaceous, and increasing by extra-calicular gemmation.
Corallites short, well circumscribed, and dispersed on the surface of a common lamellar
plate. Ccenenchyma echinulate. Septa well developed, very echinulate. Dissepiments not
numerous. No synapticule. Common basal wall imperforate, sub-costulate, and naked.

Genus ECHINOPORA.

Lamarck, Hist. des An. sans Vert., vol. ii, p. 252, 1816; Echinastrea, Blainville, Dict. des Sc. Nat.,
vol. lx, p. 343, 1830.

Corallum adherent, near the centre, and expanding into large foliaceous, lobated
lamin. Calices circular, with an exsert margin.

Typ. sp., Echinopora rosularia, Lamarck, loc. cit., p. 253; Milne Edw., Atlas du Régne Anim. de
Cuvier, Zooph., tab. lxxxiii ter, fig. 1.

Transitional Group.
PSEUDOFUNGID/.
Corallum composite and foliaceous, having a perforated plate or basal wall (as in

Fungide) and interseptal dissepiments (as in Astreide). Calices forming radiating series,
separated by lobes or ridges. No synapticule.

Genus MERULINA.
Ehrenberg, Corall. des Roth. Meeres, p. 104, 1834. Typ. sp., Merulina ampliata, Ehrenberg, loc. cit.

Family IV.
FUNGID A.
Dana, Expl. Exped., Zooph., p. 283, 1846.

Corallum simple or composite, very short and expanding, so as to constitute a disc
or foliaceous lamina. Calice very shallow, and open laterally in simple species ;

xlvi BRITISH FOSSIL CORALS.

confluent, and not circumscribed in the compound species. Septa not distinct from the
coste, and formed by complete, imperforate laminze, with the edge dentate, and the sides
covered with styliform or echinulate processes, which, in general, meet so as to constitute
numerous synapaticule, or transverse props, extending across the loculi like the bars of
a grate. No dissepiments or tabulz, so that no part of the visceral chamber is completely
closed. Wa/’s basal, in general porous. The compound species increasing by sub-
marginal gemmation, and not by fissiparity.

First Tribe—CYCLOLITIN A.

Corallum simple. Plate or basal wall having a well-developed epitheca, presenting
concentric folds.

1. Genus CYCLOLITES.
Lamarck, Syst. des Anim. sans Vert., p. 369, 1801.

Corallum circular, or nearly so, and covered with an immense number of very thin
septa. Fossula oblong, narrow, and shallow. The small septa in general united to those
of the older cycla.

Typ. sp., Cyclolites elliptica, Lamarck, loc. cit., p. 234.

2. Genus PALMocYcLUs.
Milne Edw. and J. Haime, Comptes rend. de l’Acad. des Sc., vol. xxix, p. 71, 1849.

Corallum circular. Fossula deep, very broad, and circular. Septa thick and not
numerous ; none of them cemented together. ;

Typ. sp., Paleocyclus porpita, Milne Edw. and J. Haime, loc. cit.; Madrepora porpita, Fougt, Lin.
Ameen. Acad., t. i, tab. iv, fig. 5.

Second Tribe—FUNGIN 4.

Corallum simple or composite. Plate or basal wall without an epitheca, in general
strongly echinulate, and porous.

3. Genus FuNGIA.

(In parte) Lamarck, Syst. des An. sans Vert., p. 369, 1801; Dana, Zooph., p. 287, 1846; Milne Edw.
and J. Haime, Comptes rend., t. xxix, p. 71, 1849.

Corallum simple, subdiscoidal. Septa very numerous, and united so as to appear
ramified. Basal wad/ strongly echinulate, and perforated in an irregular manner.

Typ. sp., Fungia patellaris, Lamarck, loc. cit., p. 236; Milne Edw. and J. Haime, Ann. des Sc. Nat.,
teixs pl avij onal).

INTRODUCTION. xlvui

4. Genus MIcRABACIA.
Milne Edw. and J. Haime, Comptes rendus, vol. xxix, p. 71, 1849.

Corallum simple, lenticular, plano-convex. Sepfa not extremely numerous, straight.
Wail scarcely echinulate, and perforated in a regular manner.

Typ. sp., Micrabacia coronula, nob. ; Fungia coronula, Goldfuss, Petref. Germ., vol. i, tab. xiv, fig. 10.

5. Genus ANABACIA.
D’Orbigny MSS.; Milne Edw. and J. Haime, loc. cit., p. 71, 1849.

_ Corallum simple and lenticular. Sepfa extremely numerous, thin, and projecting on
the under side of the corallum without forming a distinct basal wall or plate. Fos-
sula shallow.

Typ. sp., Anabacia orbulites, nob. ; Fungia orbulites, Lamouroux, Exp. Meéth., tab. lxxxiii, figs. 1, 2, 3.

6. Genus GENABACIA.
Milne Edw. and J. Haime, Comptes rend., t. xxix, p. 71, 1849.
Corallum composite, formed by a parent corallite similar to Anabacia, bearing young
calicula arranged circularly.
Typ. sp., Genabacia stellata, nob. ; Fungia stellata, D’ Archiac, Mém. Soc. Géol. France.

7. Genus HERPOLITHA.

Eschscholtz, Isis, 1825 ; Haliglossa, Ehrenberg, Corall., p. 50, 1834 ; Herpetolithus, Leuckart ; Dana, Zooph.,
p. 306, 1846.

Corallum composite, free. Calicula sub-radiate, and of two sorts; the central ones
multi-lamellate, and arranged in a line ; the others pauci-lamellate, and dispersed irregularly.
Septa strong, and alternately thick and thin. Under surface of the common basal wall
very echinulate.

Typ. sp., Herpolitha limacina, nob. ; Madrepora pileus, Ellis and Solander, op. cit., tab. xlv.

8. Genus CRYPTABACIA.
Milne Edw. and J. Haime, Comptes rend., t. xxix, p. 71, 1849.

Corallum composite, free and convex above. Calices distinctly radiate ; the central
ones arranged in a line, and more distinct than the others. Septa short, and not numerous.
Under surface of the common basal walls strongly echinulated.

Typ. sp., Cryptabatia talpa, nob. ; Fungia talpa, Lamarck, Hist. des An. sans Vert., t. ii, p. 237.

xlvin BRITISH FOSSIL CORALS.

9. Genus HALOMITRA.
Dana, Zooph., p. 311, 1846.

Corallum composite, differmg from the preceding genus by its very long and numerous
septa.

Typ. sp., Halomitra pileus, Dana, loc. cit., p. 311; Fungia pileus, Lamarck, Hist. des An. sans Vert.,
t. ii, p. 237.

10. Genus Popopacta.
Milne Edw. and J. Haime, loc. cit., p. 71, 1849.

Corallum composite, cyathiform, and adherent by its basis. Calices as in Halomitra.

Typ. sp., Podobacia cyathoides, nob.; Agaricia cyathoides, Valenciennes MSS., in the Gallery of the
Paris Museum.

ll. Genus LitHactinia.
Lesson, Ilustr. Zool., 1833.

Corallum composite, free. Calices of one sort only, and not radiate. Septa short,
and separated by very thin, transverse laminz, which appear to be analogous to columelle.
Typ. sp., Lithactinia novehybernia, Lesson, loc. cit., vi, figs. 1, 2.

12. Genus PoLYPHYLLIA.
Quoy and Gaimard, Voy. de l’Astrolabe, Zooph., p. 184, 1833.

Corallum composite, free, and having calices of two sorts ; the central ones sub-radiate,
and arranged in a line.

Typ. sp., Polyphyllia pelvis, Quoy and Gaimard, loc. cit., pl. xx, figs. 8—10.

13. Genus ZooPri.us.
Dana, Zooph., p. 318, 1846.

Corallum composite. Septa of two sorts; the large ones radiately prolonged quite to
the margin ; the intermediate much smaller, and those only interrupted by the calicular
fossulee or oririms.

Typ. sp., Zoopilus echinatus, Dana, op. cit., p. 319.

Third Tribe— LOPHOSERIN A.

Plate (or basal wall) not perforate nor echinulate. No epitheca.

INTRODUCTION. xlix

14. Genus CYcLosERIs.
Milne Edw. and J. Haime, Comptes rend., t. xxix, p. 72, 1849.

Corallum simple, free, and discoidal. Seta very numerous, and united by their inner
edge. Wail completely horizontal.
Typ. sp., Cycloseris cyclolites, nob. ; Fungia cyclolites, Lamarck., Hist. des Anim. sans Vert.,t. ii, p. 236.

15. Genus DIAsERISs.
Milne Edw. and J. Haime, loc. cit., p. 72, 1849.

Corallum simple, free, and discoidal ; when young, composed of a certain number of
separate, radiating lobes, which, in the adult state, become cemented together. General
structure as in Cycloseris.

Typ. sp., Diaseris distorta, nob.; Fungia distorta, Michelin, in Guerin’s Mag. Zool., t. vy, Zooph.,
pl. v, 1843.

16. Genus TRocHosetRIs.
Milne Edw. and J. Haime, loc. cit., p. 72, 1849.

Corallum simple, trochoidal, adherent. Septa very numerous, and strongly granulated.
Typ. sp., Trochoseris distorta, nob.; Anthophyllum distortum, Michelin, Icon. Zooph., pl. xliii, fig. 8.

17. Genus CYATHOSERIS.
Milne Edw. and J. Haime, loc. cit., p. 72, 1849.

Corallum composite, trochoid, adherent. Ca/dices rather distinctly radiate. Seta long
and thick. Common basal walls, sometimes forming folds, which rise up so as to con-
stitute lobes or ridges on the upper surface of the corallum.

Typ. sp., Cyathoseris infundibuliformis, nob.; Agaricia infundibuliformis, Michelin, op. cit., tab.
xlii, fig. 12.

18. Genus LopHoseERis.
Milne Edw. and J. Haime, loc. cit., p. 72, 1849 ; Pavonia (ex parte), Lamarck, op. cit., t. ii, p. 238, 1816.

Corallum composite, foliaceous, and adherent, rising in the form of irregular criste or
of lobes, with confluent, radiate calicules on each side. Colwmella tubercular.

Typ. sp., Lophoseris boletiformis, nob. ; Pavonia boletiformis, Lamarck, loc. cit., p. 240.

19. Genus AGARICIA.
(Pars) Lamarck, Syst. des Anim. sans Vert., p. 375, 1801.

Corallum composite, foliaceous, and irregular. Calices arranged in concentric series,
separated by unequal ridges. Co/umella tubercular.
Typ. sp., Agaricia undata, Lamarck, loc. cit.; Madrepora undata, Solander and Ellis, Zooph., tab. xl.
7

—

BRITISH FOSSIL CORALS.

20. Genus PACHYSERIS.
Milne Edw. and J. Haime, loc. cit., p. 72, 1849.

Corallum similar to Agaricia, excepting that the corallites belonging to the same
trench are completely blended together. Co/wmedla well developed and dense.

Typ. sp., Pachyseris rugosa, nob. ; Agaricia rugosa, Lamarck, Hist. des An. sans Vert., t. i, p. 243.

21. Genus PHYLLASTREA.
Helioseris, Dana, Zooph., p. 269; Milne Edw. and J. Haime, loc. cit., p. 72, 1849.

Corallum composite, composed of frondiform expansions. Calices circumscribed, sub-
mammillate, and arranged around the parent corallite, which remains larger than the others.
Columella tubercular.

Typ. sp., Phyllastrea tubifex, Dana, loc. cit., tab. xvi, fig. 4.

22. Genus iLALOSERIS.
Milne Edw. and J. Haime, loc. cit., p. 72, 1849.

Corallum composite, forming foliaccous, crispate, lobulate expansions, the upper surface
of which is covered with very long radu, and shows only obsolete calices. Columella
rudimentary.

Typ. sp., Haloseris lactuca, Milne Edw. and J. Haime, loc. cit.

23. Genus LEPTOSERiS.
Milne Edw. and J. Haime, loc. cit., p. 72, 1849.

Corallum composite and adherent; the basal walls rising so as to constitute a sub-
crateriform disc, mm the centre of which is situated a large parent corallite, surrounded by
smaller ones. Cal’ces very imperfectly circumscribed, but well radiated. Septa very long.
Columella rudimentary.

Typ. sp., Leptoseris fragilis, Milne Edw. and J. Haime, loc. cit.

Sub-order 2.
ZOANTHARIA PERFORATA.

Corallum composed essentially of porous sclerenchyma; with the septal apparatus well
characterised, and consisting of six primitive elements, but beg sometimes represented

only by series of trabicule. Dissepiments rudimentary ; no tabule.

The principal character of this section of Zoantharia is furnished by the structure of

INTRODUCTION. hi

the sclerenchyma, which, instead of forming imperforated lamella as in the preceding
groups, is always porous, or even reticulate. In general the mural apparatus constitutes
here the greatest part of the corallum, and does not consist of costal lamine; the walls
are always perforated, and completely or nearly completely naked. It is also to be
remarked, that the visceral chamber is almost completely open from top to bottom, and
never filled up with dissepiments or synapticulz, as in most of the Zoantharia aporosa, or
with tabul, as will be seen in the next two sections of this order.

The perforated Zoantharia form three natural families: Hupsammide, Madreporide,
and Poritide.

Family V.
EUPSAMMID &.

Milne Edw. and J. Haime, Ann. des Sc. Nat., 3™° serie, vol. x, p. 65, 1848.

Corallum simple or complex, with well-developed lamellar septa, a spongiose columella,
and perforated, granular, subcostulated walls.

The septa are always numerous, and those of the last cyclum are never situated in the
direction of a line drawn from the centre of the calice to its circumference, but are bent
towards those of the penultimate cyclum, so as to produce the appearance of a six- or
twelve-branched star. The interseptal loculi are completely open from top to bottom, or
divided only by a few incomplete trabiculz. The walls have a granulate vermiculate surface,
and become often very thick in advanced age, but never constitute.a loose spongy mass,
as in Madreporide and Poritidz, or a compact ccenenchyma, as in Oculinide.

The star-hke arrangement of the septa, which is visible in transverse sections of these
corallums, as well as in the calice, is not met with in any other family. The principal
septa are sometimes imperforate, but those of the succeeding cycla are more or less porous.
It is also to be noted that there are never any pali, and that the coste are always rudi-
mentary ; sometimes there is a rudimentary epitheca.

1. Genus EvPSAMMIA.
Milne Edw. and J. Haime, Ann. Sc. Nat., 3° serie, vol. x, p. 77, 1848.

Corallum simple, subturbinate, free, and not presenting any lateral mural expansions.
Calice oval and rather deep. Septa broad, slightly exsert, granulate, closely set, and
forming four or five cycla. Coste simple, distinct from the basis of the corallum, nearly
equal, slightly vermiculate, and composed of a series of distinct, projecting granule.

Typ. sp., Eupsammia trochiformis, Milne Edw. and J. Haime, loc. cit., tab. i, fig. 3; Madrepora trochi-
Sormis, Pallas; Turbinolia elliptica, Brongniart.

In BRITISH FOSSIL CORALS.

2. Genus ENDOPACHYS.
(Pars) Lonsdale, Journ. of the Geol. Soc. of London, vol. i, p. 214, 1845.

Corallum simple, free, and organized as in the preceding genus, but much compressed
towards its basis, which is carinate, and continued laterally into two vertical lobiform or
cristate expansions. Calice arched ; fossula long and narrow.

Typ. sp., Endopachys Maclurii, Milne Edw. and J. Haime, Aun. Sc. Nat., vol. x, tab. i, fig. 1; Tur-
binolia Maclurw, Lea, Contrib. to Geol., tab. vi, fig. 206.

3. Genus BALANOPHYLLIA.
Searles Wood, Ann. and Mag. of Nat. Hist., vol. xiii, p. 11, 1844.

Corallum simple and adherent, sub-pediculate, or sub-cylindrical, with a very broad
basis. Columetla well developed, but not projecting at the bottom of the fossula. Sepia
thin, and closely set ; those of the last cyclum well developed, and complete in number.
Coste narrow, crowded, and nearly equal; no mural expansions.

Typ. sp., Balanophyllia caliculus, Searles Wood, loc. cit.

4. Genus HETEROPSAMMIA.
Milne Edw. and J. Haime, Ann. Sc. Nat., 3™° série, vol. x, p. 89, 1848.
Corallum simple, adherent, and growing by its basis so as to cover completely the
shell on which it is fixed, and to assume the appearance of being free. Calice smaller
than the basal part of the corallum. Colwmella well developed. Septa thick, slightly

exsert, and closely set. Walls not having distinct coste, but presenting small stric or
small papillze, composed of minute granule, and arranged in an irregular manner.

Typ. sp., Heteropsammia Michelinii, Milne Edw. and J. Haime, loc. cit., p. 89.

5. Genus LEPTOPSAMMIA.
Milne Edw. and J. Haime, loc. cit., p. 90, 1848.
Corallum simple, adherent. Ca/sce elliptical. Colwmella much developed, and
projecting at the bottom of the fossula. Septa neither exsert nor crowded, very thin, and

presenting scarcely any granulations ; those of the fifth order rudimentary. /Vad/s thin and
translucid. Coste distinct from the basis, and formed by series of small granule.

Typ. sp., Leptopsammia Stokesiana, Milne Edw. and J. Haime, loc. cit., tab. 1, fig. 4.

6. Genus ENDOPSAMMIA.
Milne Edw. and J. Haime, loc. cit., p. 91, 1848.

Corallum simple, erect, and adherent. Calice circular. Colwmella much developed,
but not projecting. Septa thick, strongly granulated, and slighty exsert, forming four
cycla, the last of which is almost rudimentary. Wadl/s covered with an indistinct pellicular
epitheca, and having broad, straight coste.

Typ. sp., Endopsammia Philippensis, Milne Edw. and J. Haime, loc. cit., tab. i, fig. 5.

INTRODUCTION. Hu

7. Genus STEPHANOPHYLLIA.
Michelin, Dict. des Sc. Nat., Suppl., vol. i, p. 484, 1841.

Corallum simple, free, and presenting no trace of adherence. /Vail discoidal,
horizontal. Calice circular and open. Sepéa tall, thin, crowded, not projecting laterally
beyond the edge of the mural disc, covered with conical granulations on each side, and
all, excepting those of the first cyclum, united by the inner edge. Cos? delicate, straight,
composed of simple series of obscure granulations, and radiating regularly from the
centre of the mural disc to its circumference. No epitheca.

Typ. sp., Stephanophyllia elcgans, Michelin, Icon. Zooph., pl. viii, fig. 2. Milne Edw. and J. Haime,
loc. cit., tab. i, fig. 10.

8. Genus DENDROPHYLLIA.
Blainville, Dict. des Sc. Nat., vol. lx, p. 319, 1830.

Corallum composite, and in general arborescent. Corallites cylindrical, or cylindrico-
turbinate, and formed by lateral gemmation. C/ices circular, or nearly so ; fossula dec.
Columella well developed, and in general projecting much at the bottom of the fossula.
Septa not exsert, thin, and closely set ; those of the fourth cyclum well developed. a//s
becoming very thick, and presenting narrow vermiculate coste, formed by series of granulix.

Typ. sp., Dendrophyllia ramea, Blainville, loc. cit.; Milne Edw., Atlas du Régne Animal de Cuvier.
Zooph., pl. lxxxiii, fig. 1.

9. Genus LOBOPSAMMIA.
Milne Edw. and J. Haime, Ann. des Sc. Nat., 3™° serie, vol. x, p. 105, 1848.
Corallum composite, arborescent, increasing by successive fissiparity. Ca/ices irregwiar
inform. Septa formimg four complete and well-developed cycla. In other respects similar
to Dendrophyllia.

Typ. sp., Lobopsammia cariosa, Milne Edw. and J. Haime, loc. cit.; Zithodendron cariosum, Goldfuss,
Petref. Germ., vol. i, tab. xiii, fig. 7.

10. Genus C@NoPsAMMIA.

Milne Edw. and J. Haime, loc. cit., p. 106, 1848; Tubastrea, Lesson, Voyage aux Indes orient. par
Belanger, 1834.

Corallum composite, dendroid, or sub-globose, increasing by lateral or sub-basal gemia-
tion. Corallites cylindrical. Calices circular, or nearly so. Columel/a tubercular, well
developed. Seta not exsert, distant, and forming three cycla; those of the fifth order
rudimentary. Coste narrow, sub-vermiculate towards the bases, simple, and formed of «
series of granule near the calice.

Typ. sp., Cenopsammia coccinea, Milne Edw. and J. Haime, loc. cit., p. 107 ; Tubastrea coccinea, Lesson.
op. cit., Zooph., tab. 1; Astrea calicularis, Blainville, Manuel d’Actinol., tab. liv, fig. 2

ol

11. Genus STEREOPSAMMIA.

Corallum presenting most of the characters of Ceenopsammia, but not having any
Columella.

Typ. sp., Stereopsammia humilis, nob., tab. v, fig. 4.

iw BRITISH FOSSIL CORALS.

Family VI.
MADREPORID/.

Corallum composite, increasing by gemmation and not by fissiparity. Cenenchyma

abundant, spongy, and reticulate. Walls very porous, and not distinct from the coenenchyma.
Senta lamellose, and well developed ; loculi free.

First Tribe—MADREPORIN A.

Visceral chambers divided into two equal parts by two of the principal septa, which
ave more developed than the others, and meet by their inner edge.

1. Genus MADREPORA.
Lamarck, Hist. des Anim. sans Vert., t. xi, p. 277, 1816.

Corallum composite, forming ramified, lobate, or fasciculate masses. Cenenchyma
loose, and delicately echinulate. Ca/ices projecting, with a thick margin. No columella.

Typ. sp., Madrepora muricata, Ellis and Solander, Zooph., tab. lvii; Madrepora abrotanoides,
Lamarck, loc. cit., p. 280.

Second Tribe—EX PLANARIN A.

Visceral chamber presenting at least six equally developed principal septa.
2. Genus EXPLANARIA.

(Pars) Lamarck, Hist. des Anim. sans Vert., vol. ii, p. 254, 1816 ; Gemmipora, Blainville,
Dict. des Sc. Nat., vol. Ix, p. 352, 1830.

Corallum in general foliaceous. Ccenenchyma abundant, rather dense, and delicately
echinulate. Sepa almost all of the same size. Co/wmella spongy.

Typ. sp., Hxplanaria crater, nob.; Madrepora crater, Pallas, Eleuch. Zooph., p. 332.

3. Genus ASTREOPORA.
Blainville, Dict. des Sc. Nat., vol. lx, p. 348, 1830.

Corallum massive. Canenchyma of a loose texture, and strongly echinulated. Septa
unequally developed. No columella.

Typ. sp., Astreopora myriophthalma, Blainville, loc. cit. ; Astrea myriophthalma, Lamarck, op. cit.,
p. 260.

INTRODUCTION. ly

Family VII.

PORITID/.
Corallum entirely composed of reticulate sclerenchyma. Septal apparatus well
developed, but never lamellar, and composed only of series of styliform processes or
trabicule, constituting by their junction a sort of irregular trellis. /a//s presenting the

same structure, and “not distinct from the ccenenchyma. Visceral chamber contammng
some small dissepiments, but never divided by tabule.

First Tribe—PORITIN &.

Coenenchyma rudimentary. or not existing.

1. Genus PoritTEs.

(Pars) Lamarck, Hist. des An. sans Vert., t. i, p. 267, 1816; Milne Edw. and J. Haime, Comptes rend.,
t. xxix, p. 258, 1849.

Corallum composed of sclerenchyma, very wregularly reticulated. Ca/ies shallow.
Septa sot numerous, rudimentary, and appearing to be represented bv a circle of poli, the

apex of which is papillose.
Typ. sp., Porites conglomerata, Lamarck, loc. cit., p. 269.

2. Genus LITHAREA.
Milne Edw. and J. Haime, Comptes rend., t. xxix, p. 258, 1844.

Sclerenchyma very irregularly reticulated. Ca/ices not very deep. Colwuella spongy.
Pali rudimentary, or not existing. Sepia well developed, particularly towards the wall.

Typ. sp., Litharea Websteri, nob. ; Astrea Websteri, Bowerbank, Mag. of Nat. Mist., new scries,
vol. iv, p. 27, figs. A, B, 1540.
3. Genus COSCINAR EA.
Milne Edw. and J. Haime, Comptes rend. de l’Ac. des Se., t. xxvii, p. 496, JS4s.

Corallum of a dense structure. Ca/ices vather deep ; neither pali, nor distinct walls.
Septa crowded, very regularly fenestrate, and with crispate edges, passing without any
interruption from one visceral chamber to the adjacent one. No epithevn.

iy

Typ. sp., Coscinarea Botte, Milne Edw. and J, Haime, Ann. des Se. Nat., 3° serie, vol. ix, tab. y, fig.

fyi BRITISH FOSSIL CORALS.

4.. Genus MicROSOLENA.
Lamouroux, Exp. méth., p. 65, 1821.

_Corallum differmg from the preceding genus by the structure of the septa, the perfo-
rations of which are much larger than in Coscinarea, and by the existence of a strong,
common. epitheca.

Typ. sp., Microsolena porosa, Lamouroux, op. cit., tab. Ixxiv, fig. 24.

5. Genus GONIOPORA.

Quoy and Gaimard, Voy. de l’Astr., Zooph., p. 218, 1833 ; Goniopora and Porastrea, Milne Edw. and
J. Haime, Comptes rend. de l’Acad. des Sc., t. xxvii, p. 496, 1848.

Corallum having distinct, elevated walls, of a fenestrate structure. Calices deep.
Columella spongy. Septa well developed, and fenestrate. No pali.

Typ. sp., Goniopora pedunculata, Quoy and Gaimard, Voyage de l’Astrolabe, Zooph., tab. xvi,
figs. 9-1].

6. Genus RHODARAA.
Milne Edw. and J. Haime, Comptes rend., t. xxix, p. 259, 1849.

Corallum with thick walls, rather high. Septa rudimentary. Pali greatly developed,
and forming a rosette in the centre of the calice.
Typ. sp., Rhodarea calicularis, nob.; Astrea calicularis, Lamarck, Hist. des An. sans Vert., t. ii,

p- 266.

7. Genus PORARAA.
Milne Edw. and J. Haime, Comptes rend., t. xxix, p. 259, 1849.

Walls thm, and widely fenestrated. Septa formed by a series of spiniform processes,
which sometimes ramify towards the centre of the visceral cavity, so as to constitute a sort
of spurious columella.

Typ. sp., Porarea fenestrata, nob.; Pocillopora fenestrata, Lamarck, Hist. des An. sans Vert., t. ii,
p. 275 ; Milne Edw. and J. Haime, Ann. des Sc. Nat., 3™° serie, vol. ix, fig. 1.

S. Genus Hoare.
Milne Edw. and J. Haime, Comptes rend., t. xxix, p. 259, 1849.

Calices with distinct polygonal margins, rather deep. Septal apparatus composed of
ivegular trabicula, completely blended with the walls, and constituting thus a delicate
spongy mass. Co/umella fasciculate and short.

Typ. sp., Holarea Parisiensis, nob. ; Alveolites Parisiensis, Michelin, Icon. Zooph., pl. xlv, fig. 10.

INTRODUCTION. lv

Second Tribe—MONTIPORIN &.

Cenenchyma abundant and spongy.

9. Genus ALVEOPORA.
Quoy and Gaimard, Voyage de l’Astrolabe, Zooph., p. 240, 1833.

Corallum arborescent. Cxenenchyma very porous and echinulate, but not bearing large
excrescences. Margins of the calices scarcely distinct. Septa not numerous, and formed
by series of spimiform processes. No columella.

Typ. sp., Alveopora rubra, Quoy and Gaim., loc. cit., Zooph., tab. xix, figs. 11-14.

10. Genus Montirora.
Quoy and Gaimard, op. cit., p. 247, 1833; Manopora, Dana, Zooph., p. 489, 1846.

Corallum of various forms, differmg from Alveopora by the existence of large projec-
tions of the canenchyma between the calicules. Cenenchyma much more abundant, and
more delicately spongy.

Typ. sp., Montipora verrucosa, Quoy and Gaim., op. cit., Zooph., pl. xx, fig. 11.

ll. Genus PSAMMOCORA.
Dana, Zooph., p. 344, 1846.

Ceenenchyma somewhat compact, of a fasciculate structure, and having its surface
papillose. Ca/ices very shallow, confluent, and without distinct walls. Septa thick, and
formed by strong spiniform processes.

Typ. sp., Psammocora obtusata, Dana, loc. cit., p. 345; Pavonia obtusangula, Lamarck, Hist. des
An. sans Vert., t. ii, p. 240.

Sub-order 3.
ZOANTHARIA TABULATA.

Corailum essentially composed of a well-developed mural system, and having the visceral
chambers divided into a series of stories by complete transverse tabula or diaphragms.
Septal apparatus rudimentary.

The principal character of this sub-order is founded on the existence of the lamellar
diaphragms that close the visceral chamber of the corallites at different heights, and differ
from the dissepiments of the Astreidee by not bemg dependent on the septa, and forming
as many complete horizontal divisions extending from side to side of the general cavity,

5

lviii BRITISH FOSSIL CORALS.

instead of occupying only the one or two loculi. It is also to be remembered that the
septal apparatus, although more or less rudimentary, has the same general mode of
arrangement as in the preceding sub-orders, and never presents the crucial character which
we shall find in Zoantharia rugosa.

This section comprises four families: Favositide, Milleporide, Seriatoporide, and
'Thecidee.

Family VIII.
MILLEPORID Ai.

Corallum principally composed of a very abundant ccenenchyma, distinct from the
walls of the corallites, and of a tubular or cellular structure. Sepfa not numerous ;
tabulee numerous, and well formed.

1. Genus MILLEPORA.

(Pars) Lamarck, Syst. des An. sans Vert., p. 373, 1801; Palmipora, Blainville, Dict. des Sc. Nat., t. lx,
p- 356, 1830.

Corallum of various forms, but more or less foliaceous. Cenenchyma extremely
abundant, of an irregular subtubular structure. Ca/ices of very different dimensions in
the same corallum. No distinct septa. Zudule horizontal.

Typ. sp., Millepora alcicornis, Lamarck, loc. cit.; Milne Edw., Atlas du Regne Anim. de Cuvier,
Zooph., tab. Ixxxix, fig. 1.

2. Genus HELIOPORA.
(Pars) Blainville, Dict. des Sc. Nat., vol. Ix, p. 357, 1830; Dana, Zooph., p. 539, 1846.

Corallum lobulate, somewhat massive, and differing from Miullepora by the regular
tubular structure of the ccenenchyma, and the existence of small but distinct septa.

Typ. sp., Heliopora cerulea, Blainville, loc. cit., p. 357.

3. Genus HELIOLITES.

Dana, Zooph., p. 541, 1846; Paleopora, M’Coy, Ann. and Mag. of Nat. Hist., 2d series, vol. ii, p. 129,
1849; Geoporites, D’Orbigny, Prodr. de Palzeoni. stratif. Univers., t. i, p. 49, 1849.

Corallum sub-globose. Canenchyma regularly tubular. Septal radu advancing
almost to the centre of the visceral chamber on the upper surface of the tabula, which
are horizontal.

Typ. sp., Heliolites pyriformis, Dana, loc. cit., p. 542; Heliolite pyriforme, etc., Gucitard, Mem. sur
les Sc. et les Arts, vol. iii, pl. xxii, figs. 13, 14.

INTRODUCTION. hix

4. Genus FisTULIPORA.
M’Coy, Ann. and Mag. of Nat. Hist., 2d series, vol. iii, p. 130, 1849.

Corallum with vesicular coenenchyma ; thick walls and mfundibuliform tabule.

Typ. sp., Fistulipora minor, M’Coy, loc. cit., figs. a, 6.

5. Genus PIASMOPORA.
Milne Edw. and J. Haime, Comptes rend., t. xxix, p. 262, 1849.

Corallum free, sub-hemispheric, with a basal plate covered with an epitheca presenting
concentric folds. Calices immersed. Septa rudimentary. Zadbule horwontal. Walls
thin. Canenchyma composed of large, vertical, radiate lamine, united by smaller hori-
zontal plates, and resembling much the costal ccenenchyma of the Astreide.

Typ. sp., Plasmopora petaliformis, nob.; Porites petaliformis, Lonsdale, in Murchison, Sil. Syst.,
pl. xvi, fig. 4.

6. Genus PRopora.
Milne Edw. and J. Haime, Comptes rend., t. xxix, p. 262, 1849.
Corallum differmg from the preceding genus by the calices having exsert margins ; the
septa being more developed, and extending outwards so as to constitute small coste.

Typ. sp., Propora tubulata, nob.; Porites tubulata, Lonsdale, Sil. Syst., pl. xvi, figs. 3, 3°, 3° (ceeteris

exclusis).

7. Genus AXOPORA.

Corallum composite, incrusting, and forming thin expansions, which are often super-
posed. Canenchyma abundant, and forming wregular ridges between the calices, which
are small and deep. Septa rudimentary. Colwmella well developed, fasciculate, and
expanding at its passage through each of the tabulz.

Typ. sp., Avopora pyriformis, nob. ; Geodia pyriformis, Michelin, Icon., tab. xlvi, fig. 2.

8. Genus LOBOPORA.

Corallum having the same structure as in the preceding genus, but forming large,
thick, foliaceous expansions, the two surfaces of which are covered with calices.

Typ. sp., Lobopora Solanderi, nob. ; Palmipora Solanderi, Michelin, op. cit., tab. xlv, fig. 9.

Ix BRITISH FOSSIL CORALS.

amily IX.
FAVOSITID.

Corallum essentially formed by lamellar walls, with little or no ccenenchyma. Visceral
chambers divided by numerous and well-developed complete tabule.

First Tribe—FAVOSITID A.

Corallum massive. Walls perforated. Septarudimentary. No ccenenchyma.

1. Genus Favosttes.

Lamarck, Hist. des An. sans Vert., vol. ii, p. 204, 1816; Calamopora, Goldfuss, Petref. Germ.,
vol. i, p. 77, 1826-30.

Corallum composed of basaltiform corallites, and having a basal plate covered with an
epitheca, and no radiciform appendices. Ca/ices at right angle with the axis of the corallite,
and in general hexagonal. /Va//s perforated in a very regular manner. Zadule horizontal,
and very regularly superposed. No canenchyma.

Typ. sp., Favosites Gothlandica, Lamarck, loc. cit., p. 206.

2. Genus MICHELINIA.
De Koninck, Descr. des Anim. foss. des Terr. houilliers de la Belgique, p. 30, 1842-44.

Corallum having a basal plate with radiciform prolongations. Zadule very irregular,
and subvesicular. ‘The other characters as in Favosites.

Typ. sp., Michelinia tenuisepta, De Koninck, loc. cit., pl. c, fig. 3 a, 6.

3. Genus KCNINCKIA.
Milne Edw. and J. Haime, Comptes rend., t. xxix, p. 260, 1849.

Corallum resembling Favosites, but having the walls larger and less regular, and the
septa constituted by series of distinct and spiniform processes, interrupted at certain
distances by the tabulz, which are horizontal.

Typ. sp., Koninckia fragilis, Milne Edw. and J. Haime, loc. cit.

4. Genus ALVEOLITES.
(Pars) Lamarck, Syst. des An. sans Vert., p. 375, 1801 ; Steininger, Mem. Soc. Geol. France, vol. i.

Corallum composed of superposed strata of corallites very similar to those of Favosites,

INTRODUCTION. Ix

but much shorter, and terminated by an oblique semicircular or subtriangular ealice. the
edge of which projects on one side.

Typ. sp., Alveolites spongites, Steininger, Mem. de la Soc. Geol. de France, vol. i; Calamopora
spongites, Goldfuss, Petref. Germ., pl. xxviii, figs. 1%, 1’, 1°.

Second Tribe—CH ASTETIN AL.

Corallum massive. /Va/ls not perforated. Neither septa nor coenenchyma.

5. Genus CHETETES.
Fischer, Oryct. du Gouy. de Moscou, p. 159, 1837.

Corallum glomerate. Cora/lites very long, basaltiform, and in general more or less
bent. Calices polygonal. Zabule independent, not connected in the adjoiing corallites,
nor placed on the same level throughout the corallum.

Typ. sp., Chetetes radians, Fischer, loc. cit., pl. xxxvi, fig. 6.

6. Genus Danta.
Milne Edw. and J. Haime, Comptes rend., t. xxix, p. 261, 1849.

Corallum having most of the characters of Cheetetes, but with the tabula connected
through the different corallites so as to constitute a series of common plates, and to divide
the whole mass into a great number of parallel strata.

Typ. sp., Dania Huronica, Milne Edw. and J. Haime, loe. cit.

7. Genus STENOPORA.
(Pars) Lonsdale, Geol. of Russia and Ural Mount., vol. i, p. 631, 1845.

Corallum very similar to Cheetetes, but having small styhform processes at the angles
of the calices.

Typ. sp., Stenopora spinigera, Lonsdale, loc. cit., pl. a, fig. 2.

8. Genus CoNSTELLARIA.

Dana, Zooph., p. 537, 1846.

Third Tribe—HALYSITIN A.

Corallum composed of corallites constituting vertical laminz or fasciculi, but more or
less free laterally, and united by means of connecting tubes or mural expansions. Valls
wel] developed, and not porous. Sepa distinct, but small.

Ix BRITISH FOSSIL CORALS.

8. Genus Hanysirns.

Fischer, Zoognosia, 3d edit., vol. i, p. 387, 1813; Catenipora, Lamarck, Hist. des An. sans Vert., t. ii,
p. 206, 1816.

Corallites extremely long, arranged in a single. series, and united laterally, so as to
constitute large flabelliform expansions, which remain free laterally, but often meet, and
thus form a lacunous mass. Kipitheca very thick. Septa almost rudimentary, but very
distinct in perfect specimens. Zudu/@ horizontal.

Typ. sp., Halysites escharoides, Fischer ; Catenipora escharoides, Lamarck, loc. cit., p. 207.

9. Genus Haritopitns.
Fischer. Notice sur les Tubipores fossiles, 1828 ; Syringopora, Coldfuss, Petref. Germ., vol.i, p. 75, 1826-33.

Corallum fasciculate. Corallites irregularly cylindrical, very long, and united by hori-
zontal connecting tubes. Zadbule infundibuliform.

Typ. sp., Harmodites ramulosa, nob. ; Syringopora ramulosa, Goldfuss, loc. cit., pl. xxv, fig. 7.

10. Genus THEcostrcitrs.
Milne Edw. and J. Haime, Comptes rend., t. xxix, p. 261, 1849.

Corallites cylmdrical, short, and united by strong mural expansions situated at various
heights. Zadule horwontal.

Typ. sp., Thecostegites Bouchardi, nob.; Harmodites Bouchardi, Michelin, Icon. Zooph., pl. xlvii, fig. 3.

Fourth Tribe—POCILLOPORIN A.

Corallum massive, gibbous, or subdendroid, with thick imperforated walls, forming,
towards the surface, an abundant compact cocnenchyma. Sep/a quite rudimentary.

11. Genus PoctuLopora.
(Pars) Lamarck, Hist. des An. sans Vert., t. ii, p. 273, 1816; Dana, Zouph., p. 523, 1846.

Calices shallow, and presenting, at their bottom, a transverse, thick, projecting ring,
resembling a columella.

Typ. sp., Pocillopora acuta, Lamarck, loc. cit., p. 274; Mine Edw., Atlas du Régne Animal de Cuvier,
Zooph., pl. Ixxxi, fig. 3

INTRODUCTION. Ix

Family X.
SERIATOPORID#.

Corallum arborescent or bushy, with an abundant compact coenenchyma. Visceral
chambers filling up by the growth of the columella and the walls, and showing but few
traces of tabule.

1. Genus SERIATOPORA.
Lamarck, Hist. des An. sans Vert., vol. ii, p. 282, 1816.

Corallum arborescent, with echinulated branches. Cadices arranged in ascending series.
Septa scarcely visible. Columella large and compact.

Typ. sp., Seriatopora subulata, Lamarck, loc. cit., p. 282.

2. Genus DENDROPORA.
Michelin, Icon. Zooph., p. 187, 1845.

Corallum arborescent, with very delicate smooth branches. Ca/ices distant, and sur-
rounded by a narrow, obtuse margin. Septa small, but distinct.
Typ. sp., Dendropora explicita, Michelin, op. cit., pl. xlviui, fig. 6.

3. Genus RHABDOPORA.
Milne Edw. and J. Haime, Comptes rend., t. xxix, p. 262, 1849.

Corallum with prismatic echinulate branches. Ca/ices arranged in series. Septa very
distinct.

Typ. sp., Rhabdopora megastoma; Dendropora megastoma, M‘Coy, Ann. and Mag. of Nat. Hist.,
2d series, vol. ili, p. 129.

Family XI.
THECID A.

Corallum massive, with an abundant, compact, spurious ccenenchyma, produced by the
septa becoming cemented together laterally. Zadu/e numerous.

Genus 'THECIA.
Milne Edw. and J. Haime, Comptes rend., t. xxix, p. 263, 1849.

Septal system highly developed. Cadices shallow, with a very small deep fossula.

Typ. sp., Thecia Swinderniana ; Agaricia Swinderniana, Goldfuss, Petref. Germ., pl. xxxviil, fig. 3 ;
Porites expatiata, Lonsdale, ap. Murchison, Sil. Syst., p. 678, tab. xv, fig. 3.

Ixus BRITISH FOSSIL CORALS.

Sub-order 4.
ZOANTHARIA RUGOSA.

Corallum simple or composite, with a septal apparatus never forming six distinct
systems, as in all the preceding Zoantharia, but appearing to be derived from four primary
elements. Sometimes this disposition is rendered manifest by the existence of four well-
characterised primary septa, or of an equal number of depressions occupying the bottom of
the calice, aiid assuming a crucial appearance: in other cases only one of these primary
septa or excavations is well developed so as to interrupt the radiate form of the system ; and
in others, again, no trace of septal groups can be discovered, and the whole apparatus is
represented by numerous equally developed radiate strize rising on the surface of the tabula,
and extending up the inner side of the walls. The corallites are always perfectly distinct,
and are never united by means of a coenenchyma; nor do they ever form linear series,
which is often the case in the preceding sections. They multiply by gemmation, and the
reproductive buds are in general developed on the surface of the calices of the parents :
this often arrests the growth of the latter, and gives rise to a superposition of generations.
It is also to be noted that the septa, although in general very incomplete, are never porous,
and never bear synapticule, but that the visceral chamber is in general filled up from the
bottom by a series of transverse tabul, or by a vesicular structure, which often constitutes
the principal part of the corallum.

Family XII.
STAURIDA.

Corallum with well-developed septa, extending without any interruption from the bottom
to the top of the visceral chamber, united by lamellar dissepiments, and arranged in four
systems, characterised by 2n equal number of large primary septa.

1. Genus STAURIA.

Corallum composite, massive, astreiform, and increasing by calicular gemmation.
Corallites united by their walls, or free in part, and not presenting any coste. Septa large,
and with undivided edges, united along the axis of the visceral chamber. No columella.

Typ. sp., Stauria astreiformis, nob.

2. Genus Howiocystis.

Lonsdale, in the Quarterly Journal of the Geol. Soc. of London, vol. v, part i, p. 83, 1849.

Corallum composite, massive, astreiform, and increasing by extra-calicular gemmation.
Corallites united by means of well-developed coste. Columella styliform.

Typ. sp., Holocystis elegans ; Cyathophora elegans, Lonsdale, loc. cit., tab. iv, figs. 12, 13, 14, 15.

INTRODUCTION. Ixv

Family XIII.
CYATHAXONID/.

Corallum with well-developed, complete septa, which extend without interruption from
the bottom to the top of the visceral chamber, and not forming a regular radiate circle ;
those of the primary cyclum not much larger than the others, and not forming a four-
branched cross, as in the Stauride ; one well-characterised septal fossula. No dissepiments
nor tabule.

Genus CYATHAXONIA.
Michelin, Icon. Zooph., p. 258, 1846.

Corallum simple. Calice deep. Columeila styliform, strong, and very prominent.
Septa extending to the columella; the place of one of them occupied by a deep depression
or septal fossula.

Typ. sp., Cyathazxonia cornu, Michelin, loc. cit., p. 258, pl. lix, fig. 9.

Family XIV.
CYATHOPHYLLID.

Corallum with incomplete septa, that do not extend from the bottom to the top of the
visceral chamber, in the form of uninterrupted laminz ; those of the primary cyclum similar
to the others, and not forming a central four-branched cross. Septal fossulee varying in
number and in size. Visceral chamber divided by a series of superposed tabule.

First Tribe—ZAPHRENTIN A.

A single septal fossula, well developed, or replaced by a sulcus or a crestiform process,
and occasioning more or less irregularity in the radiate arrangement of the septal, apparatus.
The corallum is simple, and free in all the known species.

1. Genus ZAPHRENTIS.

Rafinesque and Clifford, Ann. des Sciences physiques de Bruxelles, vol. v, p. 234, 1820; Caninia,
Michelin, Dict. des Sc. Nat., Supplém., vol. i, p. 485; Siphonophyllia, Scouler, in M‘Coy’s Carbonif. Foss.
of Ireland, p. 187, 1844.

Corallum simple and trochoid. Calice deep. Septal fossula strongly developed, and
occupying the place of one of the septa. No columella. Tabule moderately developed,
and bearing on their upper surface a series of septa, which extend from the wall to the
centre of the visceral chamber, and are denticulate all along their calicular edge.

Typ. sp., Zaphrentis patula; Caninia patula, Michelin, Icon. Zooph., tab. lix, fig. 4.

Ixvl BRITISH FOSSIL CORALS.

2. Genus AMPLEXUS.

Sowerby, Miner. Conchol., vol.i, p. 165; Ampleaus and Cyathopsis, D’Orbigny, Prodrome de Paléontol.,
vol. i, p. 105, 1850.

Corallum resembling Zaphrentis, excepting that the septa do not extend to the centre
of the visceral chamber, and leave the upper surface of the tabule naked and smooth in
that part. The septal fossula well characterised in the upper portion of the corallum, but
not so on the lower floors. Zadule highly developed.

Typ. sp., Amplexus coralloides, Sowerby, loc. cit., tab. 1xxii.

3. Genus MENOPHYLLUM.
Corallum resembling Zaphrentis, excepting that a small septal fossula is situated on
each side of the large one, and that one half of the central part of the calice is occupied by
an elevated, smooth portion of the tabula, which resembles a crescent.

Typ. sp., Menophyllum tenui-marginatum, nob.

4. Genus LOPHOPHYLLUM.

Corallum resembling Zaphrentis, excepting that a crestiform co/wmella occupies the
centre of the calice, and is in continuity by one of its ends with a small septum, placed in
the middle of the septal fossula, and by the other end with the opposite primary septum.

Typ. sp., Lophophyllum Konincki, nob.

5. Genus ANISOPHYLLUM.

Corallum resembling Zaphrentis, excepting by the great development of three primary
septa, one of which is placed facing the septal fossula; this fossw/a extending much towards
the centre of the visceral chamber, and ceasing there to be distinct from the bottom of the
calice.

Typ. sp., Anisophyllum Agassizi, nob.

6. Genus BARYPHYLLUM.

Corallum very short. Calice quite superficial. A slightly developed septal fossule,
corresponding to one of the branches of a cross, the three other branches of which are
constituted by well-developed primary septa. The younger septa not arranged in a regular
radiate circle, but inclined obliquely towards the primary ones.

Typ. sp., Baryphyllum Verneuilanum, nob.

INTRODUCTION. Ixvii

7. Genus HAuutia.

Corallum tall, turbinate. Septa highly developed, and extending to the centre of the
tabule. No columella. One remarkably large primary septum occupying the place of the
septal fossula, and the neighbouring septa directed towards it, so as to assume a pinnate
arrangement ; the septa belonging to the two other systems presenting the usual regular
radiate position.

Typ. sp., Hallia insignis, nob.

8. Genus AULACOPHYLLUM.

Corallum resembling Hallia by the mode of arrangement of the septa, but having the
septal fossula not replaced by a primary septum, and affecting the form of a narrow groove,
at the bottom of which the septa of the two adjomimg systems meet, and even cross each
other.

Typ. sp., dulacophyllum suleatum ; Caninia sulcata, D’Orbigny, Prod. de Paléont., vol. i, p. 105.

9. Genus 'TROCHOPHYLLUM.

Corallum simple, trochoid. Calice rather shallow. Septal fossula rudimentary, and
occupied by a small septum. The other sepfa thick, not denticulate, presenting a regular
radiate mode of arrangement, and extending almost to the centre of the visceral chamber,
where a small tabula is visible.

Typ. sp., Trochophyllum Verneuili, nob.

10. Genus HADROPHYLLUM.

Corallum short. Ca/ice superficial. One very large septal fossula, and three small
ones, representing a cross. ‘The radiate arrangement of the septa somewhat irregular.

Typ. sp., Hadrophyllum Orbiynyi, nob.

11. Genus COMBOPHYLLUM.

Corallum presenting the general form of a Cyclolites. A single septal fossula. Seta
exsert, and regularly radiate.

Typ. sp.,Combophyllum osismorum, nob.

Second Tribe—CYATHOPHYLLIN &.

Septal apparatus regularly radiate, and uninterrupted, or equally divided mto four
groups by four superficial septal fossula. No true colwmella, but sometimes a spurious
one formed by the inner edge of the septa.

Ixviil BRITISH FOSSIL CORALS.

12. Genus CYATHOPHYLLUM.
Goldfuss (in parte), Petref. Germ., vol. i, p. 54, 1826.

Corallum simple or composite. No coste. Septa well developed, extending to the
centre of the calice, and twisted together so as to produce the appearance of a small colu-
mella. Zubule occupying only the centre of the visceral chamber; the outer portion of
which is filled up with numerous vesicular dissepiments. A single wall, situated exteriorly,
and provided with a complete epitheca.

Typ. sp., Cyathophyllum helianthoides, Goldfuss, loc. cit., tab. xx, fig. 2.

13. Genus PACHYPHYLLUM.

Corallum composite, and increasing by lateral gemmation. Cora/iites united in their
lower portion by means of the great development of the costae and the exotheca, and not
delimitated by an individual epitheca. Zudule well characterised.

Typ. sp., Pachyphyllum Bouchardi, nob.

14. Genus CAMPOPHYLLUM.

Corallum simple, very tall, and protected by an epitheca. Septa well developed.

Tabule very large, and smooth towards the centre. Interseptal loculi filled with small
vesiculze.

Typ. sp., Campophyllum flecuosum ; Cyathophyllum flezuosum, Goldfuss, Petref. Germ., vol. i, tab. xvii,
fig. 3.

15. Genus STREPTELASMA.

Hall, Palzeont. of New York, p. 17, 1847.

Corallum simple, and differing from Cyathophyllum by the structure of the wall, which
is destitute of epitheca, and covered with sublamellar costz.

Typ. sp., Streptelasma corniculum, Hall, loc. cit., tab. xxv, fig. 1.

16. Genus OMPHYMA.
Rafinesque and Clifford, in Ann. des Sc. Phys. de Bruxelles, vol. v, p. 234, 1820.

Corallum simple, turbinate. Wall provided with a rudimentary epitheca, and pro-
ducing radiciform appendices. Septa very numerous, equally developed, and divided into

four groups by an equal number of shaliow septal fossule. Zabule well developed, and
smooth towards the centre.

Typ. sp., Omphyma turbinata ; Madrepora turbinata, Lin, Ameen. Acad., vol. i, tab. iv, fig. 2.

INTRODUCTION. Ixix

17. Genus GoNIOPHYLLUM.

Corallum simple, and affecting the form of a quadrangular pyramid. Calice deep and
square. Septa thick and well developed. TZaédule central, and but little developed.

Typ. sp., Goniophyllum pyramidale ; Turbinolia pyramidalis, Hisinger, Lethzea Suecica, tab. xviii, fig. 12.

18. Genus CHONOPHYLLUM.

Corallum simple, and constituted principally by a series of infundibuliform fabule, super-
posed and invaginated, the surface of which presents numerous septal radii equally deve-
loped, and extending from the centre to the circumference. No columella nor walls.

Typ. sp., Chonophyllum perfoliatum ; Cyathophyllum perfoliatum, Goldfuss, tab. xviii, fig. 5.

19. Genus PrYcHOPHYLLUM.
Strombodes (pars), Lonsdale, Sil. Syst., p. 691, 1839 (not Schweigger.).

Corallum simple, and organized as in the preceding genus, but having the septal radii
strongly twisted towards the centre of the tabulz, so as to constitute a spurious columella.

Typ. sp., Ptychophyllum Stokesi, nob.; C. Stokes, Trans. of the Geol. Soc., 2d series, vol. i, tab. xxix,
fig. 1. (‘.B. The second figure bearing this number, but not the first.)

20. Genus HELIOPHYLLUM.
Hall, in Dana, Zooph., p. 396. 1846.

Corallum simple. Septal apparatus well developed, and producing lateral lamellar
prolongations, which extend from the wall towards the centre of the visceral chamber,
so as to represent ascending arches and to constitute irregular central tadu/e, and which
are united towards the circumference by means of vertical dissepiments.

Typ. sp., Heliophyllum Halli, nob.; Strombodes heliunthoides, Hall, Geol. of New York, No. 48, fig. 3
(not S. helianthoides of Phillips).

21. Genus METRIOPHYLLUM.

Corallum simple, turbinate. Sepfa well developed, slightly twisted, and extending to
the centre of the visceral chamber, through well-developed tadule.

Typ. sp., Metriophylium Bouchardi, nob. ; Cyathophyllum mitratum, Michelin, Icon. Zooph., tab. xlvii,
fig. 7 (not C. mitratum of Schlotheim).

Ixx BRITISH FOSSIL CORALS.

22. Genus CLISIOPHYLLUM.
(Pars) Dana, Exploring Exped., Zoophytes, p. 361, 1846.

Corallum simple, turbinate. Sepa well developed, and rising towards the centre of
the calice so as to form a spurious columella, but not twisted.

Typ. sp., Clisiophyllum Danianum, nob.

23. Genus AULOPHYLLUM.

Corallum simple. Septa well developed. A double mural investment; the interior
wall dividing the visceral chamber into two portions—one central and columnar, the other
exterior and annular. No columella. Tabule but little developed.

Typ. sp., Aulophyllum prolapsum ; Clisiophyllum prolapsum, M‘Coy, in Ann. and Mag. of Nat. Hist.,

2d series, vol. ili, p. 3.

24. Genus ACRRVULARIA.
Schweigger, Handb. der Naturg., p. 418, 1820.

Corallum composite, increasing by calicular gemmation. Corallites provided with a
double mural investment; the inner wall disposed as in the preceding genus. Septal
apparatus well developed between the outer and the inner walls, but much less so in the
central area. No columella. Tabule not well developed.

Typ. sp., Acervularia Remert; Astrea Hennahi, Roemer, Verst. der Hartzgeb., tab. ii, fig. 13 (not
Lonsdale).

25. Genus STROMBODES.

(Pars) Schweigger, Handb. der Naturg., p. 418, 1820; Goldfuss, Petref. Germ., vol. i, p. 62, 1826;
Acervularia, Lonsdale, Sil. Syst., p. 689, 1839; Arachnophyllum, Dana, Zooph., p. 360, 1846 ; Strombodes
and Actinocyathus, D’Orbigny, Prod. de Paléont. stratigr., vol. i, p. 107, 1849.

Corallum composite, increasing by calicular gemmation. Coradlites constituted prin-
cipally by a series of superposed, invaginated, infundibuliform ¢abw/e, united by ascending
trabicule, so as to form a columnar mass. Cadices pentagonal, well circumscribed, and

completely covered with the septal radu. Outer «walls not well developed; the inner
mural investment rudimentary.

Typ. sp., Strombodes pentagonus, Goldf., Petref. Germ., vol. i, tab. xxi, fig. 3.

26. Genus PHILLIPSASTREA.
D’Orbigny, Note sur des Polypiers fossiles, p. 2, 1849.

Corallum composite, resembling Strombodes, but differing from them by the septal

INTRODUCTION. Ixxi

or costal radi of the neighbouring corallites, being confluent, and consequently the calices
not being definitely circumscribed. No exterior walls; the interior mural investment
well characterised. The centre of the ¢adu/e presenting a columellarian tubercle.

Typ. sp., Phillipsastrea Hennahi, D’Orbigny, loc. cit.; Astrea Hennahi, Lonsdale, in Geol. Trans.,
2d series, vol. v, tab. lviul, fig. 3.

27. Genus ERIDOPHYLLUM.

Corallum composite, and increasing by lateral gemmation. Corallites tall, cylindroid,
and provided with a thick epitheca, which gives rise to a vertical series of short and thick
subradiciform productions that extend to the next individual and unite them together.
Tabule well developed, and occupying the central area circumscribed by the inner
wall. Septal apparatus occupying the annular area situated between the outer and inner
mural investment, but not extending into the inner or central area.

Typ. sp., Eridophyllum seriale, nob.

Third Tribe—LITHODENDRONIN .

Axis of the visceral chamber of the corallites occupied by a styliform or lamellar
columella.

28. Genus LITHODENDRON.

Phillips, Geol. of Yorkshire, vol. ii, p. 200, 1835 (but not Lithodendron of Schweigger, which is not an
admissible genus) ; Siphonodendron, M‘Coy, in Ann. and Mag. of Nat. Hist., 2d series, vol. ii, 1849.

Corallum composite, arborescent, or massive. Corallites cylindrical or prismatic.
Columella styliform, compact. Septa well developed, but not reaching to the columella.
Tabule well developed. Interior wall rudimentary.

Typ. sp., Lithodendron irregulare, Phillips, loe. cit., pl. ii, figs. 14, 15.

29. Genus NEMATOPHYLLUM.

Nematophyllum and Stylaxis, M‘Goy, loc. cit., 1849.

Corallum composite, massive. Corallites prismatic, with a well-developed interior
wall. Columella \amellar. Septa well developed, and united by transverse dissepi-
ments, which extend to the columella, but do not constitute true tabule. Exterior area
vesicular.

Typ. sp., Nematophyllum arachnoideum, M‘Coy, loc. cit., p. 16.

Ixxii BRITISH FOSSIL CORALS.

30. Genus LiTHOSTROTION.

(Pars) Fleming, British Animals, p. 508, 1828; Strombodes and Lonsdaleia, M‘Coy, in Ann. of Nat. Hist.,
2d series, vol. iii, pp. 10, 11, 1849.

Corallum resembling Mematophyllum, but having the columella formed by a fasciculus
of twisted bands, and the septa subvesicular exteriorly, and joining the columella along
their inner edge.

Typ. sp., Lithostrotion floriforme, Fleming, loc. cit., p. 508.

31. Genus AXOPHYLLUM.

Corallum simple, trochoid, and resembling Lithostrotion by its structure.

Typ. sp., Avophyllum expansum, nob.

32. Genus SYRINGOPHYLLUM.
Sarcinula, Dana, Zooph., p. 363, 1846 (not Sarecinula, Lamarck).

Corallum composite, astreiform. Corallites provided with strong walls, and much
developed coste. Septa large. Tubule but little developed. Colwmedla styliform.

Typ. sp., Syringophyllum organum ; Madrepora organum, Linnzeus, Syst. Nat., ed. xii, vol. i, p. 1278.

Family XV.
CYSTIPHYLLID A.

Corallum essentially composed of a vesicular tissue, and presenting little or no traces
of septa or radiate strize.

1. Genus CYsTIPHYLLUM.
Lonsdale, in Murchison’s Silurian Syst., p. 691, 1839.

Corallum simple, turbinate ; the visceral chamber filled with small vesicular lamine.
Calice shallow. Walls vesicular.

Typ. sp., Cystiphyllum Siluriense, Lonsdale, loc. cit., tab xvi bis, fig. 1 (but not fig. 2).

Sub-order 5.
ZOANTHARIA CAULICULATA.
Antipathacea, Dana, Zooph., p. 574.

Polypi supported on a sclerobasis or epidermic stem-like corallum.

INTRODUCTION. Ixxill
The general form of the corallum is similar to that of the Isis, Gorgonia, &., in the

order of Alcyonaria; but may be distinguished from these by its surface being spinulous
or smooth, whereas it is always sulcated in Alcyonaria.

Family ANTIPATHID A.

Gray, Synop. of the Brit. Mus., p. 135, 1842; Dana, Zooph., p. 574, 1846.

1. Genus ANTIPATHES.
(In parte) Pallas, Elench. Zooph., p. 209, 1766.
Corallum arborescent ; its surface spinulous.

Typ. sp., Antipathes myriophylla, Ellis and Solander, Zooph., tab. xix, figs. 11, 12.

2. Genus CIRRHIPATHES.
De Blainville, in Dict. des Sc. Nat., vol. lx, p. 475, 1830.

Corallum not arborescent, and having the form of a simple cylindrical stem ; its surface
spinulous.

Typ. sp., Cirrhipathes spiralis, Blainv., loc. cit.; Antipathes spiralis, Ellis and Soland., Zooph., tab. xix,
fig. 1.

3. Genus LEIOPATHES.
Gray, Synops. of the Brit. Mus., p. 135, 1842.
Corallum arborescent ; its surface smooth.

Typ. sp. Leiopathes glaberrima ; Antipathes glaberrima, Esper, Pflanz., Antipathes, tab. ix.

ZOANTHARIA INCERTA SEDIS.
1. Genus HeTeROPHYLLIA.
M Coy, Palzeozoic Corals, in Ann. and Mag. of Nat. Hist., 2d series, vol. iii, p. 126, 1849.

Corallum composed of a tall, subcylindrical, irregularly fluted, stem (or tube), contaiming
a few laminz, irregularly branching and coalescing, but not presenting a radiate appearance.

Typ. sp., Heterophyllia grandis, M‘Coy, loc. cit., figs. a, B.
10

Ixxiv BRITISH fOSSIL CORALS.

2. Genus Morrinria.
De Koninck, Anim. foss. du Terr. carbon. de Belgique, p. 12, 1842.

Corallum having the form of a bi-concave disc, presenting a radiate structure and
numerous coste.

Typ. sp., Mortieria vertebralis, De Koninck, loc. cit., pl. B, fig. 3.

3. Genus CYCLOCRINITES.
Eichwald, Uber das Silurische Schichten-System in Esthland, p. 192, 1840.

Corallum composite, astreiform. Cadices hexagonal and shallow. Seta well charac-
terised, but not extending to the centre of the visceral chamber, which appears to be
occupied by small tabule. (?)

Typ. sp., Cyclocrinites Spaskit, Eichwald, Die Urwelt Russlands durch abbildungen erlaeutert, p. 48,
tab. i, fig. 8, 1842.

Order 11.
ALCYONARIA.

Alcyoniens, Audouin and Milne Edwards, Recherches sur les Anim. sans Vertébres faites aux iles Chausay,
Ann. des Sc. Nat., Ist series,, vol. xv, p. 18, 1828; Zoophytaria, Blainville, Manuel d’Actinologie, p. 496,
1834; Zoophyta asteroidea, Johnston, Brit. Zooph., p. 164, 1838; Aleyonaria Dana, Exploring Expedition,
Zooph., p. 586, 1846 ; Anthozoa asteroidea, Johnston, Brit. Zooph., 2d edit., p. 138, 1847.

Polypi with bi-pinnate tentacula, and only eight perigastric membranaceous lamine,
containing the reproductive organs.

Alcyonaria have, in general, their dermal tissue consolidated by isolated spicule or
nodular concretions only, and very rarely present a vaginal polypidom similar to that of
the Zoantharia; but even when that is the case, the visceral chamber is never subdivided
by any longitudinal septa, and consequently the casice never presents any appearance of
radii. In general, the corallum is entirely composed of epidermic tissue, (or basal secretion,
Dana,) and constitutes a sort of stem or axis in the certre of the compound mass formed
by the gemmation of the Polypi. This sclerobasis is always covered by soft dermic
tissue, and increases by the addition of concentric layers.

This order is far from being as numerous as the preceding division of Corallaria, and
comprises three natural families,—Alcyonidee, Gorgonide, and Pennatulidee.

INTRODUCTION. Ixxv

Family I.
ALCYONIDA.
Polypi adherent and not provided with an epidermic sclerenchyma.

In this family, the dermic tissue is usually consolidated by a great number of scleren-
chymous spicula imbedded in its substance, and constitutes sometimes a tubular corallum,
but there is never any trace of a central stem or axis, like that which is constituted by the
sclerobasis in Gorgonide and in most of the Pennatulide.

First Tribe—CORNULARIN &.

Polypi simple or segregate, and produced by gemmation on creeping stolons, or basal
membranaceous expansions, and having no lateral buds or connecting appendices.

1. Genus CoRNULARIA.
Lamarck, Hist. des An. sans Vert., vol. ii, p. 111, 1816.

Polypi rismg by gemmation from creeping filiform stolons, and provided with a tough
or subcorneous tubiform polypidom, the surface of which is not costulated.

Typ. sp., Cornularia cornucopia, Cuvier; Tubularia cornucopia, Cavolini, Mem. per Servire alla Storia
de Polipi Marini, tab. ix, figs. 11, 12; Cornularia rugosa, Lamarck, loc. cit.

2. Genus CLAVULARIA.

Quoy and Gaimard, ap. Blainville, Dict. des Sc. Nat., vol. lx, p. 499, 1830; Actinantha, Lesson, Zool.
de la Coquille, Zooph., p. 89, 1831.

Polypi resembling Cornularia, but having their tubular polypidoms costulated and
incrustated with long spicula.

Typ. sp., Clavularia viridis, Quoy and Gaim., Voyage de l’Astrolabe, Zooph., tab. xxi, fig. 10.

3. Genus RHIZOXENIA.

Ehrenberg, Corall. Roth. Meer., p. 55, 1834.

Polypi resembling those of the preceding genus, but not retractile.

Typ. sp., Rhizoxenia thalassantha, Ebr. ; Zoantha thalassantha, Lesson, Voyage de la Coquille, Zooph.,
tab. i, fig. 2.

Ixxvi BRITISH FOSSIL CORALS.

4. Genus SARCODICTYON.
E. Forbes ap. Johnston, Brit. Zooph., 2d ed., p. 179.
Polypi rising from creeping, filiform, anastomosing stolons, distant, uniserial, and

appearing verruciform (not tubular) when retracted. Differ from Cornularia by the short-
ness of the polypidoms.

Typ. sp., Sarcodictyon catenatum, Forbes, loc. cit., tab. xxxili, figs. 4, 7.

5. Genus ANTHELIA.
Savigny, ap. Lamarck, An. sans Verteb., vol. ii, p. 407, 1816.

Polypi not retractile, and rising from a thin fleshy incrustating plate.

Typ. sp., Anthelia glauca, Savigny, Egypte, Polypes, tab. i, fig. 7.

6. Genus SYMPODIUM.
Ehrenberg, Corall., p. 61, 1834.
Polypi resembling Anthelia, but being retractile.

Typ. sp., Sympodium fuliginosum, Ehrenb., Savigny, Egypte, Polypes, tab. i, fig. 6.

7. Genus AULOPORA.
Goldfuss, Petref. Germ., vol. i, p. 82.

The fossil corals forming this genus greatly resemble Cornularia and Sarcodictyon, but
differ from all the preceding genera by their thick, calcareous polypidom.

Typ. sp., Aulopora serpens, Goldfuss, loc. cit., tab. xxix, fig. 1.

8. Genus CLADOCHONUS.
M‘Coy, in Ann. and Mag. of Nat. Hist. Ist series, vol. xx, p. 227.

Corallum resembling Aulopora, but composed of cup-shaped calices, arranged in a
regularly alternate manner, and bent in nearly opposite directions.

Typ. sp., Cladochonus tenuicollis, M‘Coy, loc. cit., tab. xi. fig. 8.

INTRODUCTION. Ixxvil

Second Tribe—TUBIPORIN Ai.
Polypi fasciculate, and provided with independent tubular polypidoms, united at

various heights by means of horizontal connecting plates, the surface of which produces
the reproductive buds.

9. Genus TUBIPORA.
Lamarck, Hist. des Anim. sans Verteb., vol. ii, p. 207, 1816.

Typ. sp. Tubipora musica, Lamarck, loc. cit., p. 209.

Third Tribe—TELESTHIN 2.

Polypi segregate and multiplying by lateral gemmation, so as to form arborescent
tufts.

10. Genus TELESTHO.
Lamouroux, Polypiers Flexibles, p. 232.
Polypidom composed of ramified tubes of a subcalcareous structure.

Typ. sp., Telestho aurantiaca, Lamouroux, loc. cit., tab. vii, fig. 6.

Fourth Tribe—ALCYONIN /.

Polypi aggregate and multiplying by lateral gemmation, so as to constitute a ramified,
lobate or simple mass.

ll. Genus ALcyonium.

Pallas, Elenchus Zooph., p. 342, 1766; Lobularia, Savigny, ap. Lamarck, Hist. des Anim. sans Verteb.
vol. ii, p. 412, 1816.

Polypi retractile, and united by a thick tough common tissue, so as to form gibbose
or subramified masses.

Typ. sp., Aleyonium digitatum, Lin. Solander and Ellis, op. cit., p. 175.

Ixxvili BRITISH FOSSIL CORALS.

12. Genus XENIA.
Savigny, Egypte, Atlas and op., Lamarck, op. cit., vol. ii, p- 629, 1816.

Polyji forming subramified masses, as in Alcyonium, but not retractile, and not having
a thick coating of spiculze at the basis of the tentacula.

Typ. sp. Xenia umbellata, Savigny, Egypte, Polyp., tab. i, fig. 3.

13. Genus NEPHTHYA.

Savigny, Atlas de ’ Egypte; Blainville, Manuel d’Actinol, p. 523; Spoggodes, Lesson, Illustr. de
Zoologie, 1831.

Polypi forming arborescent masses, incompletely retractile, and having the borders of
the calice thick and incrustated with large navicular spiculee.

Typ. sp,, Nephthya Chabroli, Audouin, ap. Savigny, Egypte, Pol. tab. i. fig. 5.

14. Genus PARALCYONIUM.
Alcyonidia, Milne Edwards, Ann. des Sc. Nat. 2d series, vol. iv, p. 323.

Polypi resembling Nephthya, but being completely retractile, and having the lower
part of the common mass incrustated with a thick coating of long navicular spicule, but
the upper part membranaceous and retractile.

Typ. sp., Paraleyonium elegans ; Alcyonidia elegans, Milne Edwards, loc. cit., tab, xii and xin.

15. Genus SARCOPHYTON.
Lesson, Zoologie du Voyage de la Coquille, Zooph., p. 92, 1831.

Differs from the genus Alcyonium by the great abundance and the peculiar structure
of the common tissue, the cells of which are tubular, and arranged with great regularity
in fasciculi, perpendicularly to the upper surface of the mass.

Typ. sp., Sarcophyton plicatum, Valenciennes MSS.; Sarcophyton lobulatum, Lesson, loc. cit. ; Alcyo-
nium plicatum, Lamarck, Hist. des An. sans Verteb., vol. 11, p. 395.

16. Genus CESPITULARIA.
Valenciennes MSS.

Polypi non-retractile, arranged in fasciculi, and united in the greatest part of their
length by a dense, tough, common tissue, as in Alcyonium.

typ. sp., Cespitularia multipinnata, Valen.; Cornularia multipinnata, Quoy and Gaimard, Voyage de
V Astrolabe, Zooph., tab. xxii, figs. 1-4.

INTRODUCTION. Ixx1x

17. Genus DisTICHOPORA ?
Lamarck, loc. cit., p. 197.

This singular zoophyte appears to have more affinity to Alcyonium than to any other
form of polypi; but the place belonging to it in a natural system of classification is as
yet very uncertain. It is characterised by a calcareous, dendroid corallum, composed of
long tubular cells, that present no traces of septa or tubule, and are disposed in a
flabellate manner, so as to constitute a vertical plane, the two sides of which are
covered with a thick and compact ccenenchyma, and the edge assumes the appearance of
a calicular groove, limited laterally by two rows of circular pores. Nothing is known
concerning the structure of the soft parts.

Typ. sp., Distichopora violacea, Lamarck, op. cit., p. 305. (For the structure of the Corallum, see
Milne Edwards, Atlas du Regne Animal de Cuvier, Zooph., tab. Ixxxv, fig. 4.)

Family II.
GORGONID &.

Polypiers corticiferes, Lamarck, Hist. des Anim. sans Verteb. vol. ii, p. 288, 1816; Polypes corticaus,
Cuvier, Regne Animal, vol. iv, p. 78, 1817; Corallia, Blainville, Manuel d’Actinologie, p. 501, 1834;
Cerato-corallia, Ehrenberg, Corall. des Roth. Meeres, 1834; Coralliade, Gray, Synop. Brit. Mus. p. 134;
Gorgoniade, Johuston, British Zooph., p. 182, 1838; Gorgonide, Dana, Exploring Expedition, Zooph.,
p. 637, 1846; Gorgoniade, Gray, List of British Anim. of the British Museum, p. 55, 1848.

Polypi provided with a thick, suberous ccenenchyma, surrounding a central stem that
is adherent to an extraneous body by its basis, and is formed of epidermic sclerenchyma.

First Tribe—GORGONIN AL.

Gorgonia, Pallas, Elenchus Zoophytorum, ; 160, 1766; Cuvier, Regne Animal, Ist ed., vol. iv, p. S0;
Lamarck, Hist. des Anim. sans Verteb., vol. ul, p. 309; Gorgonine, Dana, Exploring Expedition, Zooph.,
p- 641, 1846.

Common axis inarticulate, horny or fasciculate, but not calcareous.

]. Genus GoRGONIA.
Pallas, loc. cit., (in parte.)

Axis corneous. Cadices disposed irregularly round the ramified cylinders formed by
the ccenenchyma, and not encircled by imbricated squamme. Polypz retractile.

Typ. sp., Gorgonia tuberculata, Esper. Pflanz., Gorg., tab. xxxvil.

Ixxx BRITISH FOSSIL CORALS.

2. Genus PTEROGORGIA.

Ehrenberg, Corall. des Rothen Meeres, p. 144, 1834 ; Dana, op. cit., p. 647, 1846.

Differs from Gorgonia by the polypi being bifarious.
Typ. sp., Pterogorgia anceps, Ehrenb., loc. cit., p. 145.

3. Genus BEBRYCE.

Philippi, Zoologesche Beobachtungen, in Archiv. fur Naturgeschichte, von Erichson, vol. viii, p. 35, 1842.

Arborescent compound polypi, resembling Gorgonia by their corneous sclerobasis, but
differing from the preceding genera by not being retractile.

Typ. sp., Bebryce mollis, Philippi, loc. cit.

4. Genus PHYLLOGORGIA.

Differs from Gorgonia by the coenenchyma not constituting a cylindrical sheath around
the ramifications of the sclerobasis, but extending between them so as to constitute large
foliaceous, frondiform laminz, the two surfaces of which are studded with the calices of
the individual polypi.

Typ. sp., Phyllogorgia dilatata ; Gorgonia dilatata, Esper, Pflanz. Gorg. tab. xli.

5. Genus PHYCOGORGIA.
Sclerobasis flabelliform, divided into digitated lobes, and composed of delicate corneous

fibres united into laminz, the two sides of which are covered with the ccenenchyma,
and densely studded with numerous non-prominent calices.

Typ. sp., Phycogorgia fucata ; Gorgonia fucata, Valenciennes, Voyage de la Venus, tab. xi, fig. 2.

6. Genus Muricnra.
Lamouroux, Exposit. Method. des Polyp. p. 36, 1821.

Differs from Gorgonia by the calices being surrounded with imbricated squammula,
but not supported on long, verruciform, moveable appendices, as in Primnoa.

Typ. sp., Muricea spicifera, Lamouroux, op. cit., tab. Ixxi, figs. 1, 2.

7. Genus Primnoa.
Lamouroux, Hist. des Polypiers Flexibles, p. 440, 1816.

Differs from the preceding genus by the polypi constituting long verruciform subpedi-
culated appendices, which are capable of motion at their bases.

INTRODUCTION. Ixxx1

8. Genus SOLANDERIA.
Duchassaing and Michelin, in Guerin’s Revue Zoologique, June, 1846.
Differs from Gorgonia by the suberous texture of the sclerobasal axis, which resembles

the non-calcified jomts of Meliteea.
Typ. sp., Solanderia gracilis, Duchassaing and Michelin, loc. cit.

9. Genus BRIAREUM.
Blainville, Manuel d’Actinologie, p. 520, 1830.

Axis soft, suberous, or composed of spicula. This genus is intermediate between
Aleyonium and Gorgonia.

Typ. sp., Briareum gorgonoideum, Blainy.; Gorgonia briareus, Lin.; Ellis and Solander, tab. xiv, figs. 1, 2.

The genus Hyatonrma established by M. Gray, (‘ Proceed. of the Zool. Soc.’ 1835,
p. 68,) is also referred by some zoologists to the tribe of Gorgonine; but the recent
observations of M. Valenciennes tend to establish that the fasciculi of siliceous threads,
which constitute the axis of this smgular production, belong to the class of Spongidz, and
the polypi which we have observed in a dried state on different parts of the axis appear to
be parasites, belonging to the order of Zoantharia.

Second Tribe—ISIN A.
Dana, Exploring Exped., Zooph., p. 677, 1846.

Common axis articulated, or composed of segments, the structure of which differ
alternately.

10. Genus Isis.
Linneus, Syst. Nat., 12th ed., p. 1287, 1767.
Axal sclerobasis composed of joints, alternately corneous and calcareous ; branches

proceeding from the calcareous joints.

Typ. sp., Isis hippuris, Lin., loc. cit.

11. Genus Mopska.

Lamouroux, Polyp. Flex., p. 466, 1816.

Axis presenting the same structure as in the preceding genus, but with the branches
proceeding from the corneous joints.

Typ. sp., Mopsea dichotoma, Lamouroux, loc. cit., p. 467.

11

Ixxxul BRITISH FOSSIL CORALS.

12. Genus Muiivaza.
Lamouroux, Polyp. Flex., p. 461, 1816.
Axis composed of joints, which are alternately calcareous and suberous.

Typ. sp., Melitea ochracea, Lamouroux, loc. cit., p. 462.

Third Tribe— CORALLIN #&.
Dana, loc. cit., p. 639, 1846.

Common axis inarticulate, solid and calcareous.

13. Genus CoRALLIUM.
Lamarck, Hist. des Anim. sans Vert., t. ii, p. 295, 1816.

Typ. sp., Corallium rubrum, Cavolini, Mem. per Servire all. Hist. des Polypi Marini, tab. ii.

Family IIT.
PENNATULID.

Pennatula, Linneus, Syst. Nat., 10th ed., p. 818; Pallas, Elen. Zooph., p. 362, 1766; Polypi natantes,
Lamarck, op. cit., p. 415; Pennatulide, Fleming, Brit. Animals, p. 507, 1828; Pennatularia, Blainville,
Manuel, p. 512, 1830; Calomides, Latreille, Fam. du Reg. Anim. p. 543; Pennatulina, Ehrenberg, loc.
cit., p. 63, 1834; Pennatulide, Johnston, Brit. Zooph., p. 175; Dana, Explor. Exped. p. 557, 1846.

Polypi aggregate, and having a common peduncle, the centre of which is occupied
by a peculiar cavity, and usually contains a solid axis; this sclerobasis styliform, and never
expanding at its under extremity, so as to adhere to extraneous bodies. The polypi mass
is consequently free.

1. Genus PENNATULA.
(In parte.) Linnzus, Syst. Nat., 10th ed., p. 818, 1760; Lamarck, Syst. des An. sans Vert., p. 380, 1801.

Polype mass plume-shaped, with the shaft composed of contractile common tissue, con-
taining a short subosseous axis, and bearing on each side of its upper part a series of
large spreading pinnules, on the upper edge of which, the retractile exhalic portion of the
polypi protudes. The axis is cylindrical at its upper part, and more or less quadrangular
towards its lower end; its structure is somewhat fibrous, and its tissue is not very brittle.

Typ. sp., Pennatula setacea, Esper, Pflanz., Pennat. tab. vii.

INTRODUCTION. IXxxiil

2. Genus VIRGULARIA.
Lamarck, Hist. des Anim. sans Verteb. vol. ii, p. 429, 1816.

Differs from Pennatula by the length of its shaft and the shortness of its pinnules,
which assume the form of lunate lobes, or simple transverse striz. Axis cylindrical,

calcareous, very long, slender, tapering, and presenting in its transverse section a radiate
structure.

Typ. sp., Virgularia mirabilis, Lamk.; Pennatula mirabilis, Miller, Zool. Danica, vol. i, tab. xi.

3. Genus PAVONARIA.
Cuvier, Regne Animal, vol. iv, p. 85, 1816; Funicularia, Lamarck, op. cit., p. 423, 1816.

Polype mass virgate; the polypi not retractile, arranged on one side of the stem.
Axis quadrangular, long, and very tapering.

Typ. sp., Pavonaria quadrangularis, Cuv.; Pennatula antennina, Lin.; Johnston, Brit. Zooph., tab. xxx.

4. Genus GRAPHULARIA.

Corallum styliform, straight, very long, cylindroid towards the lower extremity, sub-
tetrahedra] at the upper part, and presenting on one side a broad shallow furrow. _‘T'rans-
verse section showing the existence of a thin coating, and a radiate structure in the body
of the coral.

Typ. sp., Graphularia Wetherelli, nob. ; Pennatula, Sowerby and Wetherell, in Geol. Trans. 2d series,
vol. v, part i, p. 136, tab. viii. fig. 2.

5. Genus UMBELLULARIA.

Cuvier, Regne Animal, vol. iv, 1807.

Resembling Payonaria, but having all the polypi collected in a terminal bunch at the
extremity of the stem. Axis quadrangular and twisted.

Typ. sp., Umbellularia Groenlandica, Cuv.; Hydra Marina arctica, Ellis, Corallines, tab. xxxvil.

6. Genus VERETILLUM.
Cuvier, Regne Animal, vol. iv.
Resembling Pennatula, but not having any lateral pinnules, with the polypi
arranged all round the upper part of the stem. Axis rudimentary, and of a form almost

navicular.

Typ. sp., Veretillum cynomorium, Cuvier; Pennatula digitiformis, Ellis.

Ixxxiv BRITISH FOSSIL CORALS.

7. Genus LITUARIA.
Valenciennes MSS., Cat. of the Zoophytes in the Museum of Paris.

Resembling Veretillum, but having a long well-developed axis, quadrangular and
tapering towards its lower part, inflated, claviform, pitted and echinulate at its upper end.

Typ. sp., Litwaria phalloides, Valenciennes, loc. cit.; Pennatula phalloides, Pallas, Miscel. Zool.;
tab. xiii.

8. Genus CAVERNULARIA.

Valenciennes, loc. cit., MSS.

Resembling Veretillum, but having in its centre a large fibrous tube divided longi-
tudinally into four cavities, and not containing any calcareous or horny axis.

Typ. sp., Cavernularia obesa, Valenciennes MSS.

9. Genus RENILLA.

Lamarck, Hist. des Anim. sans. Verteb., vol. ii, p. 428, 1816.

Polyp-mass explanate, unifacial, reniform, with a short, slender peduncle, containing
a central cavity as in Pennatula, but not having any solid axis.

Typ. sp., Renilla Americana, Lamarck; Pennatula reniformis, Ellis and Solander, p. 67; Shaw,
Miscel. iv, tab. exxxix.

The genus Graprotitaus (Linneus, J¢er Scan. 1751,) appears to have more affinity
with Virgularia than with any other recent zoophyte. The polype mass is slender, virgate,
and often becomes bifurcate by the progress of growth. The axis projects at the inferior
extremity of the stem, and is often bifurcate.

Example, Graptolithus ramosus, Hall, Palezeont. of New York, tab. Ixxiii, fig. 3.

The genus WesstTeERia, nob. appears to be very similar to Graptolithus by its general
structure, but offers also a certain resemblance to some Sertularide and to certain Bryozoa.
In the present state of our knowledge, the natural affinities of these fossil zoophytes are
indeed so obscure, that we hesitate to place them in any of the preceding zoological
divisions, and prefer leaving them in the cxcerte sedis.

Typ. sp., Websterta Crisioides, nob., tab. vii, fig. 5.

INTRODUCTION. Ixxxv

Order 3.
PODACTINARIA.

Polypi having the gastric cavity surrounded by four vertical membranaceous septa, at
the upper end of which are placed four pairs of intestiniform reproductive organs. The
tentacula discoidal, pedunculated, not tubular as in Zoantharia and Alcyonaria, but
organized much in the same way as in Echinoderma. The mouth proboscidiform, and
the fauces surrounded by numerous internal, filiform, contractile appendices.

The genus Lucrrnaria is the only known representative of this zoological type, and
comprises no coralligenous polypi.

Sub-class 2.
EY REARS A:

Polypes sertulariens, Audouin and Milne Edwards, Recherches sur les Anim. sans Verteb., faites aux iles
Chausay, in Ann. des Sc. Nat., Ist series, vol. xv, p. 18, 1828, ap. Lamarck, Hist. des An. sans Verteb.
2d ed., vol. ii., p. 105; Sertulariacea (in parte), Blainville, Manuel d’Actinologie, p. 465, 1834;
Zoocorallia oligactinia, Ehrenberg, Coral. Roth. Meeres, p. 67, 1834; Zoophyta Hydroida, Johnston, in
Mag. of Zool. and Bot., vol. i, p. $47, 1837; Polyparia, Gray, Synop. Brit. Mus. ; Nudibranchiata, ¥arre,
on the Structure of Polypi, Phil. Trans. 1837; Hydrozoa, R. Owen, Lectures on the Comp. Anat. of the
Inyerteb. Animals, p. 82, 1843; Hydroidea, Dana, Exploring Expedition, Zooph. p. 685, 1546;
Anthozoa Hydroidea, Johnston, British Zooph, 2d ed. p. 5, 1847.

Polypi with a simple, non-lamelliferous, digestive cavity. No internal generative
organs. ‘Tentacula filiform and subverrucose.

The naked, fresh-water zoophytes of the genus Hypra constitute the type of this
group, and till very lately were considered as being closely allied to Sertnlaria, Canrpanu-
laria, &c.; but the recent observations of divers zoologists tend to establish that all the
coralligenous animals of this form belong to the class of Medusa. ‘Till this question is
decided, it would therefore be idle to make any modifications im the systematic arrange-
ment of these problematic polypi, and it will suffice for us to refer the reader to Dr.
Johnston’s valuable work on ‘British Zoophytes,’ for the characters of the generic
divisions generally adopted.

STE eat ites
ra ;
AOL eat FF

=

DESCRIPTION

OF

THE BRITISH FOSSIL CORALS.

CHAPTER I.

CORALS OF THE CRAG.

Tue Crag formation of the Hast of England is generally reputed very rich im Fossil
Corals; and the name given to the lower strata of this system is even derived from the
abundance of various organic remains of coralloid appearance which occur in some
localities. But this opinion arises from the confusion which has till lately been made
between Bryozoa and Polypi; in reality true Corals are far from being common in any of
these beds. The four species mentioned by Mr. Searles Wood, in the Catalogue of the
Zoophytes of the Crag, published in 1844 in the ‘ Annals of Natural History,’ are the only
known Polypidoms belonging to this geological division.

These fossils are found in the Red Crag as well as in the Coralline Crag, and most of
them are as yet peculiar to England; only one species has been met with on the Continent,
in the Crag of Antwerp, astrata belonging to the same geological horizon; and none of
them are known to live in the seas of the present period. The Sphenotrochus intermedius
has, it is true, been considered as existing on the coast of England as well as in the Crag ;
but the recent species, which has lately received the name of Sphenotrochus Andrewvanus,!
is perfectly distinct from the fossil Coral to which it was at first referred. It is also worthy
of remark that the Crag Corals belong to four distinct genera, each of which is represented
by different species in the other Miocene formations ; that three of these genera are also
represented by peculiar species in our actual Fauna, and that none of them have been
discovered in strata anterior to the older tertiary formations.

1 Milne Edwards and J. Haime, Monographie des Turbinolides, Ann. des Sc. Nat., 3™° serie, vol. ix,

p. 245, tab. vii, fig. 4.
1

2 BRITISH FOSSIL CORALS.

Orper ZOANTHARIA (p. ix).
Family TURBINOLID® (p. xi).
Tribe TURBINOLIN & (p. xvi).

Genus SPHENOTROCHUS (p. XVI).

1. SPHENOTROCHUS INTERMEDIUS. ‘Tab. I, figs. 1, 1 a—lz.

TURBINOLIA INTERMEDIA, Miinster, ap. Goldfuss, Petref. Germ., vol. i, p. 108, tab. xxxvii,

fig. 19, 1826. (This figure is good, excepting that the basis of
the Coral appears too truncate.)

Ch. Morren, Descrip. Corall. foss., in Belgio Repertorum, p. 52,
1828.

R. C. Taylor, in Mag. of Nat. Hist., vol. iii, p. 272, fig. 2,
1830. (A rough figure.)

— INTERMEDIA, Milne Edwards, Notes in the second ed. of Lamarck’s Anim. sans

Vert., vol. ii, p. 361, 1836.

— Galeotti, Mém. couron. par l’Acad. de Bruxelles, vol. xii, p. 188,
1837.
— Hagenow, in Neues Jarhb. fiir Miner. Geol., 1839, p. 291.
— — Nyst, Coquilles et Poly. foss. des Terr. Tert. de la Belgique,
p. 631, tab. xlviil, fig. 14, 1843. (This figure is incomplete,
and does not show the columella.)
— Minietrana, Searles Wood, Ann. and Mag. of Nat. Hist., vol. xiii, p. 12, 1844.
SPHENOTROCHUS INTERMEDIUS, Milne Edwards and Jules Haime, Monogr. des Turbinolides,
Ann. des Sc. Nat., 3™° série, vol. ix, p. 243, 1848.

Corallum simple, straight, free, presenting no trace of adherence, cuneiform, strongly com-
pressed in its lower part, and truncate at its basis, which is very broad (fig. 1) ; sometimes
even as much so as the calice (fig. 1a). This last character exists also in the Spheno-
trochus Milletianus ; but this Coral, mstead of bemg much compressed in the lower part,
is, on the contrary, very thick down to its extremity.

Coste smooth, rather thick, especially near the calicular edge, closely set, but separated
by deep grooves (fig. 16). They all occupy almost the whole length of the corallum ; and
it is therefore difficult to recognise their relative age by the height at which they begin.
This difficulty is also augmented by the form of those situated near the middle of the
flattened sides, which in their lower part are constituted by small, rather irregular papille.
The median cost are nearly straight, nearly equal, not very prominent, and narrowing as
they approach the base; the lateral coste, and those situated near them, are, on the
contrary, larger, separated by deeper grooves, slightly curved towards their lower end,
sometimes rather undulate, and thicker at their base than higher up.

CORALS OF THE CRAG. 3

It may also be worth remarking that similar smooth and simple cost exist in all the
species of this genus belonging to the present period or to the Miocene deposits; whereas
the older species, found in the Eocene formation, have the costz crispate, and composed of
series of papille.'

The calice (fig. 1a) is regularly elliptic and slightly arched, the extremities of its
great axis being lower than those of its small axis. ‘The proportion between the two
diameters is nearly constant, and the form of the ellipse, represented by the calicular
margin, is intermediate between that of Sphenotrochus granulosus, which is much shorter,
and that of Sphenotrochus Andrewianus, which is more elongated; it is approximately
as 100: 150. The size of the calice is also subject to very slight variations in individuals
which have attained their definitive form, whether they be short or tall.

The fossula is very shallow.

The columella (figs. 1a and 1c) has the form of a rather thin, vertical lamina, situated in
the direction of the long axis of the calice and of the basal edge of the corallum. Its
upper edge is nearly horizontal, and reaches almost to the level of the apex of the septa ;
it is obtuse at its angles, and divided into two equal lobes by a small notch ; sometimes
three of the lobes are visible. The structure of this part of the polypidom may be very
well shown by a vertical section corresponding to the small axis of the calice ; it is formed
by two delicate parallel lamin, applied together, thickened near its upper edge, and united,
towards its base, to the wall, so as to form with the mural sclerenchyma one compact
mass.

The septa, as in all the other species of this genus, form three complete and well-
developed cycla (fig. ia); they are consequently twenty-four in number, and they are
closely set, straight, thick exteriorly, and becoming gradually thinner towards the centre of
the calice, exsert, arched at their apex, truncate at the upper end of their inner edge, and
granulated on their surface. These granulations are easily brought to view by a vertical
section of the corallum (fig. lc); they are small, unequal in size, pointed, not numerous,
and not disposed in a regular manner, excepting near the upper edge of the septa, where
they form a curved line nearly parallel to the edge. The septa of the first and second
cycla are nearly similar; and, as is often the case in Corals with an elliptic calice, the
two primary septa, corresponding to the long axis of the calice, are a little smaller than
the four others of the same cyclum, and the six secondary ones. ‘The tertiary septa are

1 The fossil Coral figured by Mr. Isaac Lea, under the name of Turbinolia nana, and mentioned by
that author as belonging to the Eocene strata of Alabama, would appear to be an exception to this rule,
for it resembles much the Sphenotrochus Milletianus, and seems to have smooth costz; but the figure given
by Mr. Lea is not sufficiently explicit for us to be able to decide the question, or even to be quite sure
that this Turbinolida really belongs to the genus Sphenotrochus, and in the text the author says that
he could see no trace of a columella (Lea, Contrib. to Geol., p. 195, tab. vi, fig. 209). In the present state
of palzeontology, we may, therefore, consider the above-mentioned observation as still holding good; and
the distinction between the Eocene species of Sphenotrochus and the more recent representatives of the
same generic type is a result not devoid of interest for geologists as well as for zoologists.

A BRITISH FOSSIL CORALS.

not quite as much developed as the older ones, but are broad enough to reach the columella,
to which they are united, at least towards their base. The union between the septa and
the columella is not complete, but is effected by means of a double series of trabicule
extending from the inner edge of the septa, bent alternately to the right and to the left ;
so that in a vertical section of the visceral chamber a series of pores is seen along the line
of junction of each septa with the columella (fig. 1c). This mode of arrangement of the
marginal trabicule gives also to the septa, when viewed from above (fig. 1a), or by means
of a horizontal section, the appearance of bifurcation along their inner margin, and may
easily be mistaken for a disjunction of their two constituent lamine, an error which has been
committed by Goldfuss and by ourselves in our first observations.

Spyhenotrochus intermedius is the largest known species of this genus ; sometimes, how-
ever, S. Mfi/letianus and S. granulatus are almost as long. Its usual length is about three
lines, but there are individuals half an inch long. The long axis of the calice is about two
nes and a half.

Mr. Searles Wood, to whose kindness we are indebted for the specimens here described,
has collected an interesting series of these Corals, showing the changes of form which they
experience before arriving at the adult state, and has thus enabled us to study their mode
of growth, as we had already done for Fungia in a preceding memoir.’ We have not met
with any of these young Turbinolidee with only six septa and the same number of cost ,
the youngest in Mr. Searles Wood’s collection (fig. le) has twelve well-marked coste,
distinct from the top to the bottom of the corallum; but the six primary septa are the
only ones which are pretty well developed, and those of the second cyclum are still in a
rudimentary state. There is no trace of the columella, which appears at a later period
and the general form of the corallum is almost cylindrical; its height is then not more
than two thirds of a line, and its calice is circular. The base of the corallum 1s
circular ; it is truncate, but not spread out, and its adherence must have been of very short
duration.

Before the tertiary costae make their appearance, the calice begins to enlarge in one
direction more than in the other, so as to assume an oval form; a slight coarctation
becomes visible towards the middle of the corallum, its uoper part swells out laterally, and
the peduncle enlarges and becomes smooth. Soon after this the tertiary costs begin to be
formed (fig. 1/), and the calice becomes completely elliptical, but is still quite horizontal.
The coarctation above the peduncle still exists, and we at first supposed that the upper
part of the corallum became free by rupture, as is the case in Flabellum;? but the series
of specimens collected by Mr. Searles Wood shows that such is not the case, and that the
peduncle does not lose its vitality, but is gradually absorbed. Its truncate extremity is first

! Observations sur la Structure et le Mode de Développement des Polypiers, Ann. des Se. Nat., 3° série,
vol. ix, p. 76, tab. vi.
2 Loe. cit.

CORALS OF THE CRAG. 5

cicatrized, and becomes rounded, at the same time that it expands laterally, as does the
rest of the corallum, which ceases to be cylindrical, and assumes a compressed form
(figs. 1g and1/). Soon after the peduncle begins to become thin, and to shorten (fig. 17) ;
the absorption continues till it disappears completely, and the under edge of the corallum
becomes long and obtuse. While these modifications are gomg on, other changes are
produced in the internal structure of the corallum. As soon as the tertiary septa appear,
the columella begins to rise, and the primary and the secondary septa, which have become
rather broad, give off some spiniform trabiculz, that unite with the columella. The
simultaneous development of the twelve tertiary coste also determines considerable change
in the general form of the corallum; the calice, instead of beimg horizontal, becomes
arched (figs. ly, 14, 17), and the sides of the wall corresponding to the long axis of the calice
not having yet expanded towards the basis, the corallum has the form of a small battledore ;
but when the tertiary coste increase in size, the convexity of the calice diminishes, and
the base of the corallum spreads out, till it assumes the form of a broad, obtuse wedge
(figs. 1, 1a), which it retains in the adult state.

The Sphenotrochus intermedius is easily distmguished from Sphenotrochus crispus,
S. miztus, S. pulchellus, S. granulosus, and 8. semigranosus (species which all belong to the
Eocene period), by the costee beg smooth, and not formed by a series of large granules. It
resembles S. granulosus by its general form, and 8. crispus by its calice. We are acquainted
with only three other species, which have also smooth costze, and have often been confounded
with §. intermedius. One of these levicostate species is the S. Andrewianus, which lives
on the coasts of Cornwall and of the Isle of Arran, but is easily distinguished by its narrow
subconical base, and the slight elongation of its calice, the two diameters of which are as
100:120. The second levicostate species, which we designate by the name of Spheno-
trochus Remeri,: differs also from S. intermedius by its narrow base. ‘The third species,
S. Milletianus, bears great resemblance to the latter, and belongs to strata occupying the
same geological formation, a circumstance that has also contributed to create confusion
between them. But the S. I///etianus found in the Faluns of Anjou is characterised by
its lateral costee being much less prominent, and its base being more rounded and less
compressed than in the S. zztermedius.

This fossil is common in the Coralline Crag, and the Red Crag at Sutton. , We have
ascertained its identity with the species found in the Crag of Antwerp, by comparing it
with the specimens belonging to the collection of M. H. Nyst, at Louvain, and with that
of Goldfuss, in the Poppelsdorf Museum, at Bonn. Specimens of this species exist in the

1 This undescribed species has most of the characters of S. méixtus, but the coste are all similar and
smooth. The lateral ones are not notably larger than the others, and those adjacent are slightly curved
near their lower end, and sometimes interrupted. The primary and secondary septa are equal, and those
of the third cyclum are narrow; all are thick towards the outer edge, and but slightly granulate. Calice
twice as long as it is broad. Length two lines; breadth one line and a half; thickness one line. A fossil
of the Miocene strata of Cassel and Hildesheim, belonging to the Museum of Bonn. M. Nyst possesses a

specimen of the same species found in the Crag of Antwerp.

6 BRITISH FOSSIL CORALS.

collections of the Geological Society of London, and of MM. Searles Wood, Bowerbank,
and Frederick Edwards, in London; of the Museum of Natural History, and of
MM. d’Archiac, Michelin, and Milne Edwards, in Paris; M. Nyst, at Louvain; M. de
Koninck, at Liége, &c.

Genus FLABELLUM (p. xvii).

1. Fuasettum Woopit. Tab. I, figs. 2, 2a, 26.

Funeia sEMILUNATA,! Searles Wood, Ann. and Mag of Nat. Hist., vol. xiii, p. 12, 1844.
FLABELLUM Wooptl, Milne Edwards and J. Haime, Monogr. des Turbinolides, Ann. des Sc.
Nat., 3™° serie, vol. ix, p. 267, 1848.

Corallum simple, erect, rather short, much compressed, especially towards its base,
cuneiform, subdeltoid, with a peduncle short and rather thick, and lateral edges straight,
and diverging at an angle of rather less than 90°. All the coste, even the lateral ones,
simple, flat, equal, indistinct, and crossed by scarcely developed ruge and slight folds
of the epitheca, which is very thin. The surface of the wall is also marked by small longi-
tudinal sulci, corresponding to the outer edge of the septa; those referable to the small
septa but slightly marked.

Calice having the form of a very long ellipse, and rather arched. In one specimen
the proportion of its two axes was as 100 : 280, and in another as 100 : 300; the ex-
tremities of the ellipse corresponding to the great axis are obtuse, and on a level rather
lower than that of the small axis. The fossw/a is long, narrow, and deep.

The columella represented only by few large granule adhering to the inner edge of
the septa, and assuming the form of short, thick trabicule.

The septa constitute five complete cycla, very well developed, and a sixth cyclum
imcomplete, more or less rudimentary in some parts, but most apparent in the systems
situated near the long axis of the calice. The septa of the first three cycla are nearly
of the same size, and the septal apparatus is therefore divided into twenty-four groups or
apparent systems, containing each seven septa, or only five, as is often the case when those
of the sixth cyclum are missing in half of these groups. In general, these minor septa are
most developed in the half of the lateral groups adjoiming the extremities of the long axis
of the calice, and at the same time the septa of the fourth cyclum enlarge in these groups
so as to resembie the neighbouring ones of superior orders, and produce an appearance of
there being twenty-six or twenty-eight systems ; but in these lateral groups the number of
septal elements never exceeds three.

The septa are straight, thin, closely set, and do not rise quite so high as the mural

1 The Fungia semilunata of Lamarck, to which this fossil was referred by Mr. Searles Wood, belongs to
the genus Diploctenium of Goldfuss; hence the necessity of giving a new name to the above-mentioned
species. (See our Monograph of Astreide, Ann. des Sc. Nat., 3"° série, vol. x, p. 248.)

CORALS OF THE CRAG. (|

epitheca; their upper edge is slightly sinuous, and their surface covered with projecting
granule of various sizes, disposed rather regularly in rows nearly parallel to the upper
edge. These granule are much larger along the inner and inferior part of the edge of
the septa of the superior orders, where they assume the appearance of alternate trabicul
or spines. It is also to be noted that the principal septa are slightly emarginated near
the border of the calice, and that their free edge is thin and arched above, thick,
subflexuous, and obliquely truncate towards the columella. A horizontal section of the
corallum, made a little below the edge of the calice, shows the thickness of the walls,
and of the inner part of the large septa; it also renders evident the bifoliate structure of
these septa. Height twelv; lines; long axis of the calice from twelve to sixteen lines ;
the short axis from four to six lines.

The genus Flabellum contains a great number of species, and has been subdivided
into three sections, according to the state of the basis of the corallum, which is sometimes
pedicellate or truncate, and in others widely adherent. The Madbellum Woodii is easily
distinguished from the fixed /7abel/um and the truncated Fluéellum, by the permanence of its
narrow peduncle, and differs from most of the pedicellated F7adel/a by its simple non-
cristate, non-spinous cost. Seven species, F. Gallapagense, P. Michelinu, F. Thouarsia,
F. cuneatum, F. subturbinatum, F. majus, and F. Sinense, have the same character ; but
F. subturbinatum and F. Michelinii are recognisable by their horizontal calice and their
lateral costz, almost vertical. /. Gallapagense also resembles F. Woodi by the rudimentary
state of its columella, but is of a more elongated form, and is much less compressed laterally.
F. cuneatum and F. majus are still nearer allied to #. Woodi, their characters, however,
are not yet completely known; but the first of these fossil species has the septa much
thicker than in the above-described Coral, and Flabellum majus is remarkable by its great
size, its highly-compressed calice, and the peculiar structure of its principal septa.'

The Flasellum Woodii has been found in the Coralline Crag at Iken, and appears to be
very rare; for in 1844, when Mr. Searles Wood published his ‘ Catalogue of the Zoophytes
of the Crag,’ only two specimens, one belonging to Mr. Bunbury, and the other to Mr. W.
Colchester, were known, and we believe that since that time only two more specimens,
now in the possession of Mr. Searles Wood, have been found. Those figured and
described in this Monograph were communicated to us by Mr. Searles Wood.

' See our Monograph of Turbinolide, loc. cit., p. 2611.

8 BRITISH FOSSIL CORALS.

Family ASTREID/AS (p. xxii).
Tribe ASTREIN Ai (p xxxi).

Genus CRYPTANGIA (p. xliv).

1. Crypraneia Woopi. ‘Tab. I, figs. 4, 4a, 46, 4c, 4d, 4e.
Ciapocora caRiosa, Lonsdale; in Searles Wood’s Catal. Ann. of Nat. Hist., vol. xiii,
p- xu, 1844.!
Crypraneia Woopit, Milne Edwards and J. Haime, Mem. sur les Astreides, Comptes rend.
de l Acad. des Sciences, vol. xxvii, p. 496, 1848.

This singular fossil Coral is always found immersed in a mass of Cellepora, a peculiarity
which is also met with in another species of the same genus, belonging to the Faluns of
Touraine. At first sight, the vesicular mass formed by these Bryzoa may easily be mistaken
for a cellular epithecal coenenchyma, resembling that of Sarcinula ; but an attentive examina-
tion of the cells will lead to a recognition of their real nature, and similar masses of Cellepora,
not contaming any Cryptangia, are often found in the same localities. It is however
remarkable, that Corals of this genus should never be found adhering to other extraneous
bodies, and should always take up their abode on a cluster of Cellepora, which, increasing
as they themselves grow up, imbeds them so completely, that the calices alone remain
free on the surface of the common mass.

The mode of multiplication of Cryptangia is also worthy of notice. These Corals
always form clusters, and must be produced by gemmiferous stolons, but the radiciform
expansions from which they must proceed do not become sclerenchymatous, and leave
little or no trace of their existence ; so that when the soft parts are destroyed, as is always
the case in fossils, the different corallites appear to be quite independent, and would be
free, were it not for the extraneous cellular mass in which they are so deeply immersed.
It is therefore easy to perceive that these Corals differ widely from Cladocora, to which
they were referred by Mr. Lonsdale, and are equally distinct from the generic forms to
which the name of Lithodendron, applied by M. Michelin to the Touraine species, had
been previously given. ‘They are nearly allied to the Astreine reptantes, for which we have
established the genera Angia and Rhizangia, yut must constitute a separate generic group,
which we have proposed calling Cryptangia.

1 The Madrepora cariosa of Goldfuss, to which this fossil was referred by the above-mentioned author,
is a true Madrepora, and neither the one nor the other can be placed in Ehrenberg’s genus Cladocora,
The typic specimen of M. cariosa, figured aud described by Goldfuss, is preserved in the Museum of Bonn,
where it was attentively examined by one of us; it is a fossil of the Parisian basin, having a spongy
coenenchyma, and the visceral cavity of the corallites divided into two parts in consequence of the great

development of two opposite primary septa.

CORALS OF THE CRAG. 9

The corallites penetrate almost perpendicularly to the surface of the celleporous mass,
and, when isolated from this extraneous body, present the appearance of small, subturbinate
cylinders, the walls of which are covered with a thick epitheca; there is no trace of costz
visible, and the epitheca forms round the calice a small exsert rim. The calice is
circular, and its fossula large, but not deep. The columella is well developed, papillose,
and not projecting, nor is it placed exactly in the axis of the visceral chamber, the septa
being more developed on one side of the corallite than on the other. The septa of
different orders are nearly equal in size, and do not form well-characterised systems ; they
vary in number from sixteen to twenty, and consequently must belong to three cycla, the
first two of which are probably complete, and the third developed only in two or four of
the six systems normal in all Astreide. It is also te be noted, that all these septa are
very thin excepting near the wall, closely set, slightly bent inwardly, and terminated by an
oblique edge, armed all along with strong dentations, the size of which increases towards
the columella. A few large granule are seen on the lateral surfaces of the septa, and the
loculi are divided by very thin dissepiments, placed at a distance of about two thirds of a
line from each other.

The length of these corallites, when adult, is about four lines; the diameter of the
calice, one line and a half; and the depth of the fossula, two lines.

Cryptangia parasita* of the Faluns of Touraie, is very nearly allied to the above-
described species, but differs from it by the small dimensions of its calices, and the
constant existence of eight principal septa.

Cryptangia Woodii is found in a good state of preservation in the Coralline Crag at
Ramsholt. Specimens which appear to belong to the same species, but are not well
preserved, are met with in the Red Crag of Sutton.

These fossils are to be seen in the collections of the Geological Society of London,
and of Messrs. Searles Wood, Bowerbank, D’Archiac, and Milne Edwards.

Family EUPSAMMID/: (p. li).

Genus BALANOPHYLLIA (p. li).

BALANOPHYLLIA caLycuLus. ‘Tab. I, figs. 3, 3a, 36, 3c, 3d.

R. C. Taylor, Mag. of Nat. Hist. vol. ili, p. 272, fig. D, 1830. (Very
rough figure.)
BALANOPHYLLIA CALYcULUS, Searles Wood, Aun. of Nat. Hist., vol. xiii, p. 12, 1844.
— Milne Edwards and Jules Haime, Annales des Scien. Nat.,
vol. x, p. 84, 1848.

Corallum simple, cylindrico-turbinate, adherent by a large basal surface, erect, and in
general not very tall. The walls, of a spongy tissue and rather thin, are covered in most

1 Lithodendron parasitum Michelin Icon. Zooph., pl. lxxix, fig. 3.
2

10 BRITISH FOSSIL CORALS.

parts by a pellicular epitheca, which Mr. Searles Wood has designated by the name of
pertostracum, and presents some slight transverse folds. In the parts where the epitheca
is worn off, the coste become visible. These are narrow, equal, closely set, and composed
of a single series of indistinct, obtuse granule. The intercostal spaces present. a series of
small mural perforations, disposed with some regularity. None of the numerous specimens
of this fossil which we have examined had the calice well preserved, and consequently
we have not been able to ascertain as yet whether its margin is crenulated or entire, the
fossula deep or shallow, and the columella projecting or not; but it is evident that the
calice must be sub-circular, or slightly elliptic, with its two axes in the proportion of
100 : 120, and that the columella is spongy, not greatly developed, and spread in the
direction of the long axis of the calice.

The septa are well developed, and always form five cycla, but do not appear ever to
constitute a sixth cyclum. ‘The mode of arrangement of these laminz, which is charac-
teristic in the family of Eupsammid, is very evident in this species: the septa of the
first four cycla are straight, but those of the fifth cyclum deviate a great deal from the
direction of the radu of the circle represented by the calice, and are bent. In this last
cyclum the septa of the sixth order are placed very close to the primary septa, and are
united to them to a certain extent, near the wall, but diverge strongly from them as they
advance towards the centre of the visceral chamber, and join the ternary septa near the
columella; those of the seventh order are disposed in the same way near the secondary
septa, and are also united to the ternary septa by their inner edge, but do not advance
quite so near the centre of the visceral chamber ; the septa of the eighth and ninth orders,
which complete the fifth cyclum, are smaller than the preceding ones, and are strongly
bent, so as to join the septa of the sixth and seventh orders; and the septa of the fourth
and fifth orders, which constitute the fourth cyclum, remain free, and advance in the
middle of the sort of irregularly circular depressed area, formed by the coalescence of the
septa of the eighth and ninth orders with those of the sixth and seventh. All the septa
are very closely set and thin, but the primary and secondary ones enlarge a little towards
their inner edge, and are almost equally developed, so that the adult corallum assumes the
appearance of having twelve septal systems instead of six, which is the real number. We
must also add, that the laminz constituting all these septa are cribriform, and not very
granulate.

The length of this corallum is commonly about eight lines, but the individual repre-
sented by fig. 3 is more than twice as tall, without being broader than usual. ‘The calice
is in general about seven or eight lines broad in one direction, and six lines in the other.

The greater development of the epitheca might suffice to distinguish Balanophyllia
calyculus from all the other species belonging to the same genus, but it differs also from
B. pralonga’* (a fossil species belonging to the Miocene deposits of Turin) by its broad basis,

! Turbinolia prelonga, Michelin, Icon., pl. ix, fig. 1.

CORALS OF THE CRAG.

whereas the B. prelonga and the B. Gravesii! of the Paris basin have a narrow peduncle ;
from B. cylindrica” (a Miocene species found at Turin and Verona), B. geniculata® (a fossil
belonging to the Nummulitic formation of Port des Basques), and B. Cumingii* (a recent
species from the Philippine Islands), by the existence of the fifth cyclum of septa ; and from
B. tenwstriata (fossil of the Paris basin), B. desmophyllum*® (fossil of the London Clay),
B. italica® (fossil of the Pliocene deposits of Asti), and B. Bairdiana (a recent species),
by its general form, and the slight elongation of its calice. The species which it most
resembles is B. verrucaria, which exists at the present period on the coast of Corsica ;
but in the latter the columella is less developed, and the arrangement of the septa of the
last orders is less regular.’

Balanophyliia calyculus is common in the Red Crag of Sutton, but has not, to our
knowledge, been met with in other localities. Mr. Searles Wood considered it as identical
with some fossil Coral found in the Faluns of Touraine, but the latter are young specimens
of the Dendrophyllia amica; they resemble B. calyculus by their epitheca, but are easily
recognisable by the mode of arrangement of their septa, and their multiplication by
gemmation when in the adult state.®

We have examined numerous specimens of this species in the collections of the
Geological Society of London, of Messrs. Bowerbank and Searles Wood, of the Museum of
Natural History, and of MM. D’Archiac, Michelin, and Milne Edwards, in Paris.

! Turbinolia Gravesii, ibid., pl. xliii, fig. 7.

® Turbinolia cylindrica, Michelin, ibid. pl. viii, fig. 19.

3 Caryophyllia geniculata d Archiac, Mem. Soc. Géol. France, 2d series, vol. ii, pl. vil, fig. 7 a.
4 See our Monograph of Eupsammidz, Ann. des Sc. Nat., 3™ série, vol. x, pl. i, fig. 8.

5 See tab. vi, fig. 1.

® Caryophillia italica, Michelin, Icon., pl. ix, fig. 19.

7 See our Monograph of Eupsammide, Ann. des Sc. Nat., 3° serie, vol. x, p. 85, tab. i, fig. 6.
8 Loe. cit., tab. i, fig. 9.

12 BRITISH FOSSIL CORALS.

CHAPTER II.
CORALS OF THE LONDON CLAY.

Tue Eocene deposits, known by the name of London Clay, contain various Corals,
most of which belong to the two subordinate forms predominant among the Polypi of the
present period, Zoantharia aporosa and Zoantharia perforata; but none of these organic
remains can be considered as appertaining to species now in existence, or even to those
found in the more recent tertiary formations. The general aspect of this portion of the
fossil Fauna of England resembles very much that of the Corals imbedded in the
“‘Calcaire grossier” of the Parisian basin. Some species, such as Zurbinolia sulcata,
Stylocema emarciata, Stylocenia monticularia, and Holarea Parisiensis, are common to
both these localities ; but most of those found in the Eocene strata of the environs of Paris
have not been met with in the London Clay, and many of the Corals belonging to these
last-mentioned deposits have not been discovered elsewhere. Thus the London Clay
appears not to contain any Milleporide, Madreporide, or Lophoserine, families which have
various representatives in the Fauna of the Calcaire grossier, and the only Parisian fossil
Coral referable to the order of Alcyonaria is a Distichopora; whereas both Pennatulide
and Gorgonide have been met with in the London Clay. At the present period similar
differences exist at small distances in the same zoological region, and appear to depend
principally on the depth of the sea and the nature of the bottom; by analogy we are
therefore led to suppose that in the Eocene marine Fauna they are only indicative of some
such local peculiarities. Indeed, most of the Corals of the London Clay belong to Polypi
nearly allied to species which are now found in very deep water, and seem to be particularly
organized for living on a loose, muddy, or sandy ground; whereas many of the fossil
Corals of the Calcaire grossier resemble those which now inhabit rocky shores, and are
seen very near the surface of the sea.

The principal localities from which our London Clay Corals have been obtained are,
Haverstock Hill, Highgate and Holloway, near London; Barton, Sheppy, Bracklesham
Bay, on the coast of Sussex, and Alum Bay (Isle of Wight); most of the species were
found by Mr. Bowerbank, Mr. Frederick Edwards, and Mr. Frederick Dixon, to whom
we are indebted for the specimens figured in this Monograph.

CORALS OF THE LONDON CLAY.

Orpzr 1—ZOANTHARIA.
Family TURBINOLID& (p. xi).

Tribe TURBINOLIN & (p. xvi).

1. Genus TURBINOLIA (p. xvi).

1. Turpinonia suucata. Tab. III, figs. 3, 3a, 34, 3c.

TURBINOLITE DE DEUXIEME GRANDEUR, Cuvier and Alex. Brongniart, Géogr. Minéral. des
Environs de Paris, pl. ii, fig. 3, 1808.

TuRBILONIA suLcaTa, Lamarck, Hist. des An. sans Vert., t. ii, p. 231, 1816; 2d edit., p. 361.

- _ Lamouroux, Expos. méth. des Genres de Polypiers, p. 51, tab. Ixxiy,
figs. 18-21, 1821. (Very bad figures.)

— — Cuvier and Brongniart, Descript. Geol. des Environs de Paris, p. 33,
tab. viii, fig. 3, 1822.

— — Deslongchamps, Encyclop. méthod. Zooph., p. 761, 1824.

— — Goldfuss, Petref. Germ., vol. i, p. 51, tab. xv, fig. 3, 1826. (This
figure is very good, excepting that the columella is not conical
enough.)

— — Fleming, Hist. of British Animals, p. 510, 1828.

— = Defrance, Dict. des Scien. Nat., vol. lvi, p. 93, 1828. (The Coral
figured under this name in the Atlas of the Dictionnaire des Sci-
ences Naturelles, tab. xxxvi, fig. 2, andin the Manuel d’ Actinologie,
by M. de Blainville, is not a Turbinolia, and appears to belong to
the genus Trochocyathus.)

— _ Holl, Handb. der Petref., p. 415, 1829.

— — Bronn, Lethea Geognostica, vol. ii, p. 899, tab. xxxvi, fig. 4, 1838.
(This figure is good, but the columella is rather too thick.)

—~ — Nyst, Descript. des Coquilles et Polypiers fossiles de la Belgique,
tab. xlvii, fig. 11, 1843. (This figure is copied from Goldfuss;
the description is referable to the Turbinolia Nystiana.)

— — Michelin, \econogr. Zooph., p. 151, pl. xliii, fig. 4, 1844.

_ — Graves, Topogr. Géogn. de I’Oise, p. 701, 1847.

= —_ Milne Edwards and Jules Haime, Annales des Scien. Nat., 3™° série,
vol. ix, p. 236, 1848.

This corallum has the form of a cylindroid, elongated cone, and is not contracted just
above its basis, nor inflated near the calice (figs. 3, 3 4); sometimes only the cone is
somewhat shorter in proportion to its length (fig. 3a). ‘The coste are very thin, sharp,
straight, and very prominent from top to bottom, but particularly so near the basis of the
corallum. ‘The secondary coste are nearly as long as the primary ones; they do not,
however, originate quite at the same level. ‘The tertiary costz begin to appear about half
way up the wall in young specimens, and occupy two thirds of the height of the Coral in

14 BRITISH FOSSIL, CORALS.

the adult state. Tue imtercostal grooves are deep and broad; near the calice a small
longitudinal line is visible in each of them, and indicates the existence of a fourth cyclum
of rudimentary cost, which do not correspond to any of the septa on the inner side of
the wall. ‘These vertical furrows also present a double series of small dimples, which are
prolonged laterally on the sides of the costa, so as to constitute a sort of transverse fluting,
and are arranged alternately ; they are very closely set, and about fifteen occupy the space
of a line. The wall is very thin. The calive is circular, and its fossula is not very deep.
The columella is terminated by a conical, pointed apex, which rises higher than the septa,
and is delicately granulaied. The septa are thin and very exsert, but not quite so much
so as in the Zurbinolia Divonii;* their upper edge is strongly arched, and their lateral
surfaces present small granulations, which form short submarginal, radiate lines near the
apex, and are arranged in nearly horizontal rows towards the lower part of the visceral
chamber. The inner edge of the apical portion of the septa is slightly concave, and soon
becomes horizontal, so as to meet the columella, to which it unites. A projecting line
extends from each of the six primary septa up the apical portion of the columella; the
secondary septa join the columella much lower down, but they are broader than the primary
ones. The tertiary septa are narrower at the apex, and less exsert than the preceding ones ;
they converge towards the intermediate primary septa, and become united to them all along
their mner edge, at about two thirds of the breadth of the latter. The height of the
corallum is usually about three or four lies, and the diameter of the calice about one line
and a half. In young specimens the calice is larger in proportion.

This fossil is the only species belonging to the genus Turbinolia as now circumscribed,
which was known at the time when Lamarck first established the group bearing that name.
Shortly afterwards, Mr. Defrance discovered a second species, and Mr. Isaac Lea has since
then found a third. In our Monograph of Turbinolide, published about a year ago, six
species were described, and we now know double that number of true Turbinoli, but
they all belong to the same geological period, and are imbedded in Eocene deposits. They
appear to be mure abundant in England than elsewhere ; the London Clay contains eight
species, only one of which (the fossil just described) has been met with in the synchronous
formation of the Parisian basin.

Turbinolia sulcata differs from Turbinolia dispar, an from Turbinolia costata,*’ by the
number of the septa which in these two last-mentinned species form four cycla. An
additional cyclum of cost distinguishes Zurbinolia Fredericiana* from it ; in Turbinoha
Prestwichii,’ T. minor,® and T. firma,' the coste are not so thin, prominent, and wide apart
as in this species, and the last of these characters separates it also from Turbinolia pharetra®

1 See plate in, fig. 1. 2 Michelin, Icon., pl. xlii, fig. 5.

3 These species, as well as the others only quoted here, have been described at full length in our
Monograph of Turbinolidz, published in the Annales des Sciences Naturelles, 3"° série, vol. ix.

4 See pl. iii, fig. 2. 5 See tab. iii, fig. 5. © See tab. ii, fig. 5.

7 See tab. ii, fig. 4. Contrib. to Geol., tab. vi, fig. 210.

CORALS OF THE LONDON CLAY. 15

and 7. Nystiana,' to which it is, however, closely allied. The species which it resembles
most are, however, Turbinolia Dixonit, T. humilis,? and T. Bowerbankii.2 The last of these
differs from 7. sulcata by its form, which is more elongate and conical, by the thickness
of the lower part of its primary coste, and by its very slender columella. Zurbinolia
Dizonir is easily distinguished by its compressed columella, by the enlargement of its walls
near the calice, and by the great prominence of its coste. To conclude this brief com-
parison, we must add, that Zvrécnolia sulcata differs from 7. humilis by its size, by its
form, which is not near so cylindrical as in the latter, and by the normal number of its
septa.

This species is extremely abundant in certain localities of the environs of Paris, such as
Grignon, Parnes, and Auvert; it is also found in the tertiary strata of Hauteville, in
Normandy, and in the London Clay at Bracklesham Bay, but it is not common in this last-
mentioned deposit. We are indebted to Mr. Frederick Edwards and to Mr. F. Dixon for
the specimens figured in this Monograph.

2. Turpinouia Dixonu. Tab. III, figs. 1, la, 14, le, ld.

Turpinotia Dixont, Milne Edwards and J. Haime, Monogr. des Turbinolides, Ann. des

Se. Nat., 3™° série, vol. ix, p. 238, tab. iv, figs. 2, 2a, 26. 1848.4

— SULCATA, Lonsdale, in the MS. work of M. Dixon on the Chalk Formations and
Tertiary Deposits of Sussex.

Corallum slightly contracted just above its basis, and rather inflated near the calice.
Coste very thin, and projecting very much, especially towards the lower part of the wall;
those of the third cyclum beginning very near the basis, and those of the first and second
ceycla beginning almost at the same height. Intercostal furrows nearly of the same size>
very broad, and very deep; intercostal dimples very distinct, separated by small transverse
laminze, disposed as usual, in two vertical rows, and prolonged laterally, so as to produce
the appearance of transverse fluting on the sides of the coste; about ten of these dimples

1 We have given this specific name to the Turbinolia described by M. Nyst, and considered by that
author as being referable to the Turbinolia sulcata (see Coquilles et Polyp. des Ter. tert. de la Belgique,
p- 629; but not the corresponding figure, which is copied from the work of Goldfuss, and belongs to
T. sulcata). In order to facilitate the comparison between the British Turbinolia and the species found in
other countries, we think it may be useful to point out the characteristic features of the 7’. Nystiana, which
were not known to us when we published our Monograph of Turbinolidz.

Turbinolia Nystiana, nob. (7. sulcata, Nyst, loc. cit.) Corallum elongated, slightly contracted a little
above its basis, and somewhat inflated near the calice. Coste very slightly prominent, and rather thick ;
the primary and secondary ones very broad towards the basis; the dimples of the intercostal furrows very
small, but distinct, and those of one series alternating with those of the other. No rudiments of a fourth
eyclum of costz. Columella small, and almost cylindrical. Septa rather thick, slightly granulated, and
forming three complete evela. Length 33 lines; diameter of the calice 14 line. Fossil from the environs of
Brussels. (Cabinet of M. Nyst at Louvain.)

2 See tab. ui, fig. 4. 3 See tab. ii, fig. 3.

4 In fig. 2@ of this plate, the principal septa are not broad enough towards the calice, and the
concavity of their inner edge is placed rather too high.

16 BRITISH FOSSIL CORALS.

occupy a line in length. No rudiments of a fourth cyclum of costz, and a well-marked
depression in calicular edge of the wall, corresponding to each of the intercostal spaces.
Calice with a very narrow, but rather deep fossula. Colwmella compressed, arched at the
apex, granulated on the surface, reaching in general to the same height as the septa, or
even higher, and presenting, in the part where it begins to become isolated, six vertical
striee, which are in continuity with the inner edge of the six primary septa; rather lower
down, the columella is slightly contracted, and a vertical section of the corallum (fig. 16)
shows that its tissue 1s compact, and that towards the bottom of the visceral chamber it
becomes united with the septa, so as to form a solid mass. Septa thin, unequal, very
exsert, having their upper edge strongly arched near the outer margin, but concave near
the centre of the calice, slightly granulated laterally, and forming three cycla; those of the
first and the second cycla nearly of the same height, but the secondary ones much broader
at_ the apex than the primary ones, and not extending so far up the columella. The
tertiary septa much narrower and shorter than the older ones; very thin towards their
inner edge, and cemented to the primary septa, as in the preceding species (fig. 16).
Interseptal loculi large. Height of the corallum about four lines. Diameter of the calice,
twe lines and one third. The form and the proportions not differmg in the young and
in the adult specimens.

Turbinolia Dixon is the largest known species of the genus, but 7. dispar and
T. Prestwichii are almost of the same size. This species is very closely allied to 7. sulcata,
from which it differs principally by its form (rather more inflated near the calice), by its
compressed columella, by its septa being more exsert, and its costae more projecting and
more distant. ‘The breadth of the intercostal furrows distinguishes both 7. Dizonii and
T. sulcata from T. Prestwich, T. minor, T. firma, T. pharetra, T. Nystiana, and
T. Bowerbanku. The existence of only three cycla of coste does not allow of its bemg
confounded with Z. Fredericiana, T. dispar, and T. costata, and, finally, 7. humilis is
easily distinguished from it by its cylindroid form, non-compressed columella, and
glabrous septa.

Turbinolia Dixonit is a fossil very abundant in the London Clay at Bracklesham Bay,
and has probably been confounded with 7. sulcata by Mr. Fleming, and some other
geologists, who mention the latter as being found in that locality, where it appears to be
very rare. In Mr. Dixon’s work, now passing through the press, Mr. Lonsdale has also
described it as a variety of the 7. sulcata of Lamarck.

The specimens, the examination of which has enabled us to recognise this new species,
were given to us by Mr. Dixon and by Mr. Frederick Edwards.

3. Tursinoria Bowrrsanxi. ‘Tab. II, figs. 3, 3a, 36.

Corallum almost conical, rather short. Coste not very prominent ; those of the first
two cycla inflated near the basis ; the tertiary ones beginning at less than a quarter of the
distance from the basis to the calicular edge of the wall; slight rudiments of a fourth

CORALS OF THE LONDON CLAY. 17

cyclum of coste appearing near the calice, and consisting in very short, prominent, thin
lines, most developed between the primary and the tertiary septa. Intercostal furrows
rather narrow, but presenting very clearly a double row of small dimples. Calicular
fossula not deep. Columella cylindrical, prominent, and very slender in proportion to the
size of the calice. Septa very thin, exsert, rather unequal, and forming six regularly-
developed tertiary systems (fig. 34). No traces of a fourth cyclum of septa corresponding
to the rudimentary costz of the fourth cyclum. The tertiary septa joing the primary
ones very near the columella. The lateral surfaces of all the septa presenting delicate
granulations. Height two lines; diameter of the calice one line and a third.

This species bears great resemblance to Turbinolia Fredericiana ; it differs from it by
the rudimentary state of the fourth cyclum of costz, and by its slender, round columella.
The existence of well-formed intercostal dimples distinguishes 1¢ from TZ: minor, T. costata,
T. Prestwichui, and T. firma; the coste are much less prominent than in Z. sulcata,
and 7. Dizonit, from which this Coral may also be distinguished by its form; the coste
are thinner than in 7. pharetra and T. Dizonii, and the complete development of its six
systems of septa does not admit of its being confounded with 7. humilis.

We have seen but one specimen of this species ; it was found at Barton, and belongs
to the collection of the fossils of the London Clay formed by Mr. Frederick Edwards. We
have dedicated it to our friend Mr. J. S. Bowerbank, whose active researches have much
contributed to the extension of our knowledge relative to this portion of British
palzontology.

4. Tursinonia Frevericiana. ‘Tab. III, figs. 2, 2a, 24.

Corallum of a regular conical form, not much elongated, and rather broad towards the
calice. Coste numerous, forming four cycla, closely set, unequal, and projecting very
little; the secondary ones beginning a little above those of the first cyclum, but very near
the basis of the corallum, and being, as well as the former, much thicker near their lower
end than higher up, where they become very delicate (fig. 2a). The tertiary cost begin
also at a short distance from the basis, but those of the fourth cyclum appear only in the
upper half of the corallum; they are also rather thinner than the others. The intercostal
furrows very narrow, and not very deep; the mural dimples not very apparent, small,
closely set, and forming towards the calice, if not from top to bottom, only a single series
in each intercostal furrow. Calicular fossula very narrow and shallow. Columella thick,
compressed, granulated, rising higher than the septa, and presenting well-marked pro-
longations of the principal septa. Three cycla of septa, and no vestiges of a fourth
cyclum corresponding to the quaternary costz (fig. 24). The septa are much like those
of the two preceding species, but they are a little thicker, and not so exsert ; the primary
ones are, as usual, narrower than the secondary ones, and these reach higher up along the
columella; the tertiary septa are small, and join the primary ones, but appear to be

3

18 BRITISH FOSSIL CORALS.

cemented to them. The sides of all the septa present granulations arranged in radiate
lines, but not very prominent. Height of the corallum two lines and a half; diameter
of the calice one line and a third.

Mr. Frederick Edwards, to whom we dedicate this species, has submitted to our investi-
gation a series of young individuals, showing the changes of form produced by age. The
young Corals are rather shorter in proportion, to their breadth, than the adult ones, and
consequently never resemble Zurbinolia humilis, whatever their size may be, for the latter
species is always much more cylindrical. ‘he tertiary costee make their appearance in
T. Fredericiana when very young, but those of the fourth cyclum exist only in individuals
that are nearly adult,

These quaternary coste, occupying at least one third of the height of the corallum,
and not corresponding to any rudiments of septa, distinguish Zwrbznolia Fredericiana
from all the other species of the same genus; in some others, such as Z. sudcata and
T. Bowerbanki, the rudiments of similar coste can be seen with the help of a strong lens,
but these intercostal limes never become cristiform, as is the case here. The great
development of quaternary costee and the general form of the corallum make this species
have some resemblance to Zurbinolia dispar; but in the latter, as well as in 7: costata, there
is always a fourth cyclum of septa corresponding to the last cyclum of coste. It is to
T. Bowerbanku that 7. Fredericiana approximates most; but in the former the columella
is perfectly cylindrical and extremely slender, whereas in the latter it is large and
compressed.

Turbinolia Fredericiana has as yet been found only in the London Clay, at Barton,

and the specimen figured in this Monograph belongs to the collection of Mr. Frederick
Edwards.

4. TurBinotia HuMiLis. ‘ab. II, figs. 4, 4 a, 4 6.

This little Turbimolia is of a much more cylindroid form than preceding species, and is
not so slender at its basis. The coste are thin, prominent, and not closely set; the
secondary ones begin very near the basis, and those of the third cyclum at about a quarter
of the way up the wall. The intercostal furrows are broad, and present each a doubie
row of small dimples, separated by transverse or oblique bars (fig. 4 a). The columella
is prominent, round, and conical. ‘The septa belong to three cycla, the last of which is
always incomplete, and is wanting in two of the systems ;! but it is nevertheless evident
that the number of systems is as usual six, and not five, as would at first appear, for the
secondary cost corres;:-uding to the two incomplete systems begin near the basis of the
corallum, at the same level as those of the other systems, and are as much developed as
these, whereas they would have been much shorter, and would have began much higher

' By an inadvertency of our artist, the third cyclum is represented in fig. 4 as if it were perfect ;
but the specimen did not in reality present tertiary septa in more than four of the systems.

CORALS OF THE LONDON CLAY. 19

up, if they had corresponded to septa belonging to the third cyclum. All the septa are
very thin, almost glabrous, exsert, and terminated by a regularly arched apex, rismg more
or less, according to the cyclum to which they belong. Height not quite a line and a half;
diameter of the calice two thirds of a line.

This Coral is one of the smallest of the genus Turbinolia, and is mdeed usually even
smaller than the species designated by the name of Zuréznolia minor, for which it may very
easily be mistaken ; its characteristic features can only be seen with the help of a lens,
but when sufficiently -uagnified, the appearance of its walls will make it immediately
recognisable ; for in 7. minor the coste are very thick, crenulated laterally, and the imter-
costal furrows do not present any dimples, whereas in 7. Awmzlis these dimples are well
marked, and the coste are thm. These two species are the only ones of the genus that
have apparently but five tertiary systems, and their diminutive size contributes also to
make them not easily recognisable.

Turbinolia humilis is found in the London Clay at Barton, where it appears to be
abundant. We are indebted to Mr. F. Dixon and Mr. Frederick Edwards for the
specimens in our possession.

6. Tursinomra minor. Tab. II, figs. 5, 5a, 50.

TURBINOLIA MINOR, Milne Edwards and J. Haime, Annales des Sc. Nat., 3™ série, vol. ix,
p- 239, 1848.

Corallum of a cylindrico-conical form, rather short, and very obtuse at its basis. Coste
very thick, closely set, and not very prominent ; those of the first and second order par-
ticularly thick near the basis; the outer edge of all very obtusc, and their sides delicately
crenulated. Intercostal furrows very narrow, linear, and presenting no trace of the dimples,
which are so apparent in the preceding species. The form of the cost is particularly well
marked near the calice (fig. 5 4), the lateral crenations of which are sometimes so developed
near the basis, that they assume a crispate appearance ; in other specimens they are scarcely
visible, but the variations met with in the form of these parts are never such as to make
them resemble the costae of Z. humilis. Calice very deep. Columella cylindrical, slender,
and exsert. Sepfa belonging to three cycla, and appearing to form only five systems.
although there are in reality six systems as usual; only in two of these there are no
tertiary septa, and the secondary septa are of the size of the other tertiary ones, but
correspond to secondary costa, the development of which are normal (fig. 54). All the
septa are thin, exsert, and slightly granulated on their lateral surfaces; the secondary ones
are nearly as large as those of the first order, but those of the third cyclum are much
smaller. ‘This species is always remarkably small; it is not more than a line and a half
high, and two thirds of a line in diameter.

20 BRITISH FOSSIL CORALS.

Turbinoia minor differs from TP. sulcata, T. pharetra, T. Nystiana, T. Dizxonit,
T. Fredericiana, and T. humilis, by not having the intercostal furrows ornamented with a
double row of dimples, a character which in these can always be ascertained with the aid
of a good lens. The imperfect development of two of the systems of septa, and the
apparent existence of only five systems which is thus produced, is also sufficient to
distinguish 7. minor from 7. costata, T. dispar, T. Prestwichii, and T. firma.

This fossil has been found only in the London Clay, at Alum Bay, in the Isle of
Wight. The specimen figured in this Monograph belongs to the cabinet of Mr. J. S.

Bowerbank.

7. Tursinotta Firma. ‘Tab. II, figs. 4, 4a, 4 6.

Corallum subturbinate, and elongated; narrow at the basis. Coste thick, obtuse,
closely set, and prominent ; those of the first two cycla very broad below the under end of
the tertiary ones. Intercostal furrows narrow, and presenting neither mural dimples nor
well-marked lateral transverse flutings or costal crenations. Colwmel/a compressed, and not
very large. Septa rather thin, delicately granulated, and forming three complete cycla ;
the tertiary ones less developed than the secondary ones, and cemented to the primary
ones at a small distance from the columella. Height three lines and a half; diameter of
the calice, one line and a half.

Turbinoha firma differs from 7. costata, T. dispar, T. Bowerbanku, and T. Fredericiana,
by the non-existence of a fourth cyclum of more or less developed coste; from Z. minor
and 7. humilis, by the complete development of the tertiary septa in the six systems, and
from 7. sulcata, T. pharetra, 7. Nystiana, T. Dixonii, and T. humilis, by the non-existence
of dimples in the intercostal furrows. It resembles very much 7: Prestwichi, but differs
from it by its general form and by its thick obtuse coste.

We have as yet seen but one specimen of this species; it was found at Barton, and
given to us by Mr. Dixon: unluckily the artist in whose hands it was placed in order to
have it figured, has broken it so much that it is no longer recognisable.

8. Tursinomia Prestwicnit. Tab. III, figs. 5, 5a, 56.

Corallum of a cylindroid form, much elongated, and very obtuse at the basis. Coste
strong, rather thick, and very prominent, especially towards the basis; those of the third
cyclum beginning much lower down than in most species (figs. 5 a), and contributing to form
the convex star seen at the basis of the corallum (fig. 5 4). Some slight vestiges of a fourth
cyclum of costz at the bottom of the intercostal furrows near the calice. These furrows
very deep, becoming very narrow near the wall, and not presenting any mural dimples

CORALS OF THE LONDON CLAY. 21

but irregularly crenulated laterally, especially towards the basis. Height four lines ;
diameter one line and two thirds.

The only specimen of this species which we have seen belongs to the collection of
Mr. Frederick Edwards, and is so much filled up with clay at its upper end, that we have
not been able to ascertain well the form of the columella and the septa; we are,
however, inclined to think that the columella is slightly compressed, and the septa rather
thick.

Turbinoha Prestwich differs from all the preceding species by its cylindrical form
and broad convex basis; it differs also from TZ. sulcata, T. Dixonii, T. pharetra,
T. Nystiana, T. humilis, and T. Bowerbanku, by not presenting any vertical rows of
intercostal dimples; from 7. dispar, 7. costata, and T. Fiedericiana, by having only three
cycla of cost, instead of four, and from 7 minor and TZ. firma, in which the intercostal
dimples are equally wanting, by its sharp-edged costz.

This remarkable species was found at Haverstock Hill, and appears to be very scarce,
for Mr. Frederick Edwards, whose collection of London Clay Fossils is extremely rich, has
only one specimen of it, and we are not aware of its existing im the cabinet of any other
paleontologist.

SuB-FAMILY oF THE CYATHININ AZ (p. xii).

1. Genus LuprocYaATuUs (p. xiv).

Leptocyatuus ELEGANS. ‘Tab. III, figs. 6, 6a, 64, 6c.

Corallum extremely short, nearly discoidal, and presenting, in the adult state, no trace
of adherence. Coste distinct from the centre of the under part of the corallum to the
calice, strong, projecting externally, cristiform, closely set, rather unequal, separated by
rather deep radiate furrows, and rendered echinulate by the presence of a multitude of
granulations crowded together (figs. 6a, 66). Calice circular, and regularly excavated
in the centre. Colvmel/a not much developed, and delicately papillose. Septa constituting
four complete cycla, closely set, broad, projecting much above and externally; very thin
near the columella, but remarkably thick towards the circumference, and rather unequal
(fig. 6c); those of the first two cycla nearly equal, and larger than the others ; the tertiary
ones broader than those of the fourth cyclum, but not so high; all are straight, and none
adhere together by their inner edge; their sides are covered with granulations, which are
obtuse towards the circumference of the calice, but become spiniform in the inner part,
where the septa themselves are slender. Pali corresponding to all the septa (even to
those of the last cyclum, a mode of structure which is very rare), very thin, slightly
echinulated, becoming broader as they correspond to younger septa, and appearing to be
lobated, as in the genus Paracyathus. Height of the corallum, one line ; diameter, three
lines and a half.

22 BRITISH FOSSIL CORALS.

The genus Leptocyathus, which we have established for this fossil, is nearly allied to
Trochocyathus, but differs from it by its subdiscoidal form, the absence of all sign of
adhesion at the basis, and the existence of pali corresponding to all the septa. The genus
Ecmesus of Philippi’ appears to present most of the same characters, but, as far as we can
judge by the very short description, and by the rough figure given by that author, the
calice appears to be eccentric, a mode of structure which is quite exceptional, and
very remarkable.

The fossil Coral from the environs of Biaritz, mentioned by Viscount d’Archiac?
under the name of Zurbinolia atalayensis, belongs probably to the same generical division
as our Leptocyathus elegans, but differs from it by its large size, by the existence of a fifth
cyclum of septa, and by the smooth surface of the central portion of its wall.

Leptocyathus elegans was found in the London Clay, at Haverstock Hill, by Mr.
Frederick Edwards, who obligingly communicated to us the only specimen that has as yet
been seen.

2. Genus 'TROCHOCYATHUS (p. xiv).

TROCHOCYATHUS SINUOSUS.

TURBINOLIA TURBINATA (pars), Lamarck, Hist. des An. sans Vert., t. ii, p. 231, 1816.
— .... Parkinson, Organic Remains, vol. ii, tab. iv, fig. 11, 1820.
— stnvosa, Alex. Brongniart, Mém. sur les Terr. du Vicentin, p. 83, pl. vi,
fig. 17, 1823.
— — Bronn, Syst. des Urweltlichen Pflanz., tab. v, fig. 12, 1825. (Bad
figure.)
— puBIA, Defrance, Dict. des Sc. Nat., vol. lvi, p. 92, 1828.
—_— stnuosa, Bronn, Lethea Geognostica, vol. ii, p. 897, 1838.
— — Leymerie, Mém. de la Soc. Géol. de France, 2™° serie, pl. xiil,
figs. 7, 8, 1845.
— — Michelin, Icon. Zooph., p. 270, pl. lxiii, fig. 1, 1846.
TRocHOcYATHUS sINUosUS, Milne Edwards and J. Haime, Ann. des Sc. Nat., 3™° serie,
vol. ix, p. 314, 1848.

We have not met with this fossil in any collection of the British Corals, but Parkinson
has figured it in a very recognisable way, and mentions it as having been found in the
Isle of Sheppy; we must therefore recall its specific characters im this monograph,
although we deem it advisable not to have it figured from a foreign specimen.

Corallum subturbinate, rather compressed, and having its inferior extremity shghtly
curved in the direction of the small axis of the calice. Coste distinct from the basis, very
narrow, numerous, closely set, simple, unequal, delicately granulated, and projecting very
little. Cadice oval, and contracted in the middle, so as to assume the form of an 8.

1 Neues Jahrb. fiir Mineral. Geol., vol. ix, p. 665, tab. xi, fig. B 1, 1841.
2 Bulletin de la Soc. Géol. de France, 2™° série, vol. ii, p. 1010, 1847.

CORALS OF THE LONDON CLAY. 23

Columella fasciculate, with very slender elements. Septa forming six complete cycla,
closely set, very thin and broad; those of the first three cycla nearly equal. Pali rather
large, and thin, scarcely thicker than the septa, and presenting laterally spiniform granu-
lations ; those corresponding to the penultimate cyclum of septa being the most developed,
the others nearly equal.

This Coral soon acquires all its septa and its final diameter, but continues growing up,
so that it becomes sometimes very tall, without expanding proportionally ; we have seen
specimens three or four inches high, or even still longer. It has been found in the lower
tertiary deposits of several localities in the south of France and the north of Italy.

3. Genus PARACYATHUS (p. XIV).
1. Paracyatnus crassus. Tab. IV, figs. 1, la, 14, le.

Corallum subturbinate, short, fixed by a very broad basis, slightly contracted just
above the lower end, and rather inflated at the upper part. Coste well marked from top
to bottom, closely set, nearly equal in breadth, but alternately more or less prominent,
especially near the calice, and covered with very delicate granulations. Calice nearly
circular when young, but becoming soon more or less oval; fossula deep. Colwmella
concave, papillose, thick, and not distinctly separated from the inner lobes of the pali.
Septa forming four complete cycla, and an incomplete rudimentary fifth cyclum, in one half
of the systems corresponding to the long axis of the calice; closely set, straight, slightly
exsert, thin towards the centre of the visceral chamber, rather thick externally, granulated
laterally, and unequally developed according to relative age. Pali corresponding to the
septa of the first three cycla, thick, tall, strongly granulated, and denticulated along the
inner edge, which is rather oblique; those corresponding to the tertiary septa larger than
the others, and those that correspond to the primary septa being the smallest of all.
Height, five or six lines ; long axis of the calice, four lines; short axis, three lines; depth
of the fossula, three lines.

This Paracyathus is easily distinguished from the other species of the same genus by
the number of the septa, which in P. procuméens,! P. Stokesii,’ and P. Desnoyersit, form an
additional cyclum ; by the size of the pali, which are much thicker than in P. caryophyllus,
and P. drevis,* and by the lobulate edge of these same organs, and the oval form of the
calice, from P. equilamellosus, P. Pedemontanus,’ and P. Turonensis.

Paracyathus crassus has as yet been found only in the London Clay of Bracklesham
Bay, and has been communicated to us by Mr. Dixon and Mr. Frederick Edwards.

1 Milne Edwards and J. Haime, Monogr. of Turbinolide, in Ann. Sc. Nat., 3dser., vol. ix, pl. x, fig. 6.
2 Idem, loc. cit., pl. x, fig. 7. 3 See tab. iv, fig. 2. * See tab. iv, fig. 3.
5 Caryophyllia pedemontana, Michelin, Icon., pl. ix, fig. 16.

24 BRITISH FOSSIL CORALS.

2. Paracyataus Caryoruyiivs. Tab. IV, figs. 2, 2a, 26, 2c, 2d, 2e.
TURBINOLIA CARYOPHYLLUS, Lamarck. Hist. des Anim. sans Verteb., t. ii, p. 232, 1816;
2d edit., p. 362.
— — Deslongchamps. Encyclop. méthod., Zooph., p. 761, 1824.
— — Lamarck. Tableau encyclop. ev méthod. des trois Régnes,
t. ili, p. 483, fig. 3, 1827.
— — Defrance. Dict. des Sc. Nat., t. lvi, p. 92, 1828.
PARACYATHUS CARYOPHYLLUS, Milne Edwards and J. Haime. Monogr. des Turbinolides,
Ann. des Sc. Nat., 3™° série, vol. ix, p. 322, 1448.

Corallum ttrbinate, elongated, almost cylindrical in the tallest specimens, usually
straight, or very slightly curved, and adhering by a moderately developed basis. Coste
well marked, distinct from top to bottom, nearly equal, rather narrow, not much more
prominent towards the calice than near the basis, separated by broad, deep furrows, and
covered with small granulations, which exist also in the intercostal furrows (fig. 2 6).
Calice circular ; fossula not very deep. Columella concave, delicately papillose, and not
distinctly separated from the pali (fig. 2.7). Septa forming four cycla, the last of which is
wanting in half of one or two systems ; closely set, not very exsert, thin, strongly granu-
lated laterally, and rather unequal in accordance with their relative age; the primary and
secondary ones rather thick externally. Pal very thin, rather tall, lobated, with the mner
edge oblique, and gradually larger as the septa to which they correspond are younger.
Height, varying from five to eight lines; diameter of the calice, three or four lines; depth
of the fossula, one line and a half.

This fossil is in general found in a bad state of preservation, being much worn away,
with its basis obtuse, its wall almost entirely destroyed, and the visceral chamber filled
with a carboniferous substance, the black tint of which contrasts with the white colour of
the septa. Lamarck, who had only seen specimens in this state, was thus led to suppose
that the corallum was free, and to consider it as appertaming to the genus Zurbinola.
But, through the kindness of Mr. Bowerbank and Mr. Dixon, we have been enabled to
examine a great number of specimens, some of which presented a complete calice, well-
preserved costz, and a basis that had evidently been adherent, so that no uncertainty
could remain as to their belonging to our genus Paracyathus. This species differs from
most other nearly allied species by the thinness of the pali, a character which is to be seen
only in one other species; the Paracyathus brevis, from which 7. caryophyllus is easily
distinguished, by its septa being also much thinner. The number of the septa can equally
serve as a character, for in Paracyathus Stokesii, P. Desnoyersii, and P. procumbens, there
is a cyclum more than in the species here described.

Paracyathus caryophyllus is avery common fossil in the London Clay at the Isle of
Sheppy; specimens of it are preserved in the collections belonging to the Geological
Society of London, Mr. Bowerbank, Mr. Dixon, Mr. Frederick Edwards, the Museum of
Paris, and M. Milne Edwards.

CORALS OF THE LONDON CLAY.

vw
Or

3. PAaRACYATHUS BREVIS, Tab. IV, figs. 3, 3a, 34, 3c.

PARACYATHUS BREVIS, Milne Edwards and J. Haime. Ann. des Scien. Nat., 37° série,
vol. ix, p. 323, 1848.

The fossil remains of this species which we have had an opportunity of examining, are
all more or less imperfect, and could not give us a complete knowledge of its character,
but are sufficient to show that it belongs to the genus Paracyathus, and differs from all
the other species of the same group. The general form of the corallum appears to be
usually subturbinate and short (as in fig. 3); but if, as we have some reason to think, the
natural interior cast represented in fig. 3c¢ belongs to this species, the proportions of
height and breadth must be very variable. The coste are closely set, nearly equal, thick,
and covered with dense granulations. The ca/ice is circular, and the fossula deep. The
columella is concave, large, and delicately papillose. The septa are but very slightly exsert,
closely set, thin towards the centre, and very thick towards the outer edge, strongly
granulated laterally, and almost equally developed. The pal: correspond to the septa of
the first three cycla, and are very thin, lobulated, and, as usual, developed in an inverse
ratio with the septa, in the prolongation of which they are placed. Height, from four to
seven lines; diameter of the calice, six lines; depth of the fossula, one line and a half.

The existence of only four cycla of septa distinguishes this species from Paracyathus
Stokesit, P. Desnoyersii, and P. procumbens, in which there are five of these cycla; the
tenuity of the pali distinguishes it from P. equilamellosus, P. Pedemontanus, P. Turonensis,
and P. crassus ; it is nearest allied to P. caryophyllus, in which the pali are also very thin,
and lobulated, but in the latter the septa are much thinner, and the general form is very
different.

Paracyathus brevis is found at the Isle of Sheppy, and the specimens figured in this
Monograph belong to the cabinet of Mr. Bowerbank.

ABERRANT Group oF THE PPEUDOTURBINOLID/ (p. xix).

Genus DasMia (p. xix).
Dasmia Sowersyi. Tab. IV, figs. 4, 4a, 4 6.
DEsMOPHYLLUM, J. Decarle Sowerby. Trans of the Geol. Soc. of London, vol. v, p. 136,
tab. viii, fig. 1, 1834.

Dasm1a Sowersyi, Milne Edwards and J. Haime. Ann. des Sc. Nat., 3™° serie, vol. ix,
p- 329, tab. vii, figs. 8, 8a, 1848.

Corallum subturbinate, straight, or slightly curved, and subpedicellate or adherent by
a very narrow basis. Coste extremely broad, separated by deep, narrow furrows, obtuse
A

26 BRITISH FOSSIL CORALS.

towards the basis, rather prominent; and subcristate near the calice, covered with
eranulations, which become much larger towards the calice, and varying in number (17 in
one specimen, 18 in another, and 22 ma third); about two thirds of them begin at the basis
of the corallum, and the others about half way up towards the calice, but all are of the same
breadth ; the position of these younger costz does not appear to be constant, for some are
separated by three longer ones, and others by two, or only one; in general, however, two
long ones are placed between two short ones, so that the latter are only about half as nume-
rous as the former. The calice is nearly circular, or slightly elliptical, and the fossula appears
to be deep; we are also inclined to think that there is no columella, and that the septa are
free all along their inner edge, but the calice being clogged up with carboniferous matter in
all the specimens that we have seen, we have not been able to determine these points with
any degree of certainty. The mode of arrangement of the septa is quite abnormal ; three
vertical plates advance from each of the coste towards the centre of the visceral chamber ;
they are all extremely thin, broad, somewhat flexuous, free from all adherence among
themselves, and rendered echinulate laterally by a few prominent granulations ; the plate
placed in the middle of each of these groups is rather thicker than the others, and the
space existing between it and the latter is rather larger than that comprised between the
lateral laminee of two neighbouring groups. Height of the corallum, about four lines;
iong axis of the calice, three lines and a half; short axis, two lines and a half; breadth of
the coste, more than half a line.

The three specimens of this species, from which we have drawn up the preceding
description, belong to Mr. Bowerbank’s paleontological collection, and were found at
Highgate; Mr. Prestwich has met with it also at Clarendon Hill.’

Dasmia Sowerby: is the only known species of this genus which by its general
characters appears to be closely allied to the family of Turbinolide, but differs from it,
and even from all the other Zoantharia, by the abnormal structure of the septal apparatus :
when our attention was first called to this pomt, we endeavoured to explain the mode of
radiation of the calice by supposing that each of the laminz corresponding to the middle
of the coste belonged to one cyclum, and that. the two lateral lamin of two neighbouring
groups, corresponding to the two sides of each intercostal furrow, represented the two
halves of septa belonging to another cyclum ;” the slight difference in the thickness of the
middle and the lateral laminz, as well as the facility with which the two constituent plates
of the septa separate from each other in some Corals, had induced us to admit that this
structure was only an exaggerated form of that which is frequently met with in certain
‘Turbinolidee, in many species of Flabellum, for example, where the line of junction of the
two laminee that constitute each septum is indicated externally by a single costal ridge.
But a more attentive study of this singular fossil has made us change our opinion, and

' Journ. of the Geol. Soc. of London, vol. iii, p. 368.
* Monogr. des Turbinolides, loc. cit.

CORALS OF THE LONDON CLAY. 27

has induced us to think that each group composed of three laminz, and corresponding to
one costa must be the homologue of a single normal septum. It also appears evident
that the first-mentioned hypothesis is incompatible with the mode of development of the
younger septa corresponding to the short coste; for wherever one of these younger costz
interposes itself between two older ones, a new group of three septal lamin makes its
appearance in the visceral chamber, between two of the old lateral plates, which, according
to this view, would belong to one septum, and a young septum, accompanied by two
half septa, would thus be included in the interior of an elder septum. Nothing of
the sort is ever met with in any known corallum, and would be contrary to the general
laws which appear to regulate the formation of the septal apparatus; but if we admit that
each group of these vertical lamme corresponds to a single septum in the ordinary
Polypidoms, all serious difficulties disappear, and a circumstance that tends to corroborate
this view of the subject, is, that in some Turbinolidz an intermediate tissue is seen between
the two lateral plates constituting each of the larger septa, so that if these three vertical
strata of sclerenchyma, instead of beg im contact, and intimately united, were separated
by a membranous fold or duct, each septum would no longer have the appearance of a
simple partition, but would resemble the trilaminate septal groups of the Dasmia. In the
present state of our knowledge concerning the structure and the mode of development of
this curious fossil, we must be cautious in our speculations concerning the signification of
the parts just described; but it is to be hoped that a complete solution of the question
will be obtained by the study of a greater number of these Corals. At all events, the
development of the septal apparatus must be very abnormal in Dasmia, and appears to
warrant the establishment of a separate zoological division for the reception of this extinct
genus.

Family OCULINIDA (p. xix).

1. Genus OCULINA (p. X1X).

Ocutina conrerta. Tab. II, figs. 2, 2a, 24.

Corallum composite, incrusting, forming an irregular, subglobose, or lobated mass, and
appearing to have always grown on some extraneous stem, which has disappeared during
the process of fossilization. ‘The corallites are not arranged in a regular way, but are
usually very closely set, and the calices are unequally prominent on the surface of the
coenenchyma, which is compact, and moderately thick ; its surface is covered with round,
unequal, crowded granulations, and presents no distinct coste. The calices are in general
quite circular, excepting when preparing to multiply by fissiparity, which is very seldom
the case; the edge is rather thin, and the fossula large, but not very deep. The columella
is sub-papillose. The sepfa (fig. 2 4) constitute three complete cycla, besides which some

28 BRITISH FOSSIL CORALS.

vestiges of an incomplete fourth cyclum often exists. The six systems, independently of
these rudimentary septa, are equally developed; the septa are thin, narrow towards the
apex, strongly granulated laterally, of unequal size according to their relative age, scarcely
exsert, and terminated by a slightly arched, almost undivided edge. The pati are thick,
narrow, and crispate; they form two coronets, and those corresponding to the secondary
septa are rather larger and more distant from the columella than those corresponding to the
primary septa. Diameter of the calice, two thirds of a line; depth of the fossula, half a line.

A vertical section of one of these corallites (fig. 2a) shows that the walls, as well as the
coenenchyma, are of a very compact structure, and are covered with minute granulations ;
thet the small tubercles arising on the lateral surface cf the septa are much less crowded ;
that the columella is constituted by small, irregular, filiform, ascending trabicule, and that
the loculi are devoid of dissepiments, or only contain very few of them.

The genus Oculina, reduced to the limits here assigned to that zoological division,
appears to have very few fossil representatives, for this is as yet the only known species
belonging to it that is not exclusively recent; and it might be almost considered as
constituting a distinct generic type, for it differs from all the recent species of Oculina by
the mode of arrangement of the corallites. In the latter the corallites affect a spiral order
in the ascending branches constituted by their union, and the ccenenchyma presents near
the calices some slight mdication of radiate costae, whereas in this fossil the corallites, as
we have already remarked, are quite irregularly grouped, and the surface of the ccenenchyma
is not at all striated.

Oculina conferta appears to be abundant in the London Clay at Bracklesham Bay.
We have received specimens of this fossil from Mr. Dixon and Mr. Frederic Edwards.

2. Genus DIPLHELIA (p. xxi).
Dippnenia papsttosa. Tab. I, figs. 1, 1a, 1 6.

Corallum composite, subdendroid, and rather tall. Corallites usually disposed alternately
in contrary directions, but appearing sometimes irregularly grouped, in consequence of
two series becoming united so as to form a single branch, or of a few individuals multiplying
by fissiparity. The calices placed far apart, quite circular, scarcely prominent, if at all so,
and united by a highly-developed mural ccenenchyma, the surface of which is covered with
closely-set, unequal, minute granulations, rather oblong, especially in the vicinity of the
calices (fig. 1a). Calicular margin very thin; fossula large, and very deep. Columella
very large, of a spongiose texture, and sub-papillose at the apex. Septa forming three
complete cycla, and six equally-developed systems; very narrow at the upper end, not
exsert, thin, granulated on their lateral surface, and presenting along their inner edge
delicate denticulations, which become larger towa ~. the columella, but do not assume the

CORALS OF THE LONDON CLAY. 29

appearance of rudimentary pali. The secondary septa are almost as large as the primary
ones, and thus give the appearance of twelve systems (fig. 14). Sometimes septa of the
fourth order exist in one of the real systems, and in that case the neighbouring tertiary
septa become at the same time as large as the elder ones, so that the septal apparatus
becomes divided into fourteen almost equal parts. Diameter of the calice, one line ; depth
of the fossula, one line, or more.

The great development and the compact structure of the coenenchyma are rendered
manifest by.a vertical section of one of these corallites (fig. 1a) ; this preparation is also
necessary to show the denticulations of the inner edge of the septa, and if continued to a
certain distance from the calice, brings to view a few irregular, incomplete, locular
dissepiments.

The new generic division, to which we have given the name of Diplhelia, comprises the
Oculinide that multiply by alternate gemmation, and have denticulated, non-exsert,
unequal septa, no pali, and a large columella. It differs from Astrhelia by the mode of
arrangement of the corallites dependent on the alternate position of the reproductive buds,
by the existence of a well-developed columella, and the absence of costal striae near the
ealices. ‘The mode of gemmation is the same in Amphelia and Enallhelia, but in these
Oculinidz the septa are entire and exsert, the columella is rudimentary, and the costal
strize are well marked near the calices.

Four species compose at present this small group ; they are all fossil, and belong to the
Eocene Fauna. ‘T'wo of these Corals have been described by M. Defrance, under the
names of Oculina raristella and O. Solanderi; the third is the Caryophyllia multostellata
of M. Nyst; the fourth is our Diplhelia papillosa. Diplhelia Solanderi, of which a pretty
good figure has been given by M. Michelin,’ differs from the latter by the existence of
numerous delicate, vermiculated, longitudinal sulci on its surface. Diplhelia raristella’”
differs from D. papillosa, by the calices beg smaller and not so deep, by the septa being
thicker, and the columella less developed. Diplhelia multostellata® is principally charac-
terised by the approximation of the calices, and their dilated form.

Dipthelia papillosa has as yet been found only at Bracklesham Bay, where it appears to
be abundant. The specimens here described have been communicated to us by Mr.
Bowerbank, Mr. Dixon, and Mr. Frederic Edwards.

1 Teon. Zooph., tab. xliii, fig. 19. 2 Michelin, loc. cit., tab. xliii, fig. 16.
3 Nyst, Coq. et Pol. foss. des Terr. tert. de la Belgique, tab. xlviii, fig. 10.

30 BRITISH FOSSIL CORALS.

Family ASTREIDAN (p. xxiii).
Tribe HUSMILIN At (p. xxiii).

(Husmiline aggregate).

1. Genus STYLOC@NIA (p. XX1X).

STYLOC@NIA EMARCIATA. ‘Tab. V, figs. 1, 1 a.

ASTROITE DEMI-CYLINDRIQUE, Guettard, Mém. sur les Arts et les Sciences, t. iii, p. 480,
tab. xxxi, figs. 40, 41, 42, 1770.
ASTREA EMARCIATA, Lamarck, Hist. des Anim. sans Verteb. t. ii, p. 266, 1816; 2™° edit.
p. 417.
= — Lamouroux, Encyclop. Zooph., p. 127. 1824.
— _— Defrance, Dict. des Scien. Nat., t. xlu, p. 389, 1826.
— cyYLiInpRIcA, Ejusd., loc. cit., p. 379. (From a worn specimen.)
— styLopora, Goldfuss, Petref. Germ., vol. i, p. 71, tab. xxiv, fig. 4, 1826. (From a
frustrate specimen.)

CELLASTREA EMARCIATA, Blainville, Dict. des Sc. Nat., vol. Ix, p. 342, 1830; and Manuel
d’Actinologie, p. 377. (The fossil figured in the atlas of this
work, pl.liv, fig.5, under the name of Cellastrea hystriz,
belongs to this species.)

AsTREA EMARCIATA, Michelin, Icon. Zooph., p. 154, tab. xliv, fig. 6, 1844.

— CYLINDRICA, Ejusd., op. cit., tab. xliv, fig. 4.
— DEcORATA, Ejusd., op. cit., p. 161, tab. xliv, fig. 8.
SryLoc@nia EMaRcrIATA, Milne Edwards and J. Haime, Monogr. des Astreides, Ann. des
Sciences Naturelles, 3™° série, vol. x, p. 293, tab. vii, figs. 2, 2a,
1848.

It is only in the Eocene deposits of the Parisian basin at Grignon and at Parnes that
this species has as yet been met with in a good state of preservation, but its existence in
the London clay is sufficiently established by two small fossils found at Bracklesham Bay,
by Mr. Frederick Edwards, which do not appear to differ from the worn specimens found,
together with the well-characterised ones in the first-mentioned localities. 'The following
description is consequently derived principally from the Parisian specimens ; but in order
to avoid introducing into this Monograph any uncertain elements, we have figured the
British specimens in preference to more perfect foreign fossils with which we consider them
as being specifically identical.

Astrea emarciata is a composite Coral, of an oval, gibbous, or subramose form, which at
first sight appears to be completely free, but was in all probability primitively fixed on
some soft, globular, extraneous body, which after having been completely covered by the
incrusting Coral, disappeared by the progress of putrefaction, and has only left a central
cavity in the middle of the irregular globose mass thus produced: it consists of a thick

CORALS OF THE LONDON CLAY. 31

lamellar expansion, bent so as to shut up completely an irregular cavity, and to have all
the calices of its constituent corallites turned outwards The basal or mner surface of
this lamellar corallum is coated with a thin, membranous epitheca, m which circular strie,
indicative of its mode of growth, are perceptible. The ca/ices are polygonal, and rather
unequal in size; they are separated by a simple edge, which is common to the two
adjoming corallites, and is thin where these corallites are crowded together, but rather
thick where the reproductive process has been less active; in the latter case these mural
edges are covered with numerous well-marked granulations, but in the former, no appear-
ance of granulations is tc be seen. Sometimes these two states are met with in different
parts of the same specuuen, but m others the whole mass presents one or the other of
these forms, and may then be easily mistaken for distinct species. It is thus that
M. Michelin has been led to consider the thick-walled variety as constituting a new species
to which he has applied the name of Astrea decorata. he calicular margins present also
at each corner a well-formed cylindro-conical columnar tubercle or process, which is not
very thick at the basis, and is usually fluted by six or eight vertical furrows. In specimens
that have been much rolled by the sea, these mural processes are often worn away, and
these dilapidated Corals have also been described by palontologists as a distinct species :
they constitute the Astrea cylindrica of M. Defrance. The columella is slender, cylindrical,
and free down to a great distance from its apex, but presents at the bottom of the fossula
vertical striz, which are produced by the prolongation of the principal septa along its
sides, and are particularly manifest in some worn-down specimens, such as those found at
Bracklesham, and figured in this Monograph (fig. 1 a). The septa form two complete
well-developed cycla; a third cyclum is rudimentary in four of the systems, but well
developed in two systems, the secondary septa of which become nearly as large as the
primary ones, so as to give to the calice the appearance of having eight systems instead of
six, which is the fundamental number. ‘The cight large septa thus formed are broad, very
thin, almost glabrous, not exsert, and terminated by regularly-arched, undivided edges ;
the other intermediate septa are very sinall. ‘The interseptal dissepiments are simple,
somewhat concave, slightly raised towards the columella, and placed at the distance of
about one third of a line apart. ‘The breadth of the calice is about one line and a third ;
the height of the mural processes two thirds of a line.

The British fossils which we refer to this species, and which we have figured in the
annexed plates, have evidently been modified by the long-continued ection of the sea; the
septa are much broken, and the granulations of the calicular margins are not visible ; it
appears not improbable that the polypi to which they belonged did not live in the locality
where these remains have been found, and that the Corals were brought there by some
marine current. ‘They are very rare at Bracklesham Bay, but extremely common in the
Calcaire grossicr of the environs of Paris. M. Michelin states that the same species is met
with at La Palarea, and the Styloconia found in this locality is certamly very similar to
S. emarciata, but all the specimens that we have been able to examme, were in such a bad

32 BRITISH FOSSIL CORALS.

condition that we cannot give any decided opinion as to their specific identity with the
Corals described above.

Stylocenia emarciata differs from S. monticularia,’ and from 8S. Taurinensis? by the
number of the large septa which in the latter species is only six ; it differs much by its
general form from S. Lapeyrousiana, and resembles most S. lobato-rotundata,* from which
it may be distinguished by a greater development of the mural tubercles, the tenuity of
the septa, and the general form of the mass.

2. StyLoce@nia MonticuuaRIA. ‘Tab. V, figs. 2, 2a, 26.

STYLOPORA MONTICULARIA, Schweigger, Beob. auf Naturg. reisen, tab. vi, fig. 62, 1819.
(Correct figure.)

ASTREA HYSTRIX, Defrance, Dict. des Sc. Nat., vol. xl, p. 385, 1826.

CELLASTREA HYSTRIX, Blaznville, Dict. des Sc. Nat., vol. lx, p. 342, 1830; Manuel d’Actin.,
p. 377.

ASTREA HystRIXx, Michelin, Icon., p. 160, tab. xlv, fig. 1, 1845.

STYLOC@NIA MONTICULARIA, Milne Edwards and J. Haime, Ann. des Se. Nat., 3™° série,
vol. x, p. 294, 1848.

Corallum composite, elongated, and nearly cylindrical when young, but becoming, by
the progress of growth, oval, sub-globose, and free, (fig. 2) ; with an empty central cavity,
the parietes of which are coated with a thin epitheca, wrimkled circularly (fig. 2 4). The
exterior surface of this hollow mass is covered with the calices, the borders of which vary in
form according to the age of the compound Coral; when the colony of polypiis young, the
margins of the calices are circular, prominent, and separated from each other by a striated
surface; but in older groups, they become polygonal and united, so as to form a single
thin ridge common to the two neighbouring corallites. The mamilliform processes that
rise at the angles of the calices are stout, conical, broad at the basis, and covered with
numerous prominent, sub-lamellar, finely-denticulated, vertical striz. The columella 1s
very slender, prominent, free a great way down, cylindrical towards the apex, and slightly
compressed at the basis. The septa form only two complete cycla ; the primary ones are
much larger than those of the second cyclum, broad, not exsert, granulated laterally, and
terminated by an undivided convex edge. Sometimes the strize of the mural processes are
continued down along the parietes of the visceral chamber, and assume the appearance of
rudimentary septa. Diameter of the calice, two thirds of a line; height of the mural
processes, half a line.

This species differs from S¢ylocenia emarciata, and JS. lobato-rotundata, by the regular
and equal development of the six systems of septa, whereas, in the latter, two of these

1 See tab. iv, fig. 2. 2 Michelin, Icon., tab. xin, fig. 2.
3 Michelin, op. cit., tab. lxx, fig. 3. + Michelin, op. cit., tab. xi, fig. 2.

CORALS OF THE LONDON CLAY. 33

systems are apparently double, so that the septal apparatus is divided into eight nearly
equal groups. Stylocenia Lapeyrousiana differs from it by its conical form, and by the
existence of three complete cycla of septa. Stylocenia Taurinensis has equally but two
septal cycla, but the mural processes are much smaller than in 8. monticularia ; the calices
are larger, and the primary septa are united to columella very high up.

Stylocenia monticularia has been found at Bracklesham Bay, by Mr. Frederick Edwards,
but appears to be rare in that locality; it is, on the contrary, very common at Grignon,
and in many other places near Paris.

2. Genus ASTROCENIA (p. XXX).
ASTROCENIA PULCHELLA. ‘Tab. V, figs. 3, 3a, 34, 3c.

Corallum composite, astreiform, massive, or subcolumnar, and presenting at its under
surface a common plate, covered with a complete epitheca, delicately wrinkled by concentric
strie. Corallites approximating more or less, according to the age of the Coral and the
degree of activity with which gemmation has been carried on. When the corallites are
not crowded together, the calices are circular, and have a distinct though not prominent
edge ; they are also separated by a pseudo-ccenenchyma, the surface of which is covered
with small costal ridges, that are usually denticulated, so as to assume the appearance of
rows of round, obtuse granule (fig. 3c). When the calices approximate, they become
somewhat polygonal, and their margins are separated only by a narrow furrow, or united
so as to appear simple. The calicular fossula is very shallow. Colwmella cylindrical,
obtuse, and free to a considerable extent, but not rising quite so high as the septa. Three
complete cycla of sepfa, and six equally-developed systems; the septa of unequal size,
according to their relative age, straight, slightly exsert, closely set, feebly granulated, rather
thick externally, and having their upper edge entire and convex. Breadth of the calices,
two thirds or three quarters of a line.

This species belongs to the division of the irregular Astroccenia,’ for independently of
the slight inequality perceptible in the size of the calices, it is evident that gemmation takes
place in this Coral simultaneously at various parts; but it differs from the other species of
the same section, by the number of the septa; here, as we have already said, the six systems
are equally developed, whereas in Astrocenia Konincku, A. Orbignyana, A. reticulata,
A. ornata, A. ramosa, and A. decaphylla, there is always apparently eight or even ten
systems.

We know of only three small specimens of this species, which were found at Bracklesham
Bay, and belong to the cabinet of Mr. Frederick Edwards.

1 See our Monograph of the Astreide, Ann. des Sc. Nat., 3™° série, vol. x.

5

34 BRITISH FOSSIL CORALS.

Family KUPSAMMID AX (p. hi).

1. Genus StEPHANOPHYLLIA (p. lin).
STEPHANOPHYLLIA DiscoIpES. ‘Tab. VI, figs. 3, 3a, 36.

STEPHANOPHYLLIA DiscorDES, Milne Edwards and J. Haime, Aun. des Sc. Nat., 3™° série,
vol. x, p. 93, 1848.

Corallum simple, extremely short, and discoidal ; its under surface almost horizontal,
somewhat prominent in the middle, and showing no trace of adhesion. Coste very narrow,
radiate, alternating with the septa, corresponding to these in number, and nearly equal in
breadth, but differmg much in length, according to the cycla to which they belong ; the
smaller ones often united to the larger ones at their basis, and thus giving to the latter a
dichotomous appearance (fig. 34). All these costal striae are composed of a single row of
rather indistinct granule, and are united by small intercostal trabicule, thus constituting
the tissue of the discoidal wall, and the radiate rows of pores that exist in this part of the
corallum, and give to it the appearance of a microscopical sieve. ‘The upper or calicular
surface somewhat convex, and presenting in the centre a small, narrow fossula, at the
bottom of which there appears to be a rudimentary papillose columella. Septa forming five
cycla, of very unequal size, thin, very slightly granulated, not projecting laterally beyond the
edge of the wall, and having the upper edge rather angular. Those of the first and second
order large, straight, and free at their inner end; all the others bent towards one another,
and cemented along their inner edge, so as to constitute a series of slightly undulated
arches, superposed and increasing in size from the circumference of the calice towards the
centre; the largest are formed by the septa of the fourth cyclum, which unite two by two,
along the inner edge of the tertiary ones (which are very short), and thus constitute on
each side of the secondary septa a single lamina, that advances still further towards the
centre of the calice, and joins the neighbouring secondary septum opposite the point of
junction of its homologue, so as to give to the central portion of the calice the appearance
of a six-branched cross of Malta; the septa of the fifth cyclum very small and marginal.
Diameter, two lines or two lines and a half; height, about half a line.

This fossil Coral differsfrom Stephanophyllia Suecica’ and S. Bowerbankii (Tab. IX, fig.4),
by the form of the septa, which in the latter two species are termimated by an arched edge,
and are spinulose laterally ; it is distinguished from Stephanophyllia astreata’ by the distance
that separates the large septa near the solmovelle, and from &. elegans,? S. imperialis,* and

' See our Monograph of Eupsammidze, Ann. des Sc. Nat., 3™° série, vol. x, p. 94.
2 Fungia astreata, Goldfuss, Petref. Germ., vol. i, tab. xiv, fig. 1.

3 Ann. des Sc. Nat., 3° série, vol. x, tab. i, figs. 10, 10a.

* Michelin, Icon. Zooph., tab. viii, fig. 1.

CORALS OF THE LONDON CLAY. 35

S. Wystii,1 by its diminutive size, and by its low, very feebly-granulated septa. It is
worthy of notice that S. discoides is the only species of this genus that has as yet been
found in the Eocene formations. We have seen four specimens of this Coral; they were
all met with at Haverstock Hill, and belong to the cabinet of Mr. Frederick Edwards.

2. Genus BALANOPHYLLIA (p. hi).

BaLANOPHYLLIA DESMOPHYLLUM. ‘Tab. VI, figs. 1, 1 a, 14, le.

BALANOPHYLLIA DESMOPHYLLUM, Milne Edwards and J. Haime, Monog. des Eupsammides,
Ann. des Sc. Nat., 3° série, vol. x, p. 86, 1848.

Corallum simple, adherent by a broad, incrustating basis, subturbinate, straight, rather
elongated, and slightly compressed. Coste almost straight, closely set, and formed of one
or more rows of irregular granule; the primary and secondary ones much taller and much
larger than the others, especially near the calice, and usually separated by five small ones,
two of which begin to appear at about two thirds down the wall, whereas the others extend
to the basis. Calice slightly arched, and almost elliptical; its long axis being to the short
one in the proportion of 100: 160. Calicular fossula deep and narrow. Columella
spongy, not highly developed, flat, and not prominent at the bottom of the calice. Septa
forming five cycla, usually complete; those of the fifth cyclum more developed than those
of the third order, and becoming cemented together two by two beyond the inner edge of
those of the fourth cyclum, and constituting thus in each half system two septal lamine,
that in their turn unite between the inner edge of the tertiary septa and the columella.
In the neighbourhood of the wall, the septa of the fifth cyclum that are situated next the
primary and the secondary ones are cemented to them, and do not usually correspond to
any distinct costz ; so that in each half system there are only five costz corresponding to
seven septa. The large septa are terminated by an undivided edge, and are much more
exsert than the others; all are thin, granulated laterally, very porous, and closely set ;
those of the younger orders are delicately denticulated. Height of the corallum about

1 We have given this name to a Stephanophyllia of the Antwerp Crag that we haye seen in M. Nyst’s
cabinet at Louvain, and had been referred by that author to the S. zmperialis (Coquilles et Polyp. foss. de
Belgique, p. 633, tab. xlviii, fig. 17). This figure is pretty good, but does not show the small septa. Not
having described it in our Monograph of Eupsammidz, we point out here its characteristic features. The
under surface of Stephanophyllia Nystti, nob., is somewhat concave. The coste are of almost equal thick-
ness, and do not appear distinctly composed of rows of granule ; they alternate with the well-developed septa,
but correspond to rudimentary septa of the sixth cyclum; the younger ones are, as usual, united by their
base to the elder ones, but this apparent bifurcation takes place only very near the centre of the corallum ;
the intercostal furrows become gradually wider from the centre towards the circumference of the wall, and
are bored with pores, that increase in size in the same manner. ‘The calicular fossula is very deep. The
septa are disposed in the same way as in S. discoides and S. elegans, but are much taller, thinner, and more
angular; they are denticulated externally, and present on their lateral surfaces radiate strize, which resemble
incomplete synapticulz ; those of the last cyclum are very small. Diameter nearly an inch ; height, 53 lines.

36 BRITISH FOSSIL CORALS.

seven lines ; long axis of the calice, three lines and a quarter ; short axis, two lines ; depth
of the fossula, two lines.

This species belongs to the section of the fixed Balanophylliz, and is consequently
easily distinguished from B. prelonga’ and B. Gravesii,’ which are only sub-pedicellate.
The nakedness of the wall, and quite rudimentary state of the epitheca, distinguishes it
also from B. calyculus, B. verrucaria,* and B. cylindrica’ In B. geniculata® and
B. Cumingit’ there are but four cycla of septa, whereas in the above-described fossil there
are five cycla. It differs from B. ctalica® by its elongated and compressed form, from
B. Bairdiana by its exsert septa, and from B. tenwistriata by the compressed form of its
lower part, and the thickness of its principal costa. It resembles most this last-mentioned
species, which belongs to the Calcaire grossier of the Parisian basin.

Balanophyllia desmophyllum is found at Bracklesham Bay, and has been communicated
to us by Mr. Dixon and Mr. Frederick Edwards.

3. Genus DENDROPHYLLIA (p. li).

DENDROPHYLLIA DENDROPHYLLOIDES. ‘Tab. VI, figs. 2, 2a, 246, 2c.
OcULINA DENDROPHYLLOIDES, Lonsdale, in Mr. Dixon’s manuscript work on the Chalk
Formations and Tertiary Deposits of Sussex.
DENDROPHYLLIA DENDROPHYLLOIDES, Milne Edwards and J. Haime, Ann. des Sc. Nat.,
3™° serie, vol. x, p. 102, 1848.

Corallum composite, appearing usually to have incrusted the stem of some marine
plant which has been destroyed during the process of fossilization. Gemmation irregular.
Corallites short, very unequal in size, rather closely set, united by their basis, and free
down to a variable distance from the calice, so as to project more or less on the surface of
the common mass, or even to form a certain number of somewhat ramified branches (fig. 2).
Coste delicate, numerous, closely set, almost equal in breadth, composed of a row of
uregular, conical granule, having a sub-vermiculate appearance, and becoming more
irregular and more flexuous in the parts where they unite with those of neighbouring
corallites (fig. 24). Mural pores large, and very distinct near the calice, but ceasing to be
so lower down, where the tissue of the wall becomes very compact. Cadices regularly cir-
cular, with the edge rather thin, and the fossula infundibuliform, but not deep. Co/umella
spongiose, not much developed, and appearing to be but slightly prominent at the bottom
of the fossula. Septa forming four complete cycla, and sometimes a rudimentary icomplete
fifth cyclum; very thin, unequal, not exsert, or only very slightly so, and granulated

' Michelin, Icon., tab. ix, fig. 2. 2 Michelin, Icon., tab. xliii, fig. 7. 3 See tab. 1, fig. 3.
+ Milne Edwards and J. Haime, Ann. des Sc. Nat., 3™° série, vol. x, tab. i, figs. 6, 6 a.

5 Michelin, op. cit., tab. viii, fig. 15.

6 D’Archiac, Mém. de la Soc. Géol., 2™° serie, vol. ii, tab. vii, fig. 7.

7 Milne Edwards and J. Haime, loc. cit., fig. 8. 8 Michelin, loc. cit., tab. ix, fig. 15.

CORALS OF THE LONDON CLAY. 37

laterally. The six systems very distinct; the primary septa much broader and taller than
the others ; the secondary ones smaller than those of the fourth order, which unite to those
of the fifth order opposite the almost rudimentary tertiary septa, and continuing to bend
towards the secondary ones, unite two by two along the inner edge of these, and so
constitute six laminee, that advance almost to the columella, and appear at first sight to be
prolongations of the secondary septa (fig. 3). The largest of these corallites are about
two lines in diameter at the calice, and project little more than a line above the common
mass ; the depth of the fossula is about one line and a quarter. The young mdividuals
very soon acquire all their septa.

The genus Dendrophyllia has many representatives in the seas of the present period,
and in the upper tertiary formations, but the species here described is the only one that has
as yet been found in Hocene deposits, and is the oldest known, for the various fossil Corals
appertaining to remoter geological periods that have been referred to this generic division
by M. Michelin, and by some other authors, do not in reality belong to it. Dendrophyllia
dendrophylloides differs from D. ramea,' D. Taurinensis,’ and D. digitalis,’ by its wregular
gemmation; from D. Cornigera,* D. irregularis, D. amica,’ and D. aaifuga, by having
fewer septa (a cyclum less) ; and from D. gracilis’ by not being arborescent. It resembles
most our Dendrophyllia Cecilliana, but this last-mentioned species, which lives in the Chinese
seas, is sufficiently characterised by its broad costa, formed by double or triple rows of
small granulations, and by its large prominent columella.

This fossil Coral is found at Bracklesham Bay, and has been communicated to us by
Mr. Bowerbank, Mr. Dixon, and Mr. Frederick Edwards.

4. Genus STEREOPSAMMIA (p. In).

STERFOPSAMMIA HUMILIS. ‘Tab. V, figs. 4, 4a, 46.

od
of an regular basal gemmation. Corallites crowded together, cylindrical, short, united

by the basis, and free down at least two thirds of their length from the calice. The costal
tissue which unites them at their basis, is in general but little developed, and does not
deserve the name of coenenchyma. Gemmation seldom takes place laterally ; sometimes,
however, reproductive buds are formed on the side of a parent corallite at a certain distance
from its basis, and will thus produce a slight appearance of ramification. Coste very

Corallum composite, incrusting, glomerulate, remaining low, and increasing by means

1 Madrepora ramea, Solander and Ellis, Zooph., tab. xxxvili.

* Michelin, op. cit., tab. x, fig. 8.

3 Michelin, loc. cit., tab. x, fig. 10; and tab. Ixxiv, fig. 4.

+ Esper, Pflanz. Madrep., tab. x.

> Michelin, op. cit., tab. lxxiv, fig. 3.

6 Milne Edwards and J. Haime, Ann. des Sc. Nat., 3° série, vol. x, tab. i, fig. 9.
7 Milne Edwards and J. Haime, loc. cit., fig. 13.

38 BRITISH FOSSIL CORALS.

slender, closely set, projecting very little, equal, sub-vermiculate, and assuming the
appearance of vertical striz irregularly broken at short distances, but not distinctly
composed of granulations, as is the case in most Eupsammide (fig. 4a). Walls perforated,
as usual in this family, near the calice, but becoming compact lower down. Calices
circular, infundibuliform, rather deep, and having an obtuse edge. Coiumella quite
rudimentary, or not existing at all. Four septal cycla, the last of which is quite
rudimentary, whereas the others are well developed proportionably to theirage. The septa
are very thin, closely set, not remarkably exsert, termmated by an oblique, nearly entire
edge, and proceed in a straight direction towards the axis of the corallum, but present an
undulate appearance, due principally io the existence of a few large lateral granule. The
six systems are equally developed, and very distinct; the primary and secondary septa
meet along their inner edge in the middle of the visceral chamber. Height of the
corallites, about two or three lines; diameter of the calices, about two thirds of a line.

This fossil, of which we have seen but one specimen, that was found at Bracklesham Bay,
and belongs to the cabinet of Mr. Frederick Hdwards, is the only known species of the genus
Stereopsammia. The regular radiate structure of its calice distinguishes it from most of
the Eupsammide, and in the genus Coenopsammia, where the same character is met with,
the columella is essential and well developed,’ whereas in Stereopsammia it does not exist,
or is quite rudimentary.

Family PCRITID (p. lv).
Tribe PORITIN A (p. lv).

1. Genus Lituaraa (p. ly).

Lirnar#a Wessteri. ‘Tab. VII, figs. 1, 1a, 16, le.

Astrea WesstTERI, J. S. Bowerbank, on the London Clay Formation, in Charlesworth’s
Mag. of Nat. Hist., new series, vol. iv, p. 23, figs. a, B, 1840.

SIDERASTREA WeBsTERI, Lonsdale, in Mr. Dixon’s unpublished work on the Chalk Forma-
tions and Tertiary Depos:ts of Sussex.

Corallum composite, incrusting, adherig in general to large pebbles, and forming a
thick convex mass, on the edge of which some traces of a rudimentary epitheca are
sometimes perceptible. Multiplication by gemmation in the spaces comprised between the
calices. Corallites sometimes united by a spongy ccenenchyma; in other parts crowded
together so as to render the calices almost polygonal, and separated only by a thin, simple,
common margin. Calices infundibuliform, but not deep. Columella well developed,

1 Annales des Sc. Nat., 3™° série, vol. x, tab. 1, figs. 11, 12.

CORALS OF THE LONDON CLAY. 39

of a spongy texture, not projecting at the bottom of the fossula, and terminated by a
somewhat papillose surface. Septa thin towards the columella, thick externally, strongly
echinulated laterally, broad, closely set, not exsert, terminated by an oblique crenulated
edge, and forming three complete cycla, besides which there is sometimes a very
incomplete fourth cyclum. The secondary septa differ but little from those of the first
order; the tertiary ones also well developed, bent towards those of the second cyclum, and
cemented to them along the inner edge near the columella. A horizontal section shows
that the visceral chamber is cylindroid at some distance from the calice, and that the
spongy tissue of the walls and the columella becomes much developed (fig. 1c). The
fenestrate structure of the septa is seen in a vertical section of the corallum, represented at
fig. 1 4.
Breadth of the calices, nearly two lines; depth, half a line.

This fossil is very abundant at Bracklesham Bay. The specimens from which we have
drawn up the preceding description belong to the collections of the Geological Society,
of Mr. Bowerbank, Mr. F. Dixon, and Mr. Frederick Edwards.

Some other Corals that have been described under the names of Astrea or of Porites,
and that belong to the Calcaire grossier of the Parisian basin, are also referable to our
genus Litharea, but all differ specifically from LZ. Webster. In L. Deshayesiana,
L. Heberti,? and L. bellula,*® the calices are smaller, and the septa less numerous; the
third cyclum, which is always complete in L. Websteri, is complete in the last of these
three species, and does not exist in the first two. In Lithare@a ameliana‘ and L. crispa,
which resemble most the London Clay fossil, the walls are thinner and more prominent,
and the septa more echinulate.

1 Porites Deshayesiana, Michelin, Icon. Zooph., tab. xly, fig. 4.

2 Litharea Heberti, nob. This undescribed species presents the following characters: Corallum
composite, convex, massive, and often formed of superposed layers. Common epitheca moderately developed.
Walls scarcely distinct. Calices polygonal and shallow. Columella not well developed, and appearing to
be formed only by the inner marginal denta of the septa. Septa not exsert, very thick, especially outwardly,
strongly echinulated laterally, terminated by an almost horizontal, spinular edge, and forming only two
cycla. The twelve septa are nearly equal in size, and of a very porous structure; the spiniform granula-
tions which cover their lateral surfaces are so highly developed, that they often become united to those of
the neighbouring septum. This fossil has been found in an excellent state of preservation at Auvert, by
M. Hebert, and appears to be specifically identical with same dilapidated corals met with at Valmondois.

3 Astrea bellula, Michelin, op. cit., tab. xliv, fig. 2.

4 Astrea ameliana, Defrance ; Astrea muricata, Goldfuss, Petref. Germ., vol. i, tab. xxiv, fig. 3.

5 Astrea crispa, Michelin, loc. cit., tab. xliv, fig. 7; (but not the Astrea crispa of Lamarck).

40 B'isTISH FOSSIL CORALS.

2. Genus THouaRrma (p. lvi).

Horara@a Parisiznsis. Tab. VI, figs. 2, 2 a.

ALVEOLITES PaRISIENSIS, Michelin, Icon. Zooph., p. 166, tab. xlv, fig. 10, 1845.

Corallum composite, and appearing to have lived fixed to the stem of some Fucus,
which it incrusted all round, so as to constitute, after the destruction of this extraneous
body, a hollow cylinder, open at both ends. The lamellar expansion thus rolled round is
very thin, and its inner or basal surface is covered by an extremely delicate epitheca. The
calices which occupy the opposite surface, and are consequently placed all round the
exterior of the above-described cylinder, are infundibuliform, deep, irregularly polygonal,
surrounded by a prominent margin, and sometimes slightly turned towards one of the
extremities of the corallum, which was probably its upper end. The fossula is small and
circular ; its centre is occupied by a fasciculated columella, composed of delicate vertical
processes, which are quite separated from each other, excepting towards the apex (fig. 2a).
The vertical section of the corallum, by means of which the composition of the columella
is seen, shows also that the tissue of the whole mass is uniformly and delicately spongy ;
no appearance of cost, of septa, or of any radiate structure is perceptible. The diameter
of the specimen that we have figured is about a line and a half, and the thickness of the

lamellar expansion that constitutes this cylinder, about half a line; the calices are also
about half a line in breadth.

‘This species has been found both in the London Clay at Barton and the Calcaire
erossier of the environs of Paris. The British specimen represented in our plates belongs
to the cabinet of Mr. Frederick Edwards. We have examined many of these fossils, but
owing to the very small size.of the corallites, and the extremely delicate structure of their
constituent parts, we fear that some of their characteristic features may have escaped from
observation, and we feel much uncertainty respecting the natural affinities of the generic
division of which it is as yet the only representative. We have not been able to ascertain
the existence of any tabulze in the interior of the visceral cavity, and therefore it would
appear to be allied to Poritidee rather than to Milleporide ; but it bears great resemblance
to the latter, and we are inclined to think that, when better-preserved and older specimens
become known, it will prove to be a tabulated Zoantharia, and if that be the case, there will
no longer be any reason for distinguishing Holarea from our genus Axopora (p. lix). It
ts therefore only provisionally that we place it in the family of the Poritide.

CORALS OF THE LONDON CLAY. Al

Orper 2.—A LCYONARIA.
Family PENNATULID&.

Genus GRAPHULARIA,

GraPHULARIA WetHERELLI. ‘Tab. VII, figs. 4, 4a, 40, 4¢, 4d, 4e.

PennatuLa, J. Decarle Sowerby and Wetherell, in Geol. Trans., 2d series, vol. y, part 1,
p. 136, tab. viii, fig. 2a, b, 1834.

Corallum styliform, straight, very long, cylindroid towards the lower extremity, sub-
tetrahedral at the upper part, and presenting on one side a broad shallow furrow. Surface
appearing smooth, when examined by the naked eye, but showing, when placed under the
microscope, a multitude of small, longitudinal, closely-set, striz, that seem to indicate a
fibrous structure. Transverse section showing the existence of a thin coating and a
radiate structure in the body of the Coral. Diameter of the thickest part, two thirds of a
line; probable length, more than a foot.

We have only seen small fragments of this styliform Coral, that evidently constituted
the central stem of some aggregate polypi of the family of the Pennatulide. Some of
these broken remains are almost cylindrical, and usually thicker than others that are
imbedded m the same mass of clay, and have a sub-tetrahedral form; others, again, are
intermediate between the former, both by their size and their form, and have the same
radiate structure and striated surface. It is therefore probable that they all belonged to
the same species, and constituted a long, slender, sclerenchymatous axis, somewhat similar
in form to that of Pennatula, but resembling that of Virgularia by its structure. The
characters of the corallum thus reconstructed are also nearly allied to those of Pavonaria
and Umbellularia, but differ from those of all the known recent genera of Pennatulide.
It is brittle, and presents a radiate section, as in Virgularia, but is not cylindrical from one
end to the other, as is the case in the latter, nor is it from top to bottom of a tetrahedral
form, as in Pavonaria; it never appears to be twisted like the stem of Umbellularia ; it
united in its different parts the two forms that are found separately in the two first-
mentioned genera, and so far resembles Pennatula ; but in the latter the square portion is
situated towards the lower end, and the apex is cylindrical, whereas in the above-described
fossil, it is the upper slender part that presents a square section, and the thick basal part is
cylindrical ; it must also be remembered that the axis of Pennatula is not very brittle, and
does not present a radiate structure when cut transversely, but appears rather of a fibrous
structure. Inthe recent genus Lithuaria, the styliform axis is tapering towards the lower
end, and inflated, pitted, and even somewhat echinulate at its upper extremity. It is also
impossible to refer the elongate stem of our London Clay Pennatulida to the genus

6

4.2 BRITISH FOSSIL CORALS.

Veretellum, for in the latter the axis is quite rudimentary. We therefore considered it
advisable to designate this fossil Coral by a peculiar generic name, but we are not as yet
sufficiently well acquainted with its characters to be able to form a complete idea of the
polypi to which it belonged.

Graphuaria Wetherelli is the only known species of this genus of Pennatulide ; it
was discovered about twenty years ago by Mr. Wetherell in the London Clay at Hampstead
Heath and at Highgate. Mr. Frederic Edwards has also found fragments of it at Barton
and at Haverstock Hill; and it is to his kindness that we are indebted for the specimens
described in this Monograph.

Family GORGONID 2.
Tribe ISIN A.

1. Genus Mopseka.

1. Mopsza costata. ‘Tab. VII, figs. 3, 3a.

Corallum arborescent, dichotomous, and composed of epidermic basal sclerenchyma,
the ossification of which is intermittent, so as to constitute a series of calcified cylinders,
separated by non-ossified discs. ‘The branches appear to spread out in one plane; they
are thin, elongated, cylindrical, and deeply fluted longitudinally ; each of them, immediately
above its separation from the parent branch or stem, is bent outwards and upwards, so as
to represent. an inverted ogival angle. The corneous articulations, which have been de-
stroyed during the process of fossilization, are very thin, and do not appear to have existed
in any of the non-bifurcated branches. The longitudinal costz are straight, thick, pro-
minent, denticulated on their sides, and separated by deep furrows. All those belonging
to the same jomt are in general of the same size; but in some of the thickest branches,
where they are the most numerous (about twelve), some very small ones are sometimes
visible between the larger ones. ‘The diameter of the thickest branches in the specimens
here described is about half aline; that of the smallest not more than a tenth of a line.

The genus Mopsea was established by Lamouroux, but more correctly characterised by
Ehrenberg, who refers to it four recent species—the MMopsea dichotoma of Lamouroux,
the M. gracilis, M. erythrea, and M. encrinula, Ehrenberg ; but it is doubtful whether
the last does in reality appertain to this division of the Isinze. Mr. Dana is of opinion
that the Isis coralloides of Lamarck is also a Mopsea; but in all of these species the
corallum is but slightly striated, and never presents anything like the strong costa which
exist in the above-described fossil.

We have seen two specimens of this Gorgonida, found in the London Clay at Holloway,
by Mr. Frederic Edwards.

CORALS OF THE LONDON CLAY. 43

2. Genus WEBSTERIA.
WEBSTERIA CRISIOIDES. Tab. VII, figs. 5, 5a.

Corallum composite, slender, and dichotomous, with its branches straight, flat, of the
same dimensions as the stem, spreading out on one plane, and forming very acute angles
with each other. Corallites subverruciform, disposed in opposite order, and forming
two vertical series, the outer edge of which is occupied by a row of small, oblique, circular
calices. These two laieral parts are separated by a median line, which usually has the
appearance of a small furrow; sometimes they disunite, and so bring to view a small,
styliform, central axis. Breadth of the branches about a fifth of a line; height of the
corallites, a quarter of a line.

It is not without much uncertainty that we refer this delicate fossil to the family of the
Gorgonidz. By its general aspect, and by the mode of arrangement of the verruciform
individuals of which it is composed, it resembles the genus Pterogorgia of Ehrenberg, and
the existence of a central axis furnishes a strong argument in favour of the opinion which
we have adopted provisionally ; but, on the other hand, the structure of the individuals is
very similar to that of some Sertularide, and still more so to divers Bryozoa, such as Crisza
denticuluta. The specimens that we have examined have not enabled us to decide the
question concerning the natural affinities of the animals to which these organic remains
belonged; but whether they be Polypi or Bryozoa, they appear to differ from all the known
generic forms, and to constitute the type of a new genus, that we dedicate to Mr.
Webster, whose observations on the formation in which they are found have been so
serviceable to geology.

Websteria crisioides is the only species belonging to this zoological division. It was
discovered in the London Clay at Haverstock Hill, by Mr. Frederic Edwards, to whose
kindness we are indebted for the communication of the specimens here described.

4d BRITISH FOSSIL CORALS.

CHAPTER IIT.
CORALS OF THE UPPER CHALK.

Tux fossil Corals found in the Upper Chalk of England are not numerous; they
belong principally to the section of simple Eusmiline, and appear to be peculiar to the
British Fauna. One species, it is true (the Parasmilia centralis), has been mentioned by
different geologists as existing also in the Chalk of Beauvais and in the north-west of
Germany, but we have great reason to think that these fossils are not specifically identical.
It is also worthy of notice, that even no species corresponding to those met with among
the Corals of the Upper Chalk of England have as yet been seen in the Chalk of Meudon,
and that a great difference exists between the predominant generical forms in the first of
these formations, and in the Chalk of Maestricht. In the latter some Cyathininz nearly
allied to that of England are met with; but the Diploctentum the Cyclolites, and the
ageregate Astreidz of Maestricht are represented by no corresponding forms in this part of
the British fossil Fauna, and the organic remains found in these two cretaceous deposits
have consequently a very different aspect. We must add, that the fossil Corals of the Chalk
of Faxoe are equally distinct from the British species, and that none of the latter have
been met with in the Lower Chalk Formations of England.

Orver ZOANTHARIA (p. ix).
Family TURBINOLID& (p. xi).

Tribe CYATHININ A (p. xii).

Genus CYATHINA (p. Xi).
Cyatuina L&vIGATA. ‘Tab. IX, figs. 1, la, 14, le, ld.

CyaTHINA L&VIGATA, Milne Edwards and J. Haime, Monogr. des Turbinolides; Ann. des Se.
Nat., 3™° série, vol. ix, p. 290, 1848.

MoNOCARYA CENTRALIS (pars), Lonsdale, in Dixon’s unpublished work on the Chalk Forma-

tions and Tertiary Deposits of Sussex, tab. xvii, figs. 12, 12a,

(perhaps also fig. 5, but not the other figures bearing the

same name, which are Parasmilia and probably Coelosmilia).

Corallum simple, elongated, adherent, cylindro-turbinate, straight, and im general
much contracted just above the basis, wnich is broad. Valls quite smooth, and polished

CORALS OF THE UPPER CHALK. Ab

towards the basis, but presenting towards the calice slightly-marked coste, which are
closely set, glabrous, or very delicately granulated, and almost equal im size. Calice
circular, or sometimes rather oval, shallow. Co/wmella moderately developed, not projecting
in the centre of the calicular fossula, composed of six or eight twisted, vertical processes,
and terminated by an equal number of papille. Septa forming four cycla, the last of
which is in general incomplete ; the septa of the fourth and fifth orders not existing in one
half of three of the systems or even of all six of these, so that the number of these
radiate laminz is reduced to 42, or even to 36>; sometimes, however, four of the systems
are complete, and the number of the septa then amounts to 48. These septa are well
developed, closely set, thin, but slightly granulated, rather exsert, and almost equal; the
principal ones are, however, a little thicker than the others. The padi are narrow, but
very thick, prominent, aud terminated by a curved edge; they all correspond to the
tertiary septa, and in the specimens where the fourth cyclum is complete, they exist in
front of all the septa of the penultimate cyclum, and are therefore twelve in number ; but
they are never so numerous when the fourth cyclum remains incomplete, and never
correspond to tertiary septa that are not separated by septa of the fourth cyclum. As
mentioned above, these latter septa are often wanting in one half of three of the systems,
and in that case there are consequently no pali corresponding to the tertiary septa of these
incomplete half systems, so that the number of pali is reduced to nine; two belonging to
each of the three complete systems, and one to each of the incomplete ones. The same
rule also holds good when all the six systems are deficient of the septa of the fourth
cyclum in one of their halves ; the tertiary septa of the complete halves are the only ones
having corresponding pali, so that the number of these organs is only six. The height of
the corallum varies between one inch and one inch and a half; in the tall specimens the
diameter of the calice is about four lines ; in the short and broad ones it is sometimes five lines.

This species is easily distinguished from the Cyathina Cyathus,' C. Smithii,? and
C. pseudoturbinolia,® by its never haying a fifth cyclum of septa. C. Guadulpensis* and
C. arcuata’ differ from the above-described fossil by the existence of distinct coste down
to the basis of the walls, and by the large size of the pali. C. /evigata most resembles
C. Koninckii,’ C. Bowerbankii;’ C. Debeyana, C. Brede, and C. cylindrica, and it may

1 See our Monograph of Turbinolide, Ann. des Sc. Nat., 3° série, vol. ix, p. 287, tab. iv, fig. 1.

2 Loc. cit., p. 288. 8 Loe. cit., p. 289, tab. ix, fig. 1. 4 Loe. cit., p. 290.

5 Loc. cit., p. 290. ® Loe. cit., p. 290. T Loc: cit.,; p. 292.

8 The three last-mentioned species were not known to us when we published our Monograph of
Turbinolide, and in order to render the comparison between the 7’. /evigata and the rest of the genus more
complete, it appears to us advisable to give a description of them here.

CYATHINA CYLINDRICA, nob. Corallum fixed by a broad basis, regularly cylindrical, straight, and not
very tall. Costz equal, flat, straight, closely set, not very broad, and very indistinct, especially towards
the basis. Calice circular, having a thick edge, and the fossula rather shallow. Columella very small, and
reduced to two or three small, almost indistinct, tubercles. Septa forming four complete cycla, very closely

46 BRITISH FOSSIL CORALS.

not be unworthy of notice that these five species are as yet the only representations of the
genus Cyathina that have been met with in the Chalk Formation. At first sight they appear
very similar, but by an attentive examination, constant and well-defined characteristic
differences are found between all. In C. Koninckhit, the corallum is always shorter, and
more regularly turbinate ; the pali are thicker, and the columella is reduced to two or three
thick, twisted processes. In C. Bowerbankii, on the contrary, the pali are much thinner,
and the surface of the walls appears granulous. C. cylindrica and C. Brede differ from
it by a very peculiar character, which exists also in C. Koninckii, but which is not met with
in any other species of the same genus, and is indeed quite an exception to the family of
Turbinolidz, the pali bemg only six in number, although the four cycla of septa be com-
plete, and corresponding to the septa of the antepenultimate cyclum, whereas they usually
correspond to those of the penultimate cyclum. ‘The thin, elongate form of C. Bred@ and
the quite cylindrical form of C. cylindrica, will also help to distinguish them from C. /evigata,
which differs also from C. Debeyana, by the latter having a well-marked epithecal fold near
the calice, a small columella, and thinner pali.

Cyathina levigata is found in the Upper Chalk at Dinton, in Wiltshire ; specimens may
be seen in the collections of the Geological Society, of Mr. Bowerbank, and of the Museum
at Paris.

set, and having stronger lateral granulations near the inner edge. The primary ones larger and rather
thicker than the others, but differing very little from the secondary ones; the tertiary ones are thinner and
smaller ; those of vhe fourth cyclum are distinct, but very small. Pali prominent, extremely thick, narrow,
strongly granulated laterally, and corresponding to the secondary septa. Height of the corallum about six
lines ; diameter of the calice three lines. Fossil from the Chalk of St. Peter’s Mountain, at Maestricht ; speci-
mens exist in the Museum of Natural History of Paris, and in the Tylerian Museum at Haarlem.
Cyatuina BrEeD&#, nobis. This fossil corallum, which we dedicate to Professor Van Breda, is adherent
by a rather broad basis, contracted immediately above, elongate, slender, much bent, and cylindrical towards
its upper part. The costz are not well marked, and the walls are almost smooth, but present sometimes
slight horizontal folds. Calice circular; fossula shallow. Columella but little developed, and sometimes
reduced to a single twisted process. Septa forming four complete cycla; but those of the last cyclum
rudimentary though distinct; the primary ones rather thick, especially towards the inner edge; the
secondary ones resembling those of the first cyclum, but rather narrower; the others very thin. The
‘granulations on the sides of the septa are conical, and very prominent. The pali corresponding to the
secondary septa, well developed, prominent, narrow, and appearing very thick, because they are flexuous.
Height, seven or eight lines ; diameter of the calice, two lines and a half. This species is also found in the
fossil state in the Chalk of St. Peter’s Mountain, at Maestricht; specimens exist in the collections of MM.
Van Riemsdyck and Bosquet, at Maestricht ; of M. Van Breda, at Haarlem ; and of the Museum at Paris.
CyaTHina DeBryaNa nob. Corallum cylindrical, elongate, slightly curved, and presenting near the
calicular margin a small but well-marked circular band, representing an incomplete epitheca. Calice
circular ; fossula not deep. Septa unequal, closely set, somewhat exsert, rather thick externally, but thin
towards the inner. edge. and forming four complete cycla; the secondary ones almost as large as those of
the first cyelum. Pali rather narrow, and not very thick. Height, one inch; diameter of the calice, three
lines ; depth of the fossula, one line. Fossil from the Chalk of Aix-la-Chapelle, discovered by M. Debay.

CORALS OF THE UPPER CHALK. AT

Family ASTREID A (p. xxiii).

Tribe EUSMILINA (p. xxiii).

1. Genus PARASMILIA (p. xxiv).
1. Parasmtiia cenTRauis. Tab. VIII, figs. 1, la, 14, le.

Mapreporite, Parkinson, Organ. Remains of a FormerWorld, vol. ii, tab. iv, figs. 15, 16, 1820.
MADREPORA CENTRALIS, Mantell, Geol. of Sussex, p. 159, tab. xvi, figs. 2, 19, 1822. (Cor-
rect figures.)
CARYOPHYLLIA CENTRALIS, Fleming, British Animals, p. 509, 1828.
— — Mantell, Trans. of the Geol. Soc., 2d series, vol. ii, p. 204, 1829.
= — Phillips, Ulust. of the Geol. of Yorkshire, part i, p. 119, tab. i,
fig. 13, 1829; 2d edit., p. 91.

_ — S. Woodward, Synoptic Table of Brit. Org. Remains, p. 6, 1830.
Caryopuytuia, R. C. Taylor, in Mag. of Nat. Hist., vol. iii, p. 271, fig. f, 1830.
LITHODENDRON CENTRALE, Ch. Keferstein, Die Naturgeschichte des Erdkorpers, vol. ii,

p- 789, 1824.
TURBINOLIA Excavata (?), Hagenow, in Leonard’s und Bronn’s Jahrbuch fiir Mineral., p. 229,
1839.
— CENTRALIS, Fred. Adolph Remer, Verstein. des Norddeutschen Kreidegebirges,
p- 26, 1840.
— —_— Bronn, Index Paleontologicus, p, 314, 1848.
PARASMILIA CENTRALIS, Milne Edwards and J. Haime, Monogr. des Astreides, Ann. des
Sc. Nat., 3° série, Zool. vol. x, p. 244. 1848.
MonocaRYA CENTRALIS (in parte), Lonsdale, in Dixon’s unpublished work on the Chalk
Formations of Sussex, tab. xvii, figs. 1,3, 7, 7 a, 9
(ceeteris exclusis).

Corallum simple, cylindrico-turbinate, fixed by a rather broad basis, above which it is
much contracted, elongate, irregularly bent in various directions, and presenting a series of
unequal contractions and circular dilatations. Coste closely set, and distinct from the
calicular margin down to the basis, where they are the most prominent; those corre-
sponding to the primary and secondary septa are rather larger than the others towards
the basis; but the tertiary ones soon become almost similar to the former, and at the
upper part of the wall all these large coste alternate with smaller ones belonging to
the fourth cyclum. All are covered with delicate granulations, which are most prominent
towards the lower part of the coste of the fourth cyclum, where they form simple series.
Calice circular, with the fossula less shallow than usual in this genus. Colwmella well
developed, somewhat prominent and crispate. Septa forming six equally developed systems
and four complete cycla; closely set, very unequal, broad, thin, slightly exsert, straight, or

48 BRITISH FOSSIL CORALS.

very slightly flexuous, and presenting laterally a few large granulations. Dissepiments
simple, almost horizontal, and few in number; about three from the top to the bottom of
each principal septum, as may be seen by means of a vertical section. Height varying
from one to two inches; diameter of the calice, four lines; depth of the fossula, two lines.

The genus Parasmilia, circumscribed within the limits assigned to it in the Introduction
to this Monograph, only contains seven species, all of which belong exclusively to the upper
beds of the Chalk Formations. Three of them (P. centralis, P. Gravesiana, and P. elongata)
have already been described in our Monograph of the Astreidee,’ and the four others
(P. Mantellu, P. Fitton, P. cylindrica, and P. serpentina) will be made known in the
present work. They all differ but little from each other, and in order to recognise them,
it is necessary that they should be compared together with attention. P. centralis, which
may be considered as the type of this small generic group, differs from P. Gravesiana,
P. elongata, P. cylindrica, and P. Mantellii, by its coste bemg always straight, rather
thick and never sub-lamellous, and rather flexuous, as in the four last-mentioned species ; it
is also to be remarked, that its costa are rather more prominent near the basis than higher
up, whereas the contrary is seen in the P. Gravesiana, and that the loculi are never
subdivided by small dissepiments, as is the case in P. elongata, P. cylindrica, and
P. Mantellit. P. serpentina, which bears more resemblance to it, is characterised by the
septa forming only three cycla, and the cost being very delicate, and rather indistinct
towards the basis. But it is with P. Fittonii that P. centralis is most closely alhed; the
former, however, is of a thicker form, its tertiary costae are more developed and more
delicately granulated, and its columella is much larger, and terminated by a sub-papillose
surface.

The specimens of P. centralis which we had the opportunity of examining were found
in the Upper Chalk at Northfleet, near Gravesend, and at Norwich. Mr. Phillips mentions
the existence of the same fossil at Dane’s Dike;? and Dr. Mantell has met with it at
Brighton, Lewes,’ Steyning, and Heytesbury.* Mr. Graves also alludes to it as being
found in the Chalk Formation of the Parisian basin at Beauvais ;? but we have much reason
to think that the species observed by that geologist is not the one here described, and must
be referred to our P. Gravesiana. M. Reemer and other authors equally apply the name
of C. centralis’ to a fossil found in the north-west of Germany, but we have not been able
as yet to verify the propriety of this determination, not having seen any of the specimens
discovered in that part of the Continent.

1 The species described in that work under the names of Parasmilia poculum, P. Fawasit, and
P. punctata, must now be referred to our genus Celosmilia, which is characterised by the entire absence of
the columella.

2 Op. cit., parti, p. 119. 3 Tllust. of the Geol. of Sussex, p. 160.

* Geol. Trans., 2d series, vol. iii, p. 204. 5 Geogn. de |’Oise, p. 701.

6 Versteinerangen des Norddeutschen Kreidebirges, p. 26.

CORALS FROM THE UPPER CHALK. 49

We must also remark that, in a note just published, our able friend M. Alcide
d’Orbigny* refers to the Caryophyllia or Parasmilia centralis as the type of a new genus,
designated under the name of Cyclosmilia, and characterised in the following terms:
“ Cyclosmilia are Parasmilia, in which the loculi are but very little divided by dissepiments,
the growth of the corallum is intermittent, the calice circular instead of being oval, and
the external coste distant from each other.” Now, with the exception of this last
peculiarity, which is not even met with in P. centralis, all these characters may be seen in
every species belonging to our genus Parasmilia, and we therefore can find no reason for

separating from it this new generic division.

2. Parasmitia Mantevii. Tab. VIII, fig. 2, 2a.

We have as yet seen but one specimen of this species, which appears to be very distinct
from all others. It is a small corallum, nearly straight, adherent by a broad basis, regularly
turbinate, and not very tall; but being in all probability susceptible of increasing much in
height by progress of age, as is the case with the other species belonging to the same genus.
The coste are narrow, sublamellar, closely set, distinct down to the basis of the corallum,
very echinulate, and somewhat crispate. Those of the primary and secondary cycla are
equally developed, and rather more prominent than the others, especially towards the basis
and the calicular margin; the tertiary ones also extend on the basal expansion of the
corallum, but are smaller ; and those of the fourth cyclum begin at a short distance above
the basis, and are very narrow at their lower part. ‘The intercostal furrows are broad, deep,
and divided by small transverse dissepiments, formed by rudiments of an exotheca. Calice
circular; fossula not deep. Colwmella, as far as we can judge by the specimen here
described, very similar to that of P. centralis. Septa forming four complete cycla; well
developed, thin, straight, closely set, rather unequal, and presenting well-marked strie
on their lateral surfaces. Height, seven or eight lines; diameter of the calice, nearly five lines.

This fossil differs from the other species belonging to the same genus, and more
especially from P. centralis and P. Fittoni, by its coste, which are equally prominent and
subcrispate, whereas in the latter they are smooth and never sublamellar. It resembles
more closely P. Gravesiana, P. cylindrica, and P. elongata ; but it differs from them by the
strong granulations of the coste. In P. serpentina the basis is almost smooth, and the
septa do not form so many cycla.

Parasmilia Mantelli was met with in the upper chalk at Bromley in Kent, by our
friend Mi. J. S. Bowerbank.

1 This paper, bearing the title of ‘ Note sur des Polypiers Fossiles,’ and published on the 10th of
October, 1849, contains the exposition of the characters of a series of new genera proposed by M. d’Orbigny.
The author assigns to most of these divisions the date of 1847, a period at which he appears to have adopted
them in the arrangement of his private collection; but in referring to them here or elsewhere, we have
considered it proper to quote the year of their publication, which is the only authentic date that could be

made use of if any question of priority should arise concerning them.

7

50 BRITISH FOSSIL CORALS.

3. Parasmitia Cytinprica. Tab. VIII, fig. 5.

It is not without some hesitation that we inscribe this species in the list of our
Parasmilia, for the specimen about to be described is extremely incomplete; but it does
not present the specific characters of any other species, and although very nearly allied to
P. elongata and P. Mantelli, it appears to differ from both in some essential points. The
fragment here alluded to is deficient both im the basis and in the calice, but it appertained
to a tall, nearly cylindrical corallum, that wassomewhat bent. The coste are almost equal,
extremely thin, sublamellar, but not very prominent, subflexuous, very slightly granulated,
and divided at short intervals by circles of small dissepiments, formed by rudiments of an
exotheca. The intercostal furrows are broad, rather shallow, almost destitute of granula-
tions, and presenting sometimes in the middle a small rudimentary costa. Columella well
developed. Septa forming four complete cycla, not very closely set, somewhat flexuous,
and slightly granulate; those of the first and second cycla equally developed, and rather
thick ; the tertiary ones smaller and thinner ; those of the fourth cyclum very small, although
the cost corresponding to them are as large as those of the other cycla. Length, above
two inches ; diameter, about six lines,

This fossil much resembles the Parasmilia elongata found at Ciply, but differs from it
by the unequal development of the septa belonging to the first two and to the last two
cycla, a mode of structure which does not exist in P. elongata. It differs from P. centralis,
P. Fittoni, and P. serpentina, by the delicacy and almost lamellar form. of the coste, and
bears greater resemblance to P. Mantelli and P. Gravesiana, from which it may, however,
be easily distinguished by the breadth of its intercostal furrows.

The specimen here described belongs to the Paleontological collection of Mr. J. S.
Bowerbank, and was found in the upper chalk at Norwich. Another fossil, which we
consider as belonging to the same species, exists in the Poppelsdorf Museum at Bonn, and
was found in the upper chalk at Darup, in Westphalia.

4. Parasminia Frrront. Tab. IX, fig. 2, 2a, 26.

Corallum stout, adherent by a somewhat broad basis, immediately above which it in
general becomes very narrow; elongate, much bent, and presenting at intervals circular
constrictions. Coste broad, closely set, not very prominent, excepting near the basis, down
to which they are quite distinct, rather unequal alternately, and covered with very numerous
and small granulations. Ca/ice circular ; fossula large and rather shallow. Co/wmella well
developed, but very slightly prominent, of a spongy structure, and terminated by a broad
subpapillose surface. Septa forming four complete cycla, rather thin, straight, not very
closely set, slightly exsert, and having but few granulations on their lateral surfaces.

CORALS FROM THE UPPER CHALK. 51

Those of the second order are nearly as large as those of the first set, and thus produce the
appearance of twelve tertiary systems. Height, from one to two inches; diameter of the
calice five lines ; its depth, two lines.

This species is easily distinguished from all the other Parasmilia by the great develop-
ment of its columella, which occupies nearly half the diameter of the calice, and by the
spongy structure of this organ. It most resembles P. centrals, from which, however, it
differs also by its thick form and the greater development of the tertiary costz towards
its basis. The breadth and delicate granulations of the costze may equally serve to dis-
tinguish it from P. Gravesiana, P. clongata, P. cylindrica, and P. Mantel. It differs
from P. serpentina by having an additional cyclum of septa, and by its basis not being
smooth, as is the case in the latter.

This fossil is found in the upper chalk of Norwich, and exists in the collections of the
Geological Society, of the Geological Survey, and of the Museum at Paris. It appears
probable that it has often been confounded with P. cextralis, and that some of the figures
referred to that species may in reality belong to it; but the engravings here alluded to are
not correct enough to enable us to decide this question.

5. PARAsMILIA (?) serPENTINA. ‘Tab. VIII, fig. 3, 3a, 30.

It is not without some doubts that we place this fossil in the genus Parasmilia, for in
the unique specimen that has come under our observation, the calice was in so bad a state
of preservation that it was impossible to decide whether the papille seen near the centre of
that part were fractured septa or remains of pali, or even trabicule belonging to the
columella. However, the first hypothesis appears most probable, and the general appear-
ance of the corallum is also very similar to that of all the other Parasmilia.

This fossil is almost cylindrical, slender, much elongated, and bent; it presents some
strongly marked circular constrictions, indicative as usual of a certain intermittence in
the progress of its growth. The coste are narrow, straight, rather unequal alternately,
scarcely distinct near the basis, but more prominent towards the upper part of each inflated
ring and near the calice; the calice is circular. ‘The septe form three complete cycla and
are rather closely set, exsert, and somewhat dilated exteriorly. The columella is well
developed. Length, one inch, seven lines. Diameter of the calice, two lines and a half.

This coral, belonging to Mr. Bowerbank’s collection, was found in the upper chalk at
Bromley, in Kent.

It is the only species of Parasmilia in which the fourth cyclum of septa does not exist ;
it is also characterised by its basis not being costulated.

52 BRITISH FOSSIL CORALS.

2. Genus CELOSMILIA (p. XXV.)

Canosminia LAXA. ‘Tab. VIII, fig. 4, 4a, 46, 4c.

Corallum simple, turbinate, slightly bent, rather intermittent in its growth, and
appearing to have been adherent. Coste distinct from the basis to the calice, very distant
from each other; those belonging to the first three cycla subcrestiform ; those of the last
cyclum flat and scarcely visible, delicately granulated and crossed by small horizontal
strie. Calice circular; fossula narrow and rather deep. No columella. Septa forming
four complete cycla; but those of the last cyclum almost rudimentary. The six systems
equally developed. The septa very unequally developed, broad, very exsert; thin, but
rather less so near the inner margin, presenting a few round granulations on their lateral
surfaces. Those of the first and second cycla united along the lower part of their inner
edge. Height, from one inch to one inch and a half; diameter of the calice, seven lines.

We have given the generic name of Celosmilia to a certain number of Kusmiline which
we formerly placed in our genus Parasmilia, but which are characterised by the absence
of the columella and the rudimentary state of the endotheca. Parasmilia poculum,
P. Faujasi, and P. punctata’ belong to this group, and differ from C. dara by their coste
being flat and granulated near the calice, whereas in the above-described fossil these parts
are subcrestiform. It is also to be remembered that in Calosmilia poculum and C. Fawast
the septa form five complete cycla, and that in the last-mentioned species, as well as in
C. punctata, the principal septa are much thicker than in C. Jaza. M. Alcide d’Orbigny
has lately discovered in the white chalk of Césanne a new species which he designates by
the name of Celosmilia Hdwardsiana, and which differs from C. axa by its coste being
rudimentary and its septa thinner.

1 See our Monograph of the Astreidz (Ann. des Scien. Nat. 3™ série, vol. x). It is possible that our
Celosmilia punctata may be only a young form of C. Faujasi, but we have not as yet seen a sufficient
number of specimens to be able to decide the question.

CORALS FROM THE LOWER CHALK. 53

CHAPTER IV.

CORALS FROM THE LOWER CHALK.

Tue number of British Corals known to belong to this formation 1s as yet so very
small, that it would be premature to speculate on their mode of distribution. We have
seen but two species, one appertaining to the family of Oculinide, the other to that
of Eupsammide ; both appear to be peculiar to the lower chalk of England.

Family OCULINID (p. xix).
Genus SYNHELIA (p. XX).
SYNHELIA SHarpeana. Tab. IX, fig. 3, 3a.

Corallum composite, dendroid, with thick, erect branches, forming acute angles with
each other, and presenting on their surface large, non-exsert, circular calices, which are not
closely set, and are united by rather mdistinct, small costal strie. Calices quite superticial,
and presenting scarcely any central depression. Co/wmel/a assuming the appearance of a
small, obtuse tubercle. Three complete cycla of septa, and in one half of each system two
quaternary septa, of which no homologues exist in the other half. The septa are thick,
very closely set, almost straight, and unequally developed, but those of the second order
differ but little from the primary ones. The upper edge of all is horizontal, and closely
denticulated ; towards the columella the denticulations are rather larger thah towards
the calicular margm, and we have not been able to decide whether some of them do
not constitute pali. The lateral surfaces of the septa present oblong transverse granu-
lations, which much resemble incomplete synapticule, but they are not prominent enough
to meet those of the adjoining septa, and to subdivide the interseptal loculi. The height
of the specimen here describec is about two inches and a half, and the diameter of the
calices two lines.

We are as yet acquainted with but two other species that can be referred to our genus
Synhelia ; one is the &. yibdosa, which was first described by Goldfuss under the name of

54, BRITISH FOSSIL CORALS.

Lithodendron gibbosum,! and which belongs also to the lower Chalk formation, but is found at
Bochum, in Westphalia, and at Blaton, near Mons, in Belgium. It differs from 8. Sharpeana,
by its calices beg more closely set; rather oblong, with a more prominent margin, and
twenty-four nearly equal, very thick septa, separated by an equal number of rudimentarv
ones. The other is the Madrepora Meyeri, found by MM. Koch and Dunker in the
Jurassic formation at Elligser-Brinke ; it has deep calices.”

The unique specimen here described appears to have been found in the lower chalk
near Dover, and was kindly communicated to us by Mr. Daniel Sharpe.

Family EUPSAMMID/® (p. li).
Genus STEPHANOPHYLIIA (p. lin).
STEPHANOPHYLLIA BoweErsBanku. ‘Tab. IX, fig. 4, 4a, 46, 4c.

STEPHANOPHYLLIA BowERBANKII, Milne Edwards and J. Haime, Monogr. des Eupsammides,
in Annales des Sciences Naturelles, 3™° série, Zool.
vol. x, p. 94, 1848.

Corallum simple, resembling, in its general form, a plano-convex lens. WVai/
discoidal and horizontal. Coste numerous, delicate, nearly equal, closely set by pairs, and
formed by a simple series of granulations, which become the most distinct near the outer edge
of the mural disc. ‘Twenty-four of these costa begin near the centre of the corallum, and soon
after bifurcate ; the forty-eight coste thus produced soon divide again, in the same manner,
and near the edge of the disc the number of these radiate ridges amounts to ninety-six.
The mural pores are small, not very distinct, and arranged in series in the intercostal
furrows. Calice quitc circular, and appearing to be regularly convex, excepting towards
the centre, where there is a slightly-marked, shallow fossula. Co/wmel/a almost rudimentary,
and formed only by two or three trabicule, which are often scarcely distinct from the edges
of the septa. These last-mentioned organs arise from the upper surface of the mural disc,
and are thin, especially outwards, closely set, and covered laterally with large, prominent
granuletions. ‘They form five complete cycla, and represent six well-characterised and
equally-developed systems. The primary and secondary septé are straight, and extend to
the columella; their upper edge is arched, or slightly angular. The tertiary septa are also
much developed, and bend towards the secondary ones, to which they become united by
their inner edge, near the columella. The septa of the fourth and fifth orders, constituting
the fourth cyclum, are united in a similar way to the tertiary septa, at about half way from
the margin of the mural disc to the columella, but not exactly at the same point, those of

1 Petref. Germ., vol. i, tab. xxxvii, fig. 9.
* Beitrage zur Kenntniss des Norddeutschen oolithgebildes, p. 55, tab. vi fig. 11, 1837.

CORALS FROM THE LOWER CHALK. 55

the fifth order being rather longer than those of the fourth order. The septa of the fifth
cyclum are small, thin, low, and unite to the neighbouring principal septa; those of the
sixth order join the primary ones; those of the seventlr order adhere by their inner and
upper edge to the secondary ones, and those of the eighth and ninth orders to the tertiary
septa; or, in other words, each element of this fifth cyclum joins the eldest of the two
septa between which it is placed. Independently of these junctions, which are normal, and
always take place along the inner edge of the septa, the interseptal loculi are irregularly
divided in some places by the projecting lateral granule of two neighbouring septa
meeting, and becoming cemented together. By this character, as well as by its general
form, this species tends to -nite the family of Eupsammide with the Fungide.

Height of the corallum, one and a half or two lines; diameter, three or four lines.
Some specimens, which were probaky not udult, were only two lines and a half in
diameter.

This delicate little Coral differs from Stephanophyllia elegans, S. imperialis, and
S. discoides’ by the form of the septa, which do not appear to be angular and laccrated,
as in the three latter species. Stephanophyllia astreata’ differs from it by having a large
fossula and a well-developed columella. It most resembles S. swecica ;* but in this species
the two tertiary septa of each system unite below the columella and the secondary septa,
which consequently do not extend to the centre of the calice; whereas in 8. Bowerbankii
these tertiary septa, as we have already stated, adhere to the secondary septa, and these
last-mentioned septa extend to the columella. The Fossil Coral figured by M. von
Hagenow, under the name of Fuagia clathrata,* and found by that geologist in the chalk
formation of Rugen, is evidently very nearly allied to the British species here described :
but as far as we can judge of it by M. von Hagenow’s engraving, it appears to differ from
it by its more elevated form, by the strongly-marked concentric striz visible on the mural
disc, and by its basis being more prominent.

We must also remark, that the section of the genus Stephanophyllia, to which this
species belongs, and to which we applied the name of Lenticular Stephanophyllia,’ las of

1 See our Monograph of the Eupsammidze, Ann. des Sc. Nat., 3° serie, vol. x.

2 Fungia astreata, Goldfuss, Petref. Germ., vol. i, p. 47, tab. xiv, fig. 1 (where it is by mistake
designated under the name of Fungia radiata). This species not having been, as yet, well characterised,
we think it may be useful to give a short description of it here. Corallum simple, very short, and having
the form of a plano-convex lens. Calicular fossula circular, and well developed. Coste very delicate
and not closely set. Septa forming five complete cycla, and appearing to be thin and strongly granulated.
Size very variable ; in the adult, diameter three lines, height about one line. Fossil found at Aix-la-Chapelle,
in Westphalia, and existing in the Museums of Bonn and Paris. All the specimens yet found are in a very
bad state of preservation.

3 Monogr. of the Eupsammide, loe. cit., p. 94.

4 In Leonhard and Bronn’s Jahrbuch fiir Mineralogie, 1840, p. 684. tab. ix, fig. 3.

> Monogr. of the Eupsammidee, loc. cit., p. 94, 1848.

56 BRITISH FOSSIL CORALS.

late been considered by M. Alcide d’Orbigny as deserving to be elevated to the rank of a
genus, and has been named by that author Discopsammia ;' but M. d’Orbigny has not
pointed out any new characters in addition to those on which this separation was primitively
established in our Monograph, and consequently we see no reason for altering the
classification previously adopted.

Stephanophylia Bowerbanku is found in the lower chalk near Dover, and does not
appear to differ from some corals which one of us’ has lately met with in a bed of chlorited
chalk at Orcher, near le Havre. ‘The specimens here described belong to the collections of
Mr. Bowerbank, Mr. D. Sharpe, and the Geological Society.

1 Note sur les Polypiers Fossiles, p. 10, 1849.
2M. Jules Haime.

CORALS FROM THE UPPER GREENSAND. 57

CHAPTER V.
CORALS FROM THE UPPER GREENSAND.

Tue class of Polypi had not, in all probability, numerous representatives in the seas
where the Upper Green Sand was deposited, for we have as yet seen only four British
species belonging to that formation, and the English geologists do not appear to have met
with many more. Most of these fossils belong to the family of Astreide, and have been
found at Haldon, at Blackdown, or at Warminster. One of these British species
appears to be identical with a coral described by Goldfuss, and found in the chalk forma-
tion of Essen; and Mr. Morris has pointed out two others as being referable to species
found in the chalk of Maestricht, but we have not had an opportunity of recognising the
specific identity of these last-mentioned fossils.

Family ASTREID/E (p. xxii).
Tribe EUSMILIN # (p. xxiii).
1. Genus PEPLosMILiA (p. xxv).

Peptosminia Avstent. Tab. X, fig. 1, la, 14.

Corallum simple, fixed by a broad basis, cylindrical, and surrounded from top to
bottom by a membraniform epitheca, presenting some slight transverse folds. Calice
circular, or somewhat oval; fossula shallow, narrow, and elongated. Columella well
developed and lamellar. Septa appearing to form four well-developed cycla, and a fourth
rudimentary one. ‘The primary and secondary ones equal, and differing but little from
the tertiary ones; they are all thick, broad, closely set, slightly exsert, not quite straight,
those on one side inclining to the right near the columella, and those of the other side
bending in an opposite direction. A vertical section of this Coral (fig. 14) shows that
the septa are granulated on their lateral surfaces, especially near their inner edge, which
joms the columella, and that these granulations form closely-set radiate rows. Dzssepr-
ments vesicular, and rather abundant. Height of the coral, one inch and a half; diameter

of the calice, above an inch.
8

58 BRITISH FOSSIL CORALS.

This species is as yet the only known representative of our genus Pepiosmilia,! and is
easily distinguished from the other true Eusmiline, either by its lamellar columella or its
complete epitheca; it may be considered as a Montlivaltia, having a lamellar columella.
We have seen but one specimen of this fossil ; it was found in the Greensand at Haldon,

and presented to the Geological Society by Mr. R. H. C. Austen.

2. Genus TROCHOSMILIA (Dp. XXIV).
TRocHosMILiA (?) TuBEROSA. ‘Tab. X, fig. 2, 2a.

TURBINOLIA CoMPRESSA, (?) Morris, Cat. of Brit. Foss., p. 46, 1843.

Corallum simple, compressed, even at its basis, cuneiform, subpedunculated, and pre-
senting on each of its lateral edges, at a short distance above the basis, a broad but not
very prominent tuberosity. Coste delicate, straight, not prominent, but very distinct from
the basis upwards, closely set and somewhat unequal. Cadice elliptic and horizontal; its
small axis only half the length of the long axis. Fossula narrow, rather shallow, and
elongated. No columella. Septa forming five complete cycla; very thin, straight, closely
set, and delicately granulated laterally ; those of the first and second cycla nearly equal in
size and larger than the others, so as to produce the appearance of twelve systems; those
of the fifth cyclum very small. Height, seven lines; diameter of the calice, eight lines by
four.

The above-described specimen was found in the Greensand of Blackdown by our able
friend Mr. J. S. Bowerbank. We have not, as yet, been able to ascertain the existence of
dissepiments in the interseptal loculi, and consequently are not quite sure that it belongs
to the genus Trochosmilia; if these parts do not exist it must be referred to the family of
the Turbinolidz, but we have not had the materials necessary for deciding that question.
We shall therefore only add here, that this coral differs from the other species of ‘Trocho-
smilia described in our ‘ Monograph of the Astreide’ by the existence of the lateral tubero-
sities, and the basis presenting scarcely any traces of adherence.

It is probably this fossil which Mr. Morris referred to the Zurbinolia compressa of
Lamarck, and mentioned as existing in the Greensand of Blackdown. 7. compressa belongs
also to our genus 'l'rochosmilia, and is found in the Greensand at Uchaux in the South of
France, but differs from 7. tuberosa by its general form.

1 The fossil described by M. Michelin under the name of Anthophyllum detritum (Icon. Zooph., tab. x,
fig. 1) might at first sight be supposed to belong to this genus, for it presents some appearance of a lamellar
columella ; but that is owing to the presence Of some extraneous matter adhering to the specimen figured by
M. Michelin, and although the epitheca does no longer exist in this fossil, we have no doubt that it is in
reality a Montlivaltia.

CORALS FROM THE UPPER GREENSAND. 59

Tribe ASTREIN A (p. xxxi).

Genus PaRASTREA (p. xlui).

ParastreEA stricta. Tab. X, fig. 3, 3a.

Corallum composite, forming a mass not very tall, and slightly convex on its upper
surface. Calices seldom circular, in general oblong or irregularly polygonal, projecting
very little, and having always distinct margins. Coste delicate, closely set, nearly equal,
almost horizontal, nearly straight or slightly bent, and united by their extremity to those of
the neighbouring corallites, which, however, remain circumscribed by a small furrow.
Calicular fossula shallow. Columella of a dense tissue, subpapillose, and not much developed.
Septa thin, broad, closely set, terminated by a series of calicular dentations, the last of
which (towards the columella) appears to be more developed than the others ; the number
of these septa seldom exceeds forty, and they are rather unequal. Mails thin, but well
developed. Diameter of the calices, usually between two lines and two lines and a half ;
distance between the calices, at least half a line.

This species, found in the Greensand at Blackdown, is characterised from a specimen
belonging to the Geological Society; it differs from all the previously described Parastrea
by the approximation and delicate structure of the septa.

Mr. Morris mentions, in his ‘ Catalogue of British Fossils,” two other species which
have been found by M. Austen in the Greensand at Haldon, and which belong to the family
of Astreide. M. Austen considers the one as being identical with the Maestricht
fossil coral described by Goldfuss under the name of Astrea elegans,’ and he refers
the other to the Astrea escharoides* of the same author*. We regret not having had
an opportunity of examining these fossils.

1 Loe. cit., p. 31.

2 Petref. Germ., vol. i, tab. xxiii, fig. 6.

3 Goldfuss, op. cit., tab. xxiii, fig. 2; fossil from Maestricht.

* Austen, on the Geol. of the South-east of Devonshire, Trans. of the Geol. Soc., Second Series, vol. vi,
p. 452.

60 BRITISH FOSSIL CORALS.

Family FUNGID/ (p. xlv).
Genus MicraBacta (p. xvii).
MicraBacta coronuta. ‘Tab. X, fig. 4, 4a, 46, de.

Cycio.ites, WV. Smith, Strata identified by Organic Fossils, p. 12; Greensand, p. 15, 1816.
FUNGIA CORONULA, Goldfuss, Petref. Germ., vol. i, p. 50, tab. xiv, fig. 10, 1826.
= = F.A. Remer, Die Verstein. des Norddeutschen Kreidegebirges, p. 25, 1840.
— — Morris, Cat. of Brit. Fossils, p. 38, 1843.
FunGra cLaTHrata (?) Geinitz, Grundriss der Versteinerungskunde, tab. xxiii, fig. 2, 1849.

Corallum simple, lenticular, short ; its under surface horizontal or slightly concave ; its
upper surface somewhat convex. Mural disc completely naked and regularly perforated
by small intercostal pores. Coste closely set, almost straight, equally narrow, not pro-
minent, and but slightly echinulated ; only twelve of them arise in the centre of the dise,
but these soon bifurcate, and the twenty-four costa so formed soon divide again; at about
half the distance from the centre to the circumference of the disc each costa bifurcates once
more, and the two terminal costee so formed are grouped two by two towards the periphery
of the disc. The granulations which form all these coste are not very distinct, and are
arranged in single lines. Cadiculur fossula small and not very deep, but well marked and
rather elongated laterally. Colwmella very small, oblong, and subpapillose. Septa forming
five complete eycla, and corresponding to the intercostal spaces; those of the last cyclum
quite rudimentary ; the others tall, thin, straight, and united by sub-spiniform trabicule.
Those of the first cyclum larger than the others, and augmenting slightly in thickness
towards the middle; the secondary ones almost as large; all delicately denticulated along
their upper edge, and much thinner towards their outer and inferior angle than in any
other part. Diameter, three or sometimes four lines ; height, one line and a half.

The above-described fossils were found in the Greensand at Warminster, in Wiltshire,
and according to William Smith, who was the first author that mentions this fossil,
are also met with at Chute Farm and Puddle Hill, near Dunstable.

By an attentive comparison with the specimens described by Goldfuss, and belonging
to the Poppelsdorff Museum at Bonn, we have ascertained the specific identity of this
British Coral with the Fungia coronula found in the chalk of Essen. Specimens exist in
Mr. Bowerbank’s cabinet, and in the collections belonging to the Geological Society,
the. Museum of Paris, the Museum of Bonn, and M. Defrance at Sceaux, who has

designated it by the unpublished name of Fungia dubia.

CORALS FROM THE GAULT. iH]

CHAPTER, VIE
CORALS FROM THE GAULT.

Tue Fossil Corals contained in the Gault are more numerous than those imbedded i
the upper greensand and the lower chalk. Most of them belong to the family of
Turbinolide, and the principal localities where they have been met with in England are
Folkstone and Cambridge.

Family TURBINOLIDE (p. xi).
Tribe CYATHININA (p. xii).

1. Genus CYAtHINA (p. xi).
Cyatnina Bowersankil. Tab. XI, fig. 1, la, 1d.

CyaruHina BowEersBankul, Milne Edwards and J. Haime, Monogr. Turbin., in Ann. des Sc.
Nat., 3™° serie, vol. ix, p. 292, 1848.

Corallum simple, elongated, turbinate, very narrow, and slightly bent near the basis.
which does not appear to have expanded much. Vall quite naked. Coste almost flat,
distinct from the basis, or nearly so, covered with small granulations, nearly equal, and
showing a slight tendency to form binary groups. Calice circular. Columella not much
developed, and composed of twisted blades. Septa forming four complete cycla; very
thin, but slightly granulated, and rather unequal. Those of the last cyclum very little
developed, and the tertiary ones rather thickened towards the inner edge. Pali covie-
sponding to the penultimate cyclum of septa, and rather broad. Height of the coral.
eight or nine lines; diameter of the calice, three lmes and a half.

This fossil was found in the Gault at Folkstone, by our friend Mr. Bowerbank. All
the specimens that we have seen were very incomplete, but some showed all the principal
characters represented in the figures which we have given.

C. Bowerbankii is easily distinguished from C. Siithit and C. pseudoturbinolia, by uot
having a fifth cyclum of septa. It differs also from C. arcuata by the delicacy of its septa.
and from C. Guadulpensis by the circular form of its calice, and its round columella.

62 BRITISH FOSSIL CORALS.

C. cylindrica, C. Breda, and C. Koninckii, have only six large pali, whereas in C. Bowerbankit
the number of these organs amounts to twelve. C. devigata! differs from the above-
described species, by the pali being narrow, and very thick, and C. Dedeyana by the
existence of a well-marked epithecal band near the calice.

M. Alcide d’Orbigny has, in a recent publication,’ referred to this species as the type
of his new genus Amblocyathus, which he defines as being Cyathina, with a circular calice
and a round columella. He adds that Amblocyathus is a lost genus, and contains three
fossil species belonging to the Neocomian and Albian’ strata. We must, however, beg
leave to remark, that the two above-mentioned characters are met with in almost every
species of our Cvyathina, and most especially in C. cyathus, which is the type of the genus
Cyathina, and is actually living in the Mediterranean sea. Only two of the species
referred to the genus Cyathina in our ‘Monograph of the Turbinolide’ present a slightly
oval calice and a transversal columella—C. pseudoturbinohia and C. Guadulpensis. In
C. Snuthii the columella is oblong, but the calice is circular, or nearly so. If it be con.
sidered necessary to separate the Cyathina with a circular calice from those that have an
oval calice, it would therefore be more proper to give a new generic name to the latter, and
not to change the denomination of the group containing the very species for which
Khrenberg first established the genus Cyathina. But this innovation, proposed by
M. d’Orbigny, appears to us as being in every respect unnecessary, for the slight deformation
of the calice and the columella which forms the sole basis of the new generic division, can
hardly be considered as characters of sufficient value; species that differ in no other
respect are often found to vary in this way, and even specimens belonging to the same
species sometimes differ much in the form of the calicular margin. ‘Thus, although the
calice is circular, or nearly so, in most specimens of C. cyathus and C. Smithii that are met
with, we have seen some that were compressed, and had the calice as oval as in C. pseudo-
turbinolia and C. Guadulpensis ; similar deviations from the normal form ave also to be -
met with in the columella; in C. Smithii, for example, this organ is sometimes quite
circular, although it is in general oblong. Differences of this kind, when not more marked
than is the case among the various species of Cyathina, can therefore scarcely be deemed
important enough to characterise generic divisions; and, as in the present case, they do
not appear to coexist with any other structural peculiarity, we see no reason for admitting
the new genus Amblocyathus.

1 Tab. ix, fig. 1.
? Note sur des Polypiers Fossiles, Paris, 1849.
3 M. d’Orbiguy employs the name of Aldian formation to designate the Gault.

CORALS FROM THE GAULT. 63

2. Genus CYCLOCYATHUS (p. Xiv).
Cyctocratuus Firronrt. Tab. XI, fig. 3, 3a, 36.

Corallum simple, discoidal, short ; mural disc horizontal, or slightly concave, and pre-
senting in its centre a small, irregular cicatrix, indicative of its primitive adherence.
Epitheca very thin, presenting some slight concentric striz, and not preventing the radiate
costz from being visible. These are straight, and not very prominent; but those of the
first and second order are well marked. The edge of the mural disc is thin, and slightly
prominent. The upper or calicular surface of the corallum is rather convex externally,
and concave towards the centre. The fossula is shallow, but large, and well marked.
Columella fasciculate, well developed, and terminated by a broad, papillose surface. Septa
forming four complete cycla. The six fundamental septal systems distinct, but the septa
of the second order not differing much from those of the first order. All the septa well
developed, straight, rather thick exteriorly, arched above, and granulated laterally ; their
outer edge somewhat crenulated, granulose, slightly concave near the mural disc, and pro-
jecting a little towards the upper part. Padi well developed, very distinct from the septa,
and corresponding to those of the third cyclum. Height of the corallum, two or three lines ;
diameter, in general not more than five or six lines.

This fossil is the only known species of the genus Cyclocyathus ; its form renders it
very remarkable. It has been found in the Gault at Cambridge, Drayton, West Malling,
and Folkstone, but appears to be most abundant in the last-mentioned locality. The
specimens here described belong to the collections of the Geological Society, of Mr.
Bowerbank, and of Mr. D. Sharpe.

3. Genus TROCHOCYATAUS (Pp. XIV).

1. Trocnocyatuus conutus. Tab. XI, fig. 5, 5a.

CARYOPHYLLIA CONULUS, (?) Phillips, Ilust. of the Geol. of Yorkshire, tab. ii, fig. 1, 1829.
(A rough figure without any description.)
— — Michelin, Mém. de la Soc. Géol. de France, vol. iii, p. 98, 1838.
TURBINOLIA ConuLUSs, Michelin, Icon. Zooph., p. i, pl. i, fig. 12, 1840.
TRocHOCYATHUS conuLUS, Milne Edwards and J. Haime, Monogr. des Turbin., Ann. des
Se. Nat., 3™° série, vol. ix, p. 306, 1848.

Corallum simple, turbinate, rather elongate, straight or slightly bent, and pedicellated.
Wail presenting in general some slight traces of an incomplete epitheca. Coste simple,
distinct from the basis, closely set, delicately granulated, not very prominent, and alternately

64 BRITISH FOSSIL CORALS.

of unequal size towards the calicular edge. Calice almost circular, or somewhat oval and
shaliow. Columella fascicular, well developed, not prominent at its apex, and terminated
by ten or fifteen papillee of equal size. Septa forming four complete cycla and six well-
marked, equally developed systems, i which, however, the secondary ones differ but little
from those of the first cyclum. The septa are slightly exsert, closely set, unequal, and
rather thicker outwards than towards the columella. Padi narrow and unequal; those
corresponding to the tertiary septa broad and rather stout; the others, and most especially
those corresponding to the primary septa, narrow and thinner. Height of the covallum,
seven or eight lines ; diameter of the calice, almost seven lines.

This species belongs to the first section of the genus Trochocyathus (Z! simplices), and
differs from 7. impari-partitus' and T. Bellingherianus by not having a fifth cyclum
of septa; its general form distinguishes it from 7. mitratus,’ T. crassus, T. simplex, and
T. costulatus, which are all short, broad, and curved ; and from 7. elongatus, T. Koninchit,
and 7. gracilis, which are much elongated, curved, and very narrow towards the basis. It
appears to resemble most, especially by its general form, 7. cupwla,’ which is also conical
and straight, but this last-mentioned species differs from it by the thickness and strong
granulations of the septa, and by the breadth of the basis.

Trochocyathus conulus appears to have been very widely spread in the seas where the
Gault formations were deposited. ‘he specimens which we most particularly studied were

’ See our Monograph of Turbinolide, loc. cit., p. 307.

? Since the publication of our Monograph of the Turbinolide (in 1848) we have recognised that the
fossils from Tortona, which M. Michelotti designates under the name of Turbinolia plicata, do not differ
specifically from the specimens existing in the Poppelsdorf Museum under the name of Turbinolia mitrata,
Goldfuss. As we already expected, the latter specific name must therefore be substituted for the one
employed by M. Michelotti, and M. Michelin.

3 This new species, designated under the name of Turbinolia cupula, by M. Alex. Rouault, (Bulletin
de la Soc. Géol. de France, 2" série, vol. ix, p. 206, 1848), was found by that geologist at Bos d’Arros, in
the department of the Lower Pyrennees, and does not appear to differ from a fossil which exists in the
collection of M. Nyst, and was found in the Eocene formation at Lacken, near Brussels. Trochocyathus
cupula belongs to the first section of our genus Trachocyathus, and presents the following characters :

Corallum straight, or almost so, subturbinate, but short, and having a broad peduncle, but not remaining
adherent in the adult state. Coste distinct from the basis, straight, unequally developed alternately,
rather prominent, especially near the calice, granulated and striated transversely ; rudimentary cost, that
do not correspond to any septa, are seen in the intercostal furrows. Calice circular; fossula not deep.
Columella cr’ te, well developed. Septa forming three complete cycla, and in general a fourth incom-
plete cyclum in one half of three of the systems; exsert, rather unequal, strong, and presenting on their
lateral surfaces large prominent granulations, which are arranged in lines nearly parallel to the upper edge.
Pali thick, strongly granulated, and unequal; those corresponding to the tertiary septa the largest in the
half systems where the septa of the fourth cyclum exist, and those corresponding to the secondary septa
most developed in the other part of the calice. Height of the corallum, three lines; diameter of the calice
almost as much. By the strong granulations of the septa, and the breadth of its basis, this species tends to
establish a transition between the genus T’rochocyathus and the genus Paracyathus.

CORALS FROM THE GAULT. 65

found near Cambridge, in England ; at Gatis de Gerodot, Dienville, near Brienne (department
of the Aude), and Etrepy (department of the Marne), in France. Other specimens, which
in all probability belong also to this species, are designated in M. Michelin’s collection as
having been found at Novion-en-Porcien ; at Macheromenil, in the Ardennes, and at the
Perte du Rhone, in the department of the Ain; but we suspect that some mistake may have
been made in the labelling of the specimen which is designated in the same collection as
belonging to the chalk of Tournay, in Belgium. We must also add, that the fossil designated
by Professor J. Phillips under the name of Zurbinolia conulus was found by that eminent
geologist at Speeton, in Yorkshire ; but its characters are not sufficiently well known for us
to be able to identify it with the above-described species, specimens of which exist in the
collections of the Geological Society, of the Museum at Paris, and of MM. d’Orbigny,
Michelin, and Milne Edwards.

M. Al. d’Orbigny has lately given the name of Aplocyathus' to those species of our
genus Trochocyathus in which the calice is circular. If this new generic division was
adopted, the species here described would be referred to it ; but that is not, in our opinion,
advisable. The calice, which is quite circular in a great many species of our genus
Trochocyathus, becomes slightly elongated in some, quite elliptical in others, and not only
would the line of separation be difficult to establish between these different forms, but
certain species which are evidently most closely allied by all their other organic
characters, would be. separated generically in the classification proposed by M. d’Orbigny.
We cannot, therefore, adopt his views in this respect ; but, m justice to that distinguished
paleontologist, we must remark that the species’ chosen by him as the type of his
genus Aplocyathus differs much in its general aspect from most species of our genus
Trochocyathus, and, when more completely known, may be found to present characters of
sufficient value to authorise the establishment of a separate generic group, which must
then be so defined as not to comprehend 7. conulus, nor most of the other species that
have a circular calice.

2. TRocnocratuus Harveyanus. Tab. XI, fig. 4, 4a, 40.

TrocHocyatuus Harveyanus, Milne Edwards and J. Haime, Monogr. des Turbinolides, in
Ann. des Sc. Nat., 3" série, vol. ix, p. 314, 1848.

Corallum simple, straight, short, almost hemispherical, and terminated by a very short
peduncle, the basal surface of which is concave. Coste distinct from the basis, and
delicately striated transversely ; the primary and secondary ones very prominent and sharp ;
those of the third cyclum well developed along the upper half of the wall, but those of
the fourth cyclum very small and obscure. Calice circular and flat; fossula shallow.
Columella well developed and papillose. Sepfa forming four complete cycla; exsert, thin,

' Note sur des Polypiers Fossiles, p. 5, 1849. 2 The Trochocyathus armatus.
9

66 BRITISH FOSSIL CORALS.

broad, straight, granulated laterally, unequally developed, but not differing much in the
first and second cycla. Pali corresponding to the septa of the first three cycla, rather
narrow, and unequally developed.in an inverse ratio to the corresponding septa ; no pali in
the radii of the septa belonging to the fourth cyclum. Height of the corallum, three lines ;
diameter of the calice, four lines.

This species belongs to the fourth section of our genus Trochocyathus (7. dreves), and
consequently its characters need not be compared with those of the various species belonging
to the sections of the 7. simplices, T. cristati, and T. multistriati, the description of which
may be found in our ‘ Monograph of the Turbinolide.’ It differs from 7. obesus, T. armatus,
and 7. perarmatus,' by not having any costal spines, and from 7. Michelini by the coste
being distinct down to the basis, and by its general form being less depressed. It appears
to be most closely allied to the fossil which we shall next describe under the name of
Trochocyathus (?) Konigi, but is of a more slender form.

T. Harveyanus was found in the Gault at Folkstone, the birthplace of the illustrious
physiologist to whom we have dedicated this species. ‘The specimens here described belong
to the collections of Mr. Bowerbank and Mr. D. Sharpe.

3. TrocHocyaTuHus (?) Kontat.

TurBINOLIA Kontel, Mantell, lust. of the Geol. of Sussex, p. 85, tab. xix, figs. 22, 24,
1822.
— — Fleming, British Animals, p. 510, 1828.
— (TRocHOCyATHUS?) Konter, Milne Edwards and J. Haime, Monogr. des Turb.,
in Ann. des Se. Nat., 3™° série, vol. ix, p. 335, 1848.

The specimens of this fossil figured by Mr. Mantell, and those which we have seen in
the collections of MM. d’Orbigny and Michelin, are in a very bad state of preservation,

1 This species, which has been lately designated under the name of Turbinolia perarmata by
M. Talavignes, but has not yet been described, and has been given to us by that geologist, was discovered
at Fabresan, in the department of the Aude. M. Alex. Rouault has since then met with the same species
at Bos d’Arros in the Lower Pyrennees. (See Bull. Soc. Géol., 2™ serie, vol. v, p. 206.) It may be
recognised by the following characters :

Corallum very short, subdiscoidal ; its under surface flat and almost smooth ; sometimes adhering to a
small shell. Coste distinct near the calice, projecting very little, closely set, almost equal, and delicately
granulated ; those of the first cyclum not differing much from the others, but bearing, at a short distance
from the calicular edge, a strong spiniform appendix, which is rather compressed, extends outwards, and
presents, on its under edge, a small pointed tubercle. Calice circular. Septa forming four complete cycla
and six equally developed systems; closely set, rather exsert, thin, and unequally developed ; but those of
the second cyclum differing very little from the primary ones. Pali narrow and rather thick. Height of
the corallum, one line and a half; diameter of the calice, two lines and a half. Fossil from the Nummulitic
formation at Fabresan and Bos d’Arros.

CORALS FROM THE GAULT. 67

and have lost their walls; we are, therefore, unable to characterise the species with
any degree of precision, and it is with much doubt that we refer it to the genus
Trochocyathus, for we are not as yet sufficiently satisfied as to the existence of pali.
M. Michelin is of opinion that these fossils are merely specimens of Zrochocyathus conulus
with their basis worn away. They are of a conico-convex form, and are broader in
proportion than 7. Harveyanus, to which they bear, however, great resemblance. Their
height is about four lines, and their diameter a little more. We have not considered
it necessary to give a new figure of these corals, for the specimens in our possession do not
show anything more than those represented in Dr. Mantell’s plates.

The specimens that we have had an opportunity of examining were found in the Gault at
Folkstone, in the environs of Boulogne-sur-Mer, at Wissant, at Les T'iz, near Chamounix,
and at the Perte du Rhone, in the department of the Ain. According to Dr. Mantell the
same species is met with at Lewes in Sussex, and Godstone in Surrey, at Malling in
Kent, in Cambridgeshire,’ at Ringmer, and at Bletchingley.”

TrocHocyaTHus (?) WaARBURTONI.

We are inclined to think that a cast found in the Gault of Cambridgeshire by
Mr. H. Warburton, and presented by that gentleman to the Museum of the Geological
Society, must belong to a distinct species of Trochocyathus. It is about six lines in
height, and seven in diameter; the number of septa is forty-eight. For the sake of
convenience we have given a specific name to it, but we are not able to characterise it.

A. Genus BatnycYatTuus (p. xill.)

BatuycyatTaus Sowersyi. ‘Tab. XI, fig. 2, 22.

=
Batuycyatuvcs Sowersyt, Milne Edwards and J. Haime, Monogr. des Turbinolides, Ann.
des Sc. Nat., 3™° serie, vol. ix, p. 295, 1848.

Corallum simple, adherent by a broad basis, straight, tall, compressed, and having its
lateral edges somewhat prominent. /MVa/l delicately granulated. Coste not very distinct
in the lower half of the corallum, but becoming rather prominent higher up, especially
those of the first and second orders. Calice elliptical and horizontal, the relative length
of its long and short axis varying much (in one specimen = 100: 170, and in another
= 100:250). FussvJa narrow, and appearing to be deep, but completely filled up with
extraneous matter in all the specimens that we have seen, so as not to enable us to obtain
any knowledge respecting the columella and the pali. It is therefore with some uncertainty
that we refer this species to the genus Bathycyathus, and in doing so we have been guided

1 Geol. of Sussex. 2 Trans. of the Geol. Soc., s. 2, vol. iii, p. 210.

68 BRITISH FOSSIL CORALS.

enly by characters of secondary value, which agree, however, very well with those of the
other Corals belonging to the same generical division. Septa forming four complete cycla ;
exsert, thick exteriorly, but thin inwardly, and presenting but few granulations on their
lateral surfaces. Those of the second cyclum almost as large as the primary ones; the
tertiary ones but little developed, although they correspond to large costa, and not as tall
as those of the last cyclum, which are grouped very closely on each side of the primary and
secondary ones. Height of the corallum, one inch two or three lines; great diameter of
the calice, six or seven lines.

The genus Bathycyathus contains two other species, which are both recent: B. Chilensis'
and B. Indicus,” which differ from B. Sowerby: in having an additional cyclum of septa,
the calice arched, and the costze more developed near the basis. We have seen but two
specimens of this fossil; one, belonging to the collection of Mr. D. Sharpe, is catalogued
as having been found in the Gault near Folkstone; the second, belonging to the museum
of the Geological Society, is referred with doubt to the upper greensand of Kidge, in

Wiltshire.

Family ASTREID Ai (p. xxi).

Tribe EUSMILIN AB (p. xxii).

1. Genus TROCHOSMILIA (p. XXIV).
TrocnuosmiLia sutcata. ‘Tab. XI, fig. 6, 6a, 64.

Corallum simple, turbinate, straight, tall, much compressed, subpedicellate, and ap-
pearing to be free. Wall presenting on each side two deep longitudinal furrows. Coste
distinct from the basis, slightly prominent, closely set, and unequal, especially towards their
upper end. Cadice elliptical, sublobulated, and slightly arched ; its long and short axis in
the proportion of 100:200. Fossula very narrow, elongated, and not very deep. No
columella. Septa forming four cycla or more, rather unequal, closely set, thin, and slightly
exsert. Dissepiments not numerous. Height of the corallum, nearly one inch; diameter
of the calice, six or seven lines by three; depth of the fossula, two lines and a half.

We have seen but one specimen of this fossil, which, although somewhat weather-worn,
appeared sufficiently distinct from all other species to authorise us in giving it a peculiar
specific name. It differs from Zrochosmilia didyma’ by its calice being straight, and not

1 See our Monograph of Turbinolide, tab. ix, fig. 5.
2 Loc. cit., tab. ix, fig. 4.
3 Turbinolia didyma, Goldfuss, Petref. Germ., vol. i, tab. xv, fig. 11.

CORALS FROM THE GAULT. 69

bent in two; from 7. Boissyana, T. Patula,’ T. cernua, and T. crassa,’ by being sub-
pedicellated, and not adherent in the adult state; from 7. irregularis, T. corniculum,
T. Fayasi, T. Gervilli, and T. uricornis, by being strongly compressed quite down to
the basis; and from 7. Saltzburgiensis, T. cuneolus, T. compressa, T. complanata,
T. Basochesu, and T. tuberosa, by the existence of the above-mentioned four deep mural
furrows. By their general form, all these corals much resemble many species belonging
to the division of Cyathinine, but differ from them, and from all other Turbinolide, by
having interseptal dissepiments.
This fossil was found in the Gault at Folkstone, by Mr. Bowerbank.

The LirHopENDRON GRACILE, Goldfuss,* is mentioned by Mr. Morris’ as having been
found in the Gault of Kent, but as yet we have not met with any specimens of that specics
in any of the British palzeontological collections.

+ See our Monogr. des Astreides, Ann. des Sc. Nat., 3** serie, vol. x, p. 236.

2 We here designate, under the name of Trochosmilia crassa, the fossil described by M. Michelin under
the name of Turbinolia cernua, Goldfuss, and by ourselves as Trochosmilia cernua ; for, on comparing it with
the specimens previously described by Goldfuss under the name of Turbinolia cernua, we have ascertained
that they are not specifically identical.

The species which must retain the name first applied by Goldfuss presents the following characters :

Corallum pedicellated and strongly compressed quite from the basis. Coste thin, alternately unequal ;
the larger ones rather prominent and somewhat lamellar. alice arched and elongated in the proportion
of 100: 230. Septa thin, very closely set, and presenting on their lateral surfaces a great number of
granulations arranged somewhat regularly in convex lines parallel to the upper edge. Forty-eight principal
septa, separated by an equal number of small ones; some indications of an additional rudimentary cyclum.
Height of the corallum, one inch and a half; long diameter of the calice, twelve lines ; short axis, five lines.
(The figure given by Goldfuss, tab. xv, fig. 8, is not quite accurate.)

° Monogr. des Astreides, loc. cit.

* Petref. Germ., vol. i, tab. xiii, fig. 2.

> Catalogue of British Fossils, p. 40.

70 BRITISH FOSSIL CORALS.

CHAPTER VIL.
CORALS FROM THE LOWER GREENSAND.

Tue remains of true Polypi are very rare in this part of the British geological strata ;
the fossil which Mr. Lonsdale has lately described under the name of Choristopetalum impar,’
and which was found in the lower greensand at Atherfield, does not appear to us to
belong to this class, and is, in our opinion, a Bryozoon. We have as yet met with but

one species of Zoantharia, which can be referred with any degree of certainty to this
formation.

Family STAURID/ (p. Ixiv).

Genus. Hotocystts (p. lxiv).

Hotocystis rteaans. ‘Tab. X, fig. 5, 5a, 56.

AstrEA, Fitton, On the Strata below the Chalk, in Geol. Trans., s. 2, vol. iv, p. 352, 1843.

ASTREA ELEGANS, Fitton, in Quarterly Journ. Geol. Soc., vol. iii, p. 296, 1847.

CyaTHoPHoRa (?) ELEGANS, Lonsdale, Proceed. of the Geol. Soc., vol. v, part i, p. 83, tab. iv,
fig. 12, 15, 1849.

Corallum complex, astreiform, constituting a convex mass, and augmenting by extra
calicular gemmation ; the young individuals being produced at the poimt of junction of the
surrounding calices. Corallites somewhat prismatic, and cemented together laterally,
either by the direct union of their walls, or by means of the costz, which are thick, and in
general pretty well developed. Calices subpolygonal, separated in general by a simple but
thick mural ridge ; sometimes by walls that remain distinct, and are in their turn separated
by a small intermural furrow. ossula deep. Columella very small, and appearing to be
styliform. Septa forming three complete cycla, and four well-characterised systems. The
four primary ones much more developed than the others, reaching almost to the centre of
the fossula, and giving to the calice a crucial character, which is never met with in Astreide,
Oculinide, 'Turbinolide, &. The septa are slightly exsert, closely set, thick exteriorly,
and very slightly granulated laterally; they appear to have undivided edges, and they
differ much in size, according to the cycla to which they belong. ‘The interseptal dissepiments
are simple, horizontal, or slightly convex, and placed at the same level in the different loculi,

1 Proceedings of the Geol. Soc., vol. v, part i, p. 69, tab. iv, figs. 5 to 11, 1849.

CORALS FROM THE LOWER GREENSAND. 71

so as to constitute by their union a series of complete tabule, subdivided by the primary
septa, and distant from each other about one fifth of a line. Exothecal dissepiments much
resembling the preceding ones. Diameter of the calices, and depth of the fossula, about
one line and a fourth.

Fossil from the lower greensand at Redhill cutting, Atherfield, in the Isle of Wight,
and at Peasemarsh.

The specimens here described belong to the Museum of the Geological Society, and had
been named by Mr. Lonsdale. The propriety of establishing a new generical division for
this remarkable coral, was very judiciously pointed out by that indefatigable paleontologist ;
but, guided by reasons which we do not quite understand, he refers, with a sign of doubt,
this same species to the Cyathophora of M. Michelin, a genus which, in our opinion, does
not differ from true Stylina. The genus Holocystis differs from our genus Stauria by its
extra calicular gemmation, and its costulated walls. It is the most modern representative
of the great division of Zoantharia rugosa, which becomes predominant in the Paleozoic
formations, and is principally characterised by the tendency to a quadrate arrangement of
the constitutive parts of the Corallites, whereas in the other sections of Zoantharia, six is
the fundamental number of the radiate organs.

TAB. I.
CORALS FROM THE CRAG.

‘SSPHENOTROCHUS INTERMEDIUS (p. 2).
Fig. 1. An adult specimen ; natural size.
1a, An adult specimen ; variety having a dilated basis ; natural size.
1 4. A magnified view of the specimen represented at fig. 1.
lc. A vertical section of the same, corresponding to the short diameter of the
calice ; magnified.
| d. Calice of the same; magnified.
le. A very young individual, magnified; the natural size is indicated by the
length of the line placed near this figure.
Lf, 1g, 14, 17. A series of young individuals, at different periods of their growth ;
magnified.

PLabetLuM Wooptt (p. 6).
Fig. 2. <A side view of the corallum ; natural size.
2a. A specimen magnified, and showing the mode of arrangement of the septa;
one half of the calice has been cut away down to the bottom of the fossula.

26. Calice entire; natural size.

BALANOPHYLLIA CALYCULUS (p. 9).
Fig. 3. Two individuals cemented together by their basis ; natural size.

3a. A variety with a narrow basis; natural size.

36. An individual remarkably tall, with its calicular extremity worn away ; natural
size.

3c. Horizontal section of the same, near the calice, magnified so as to show the
mode of arrangement of the septa.

3d. A fragment of the wall deprived of its epitheca, and much magnified.

Crypraneia Wooptt (p. 7).
Fig. 4. A small aggregation of corallites imbedded in a mass of Cellepora; natural

sIZe.

4a. One of these corallites extracted from the mass of Cellepora, and showing its
epitheca ; natural size.

46. One of the same separated from the extraneous mass, and having its epitheca
dimpled by pressure of the surrounding Cellepora.

4c. Calice, magnified.

4d. One of the same corallites magnified, and having part of its calice cut away
so as to show the denticulate edge of the septa; the dimples of the epitheca
are accidental and produced by the investing Cellepora.

Ge ae
et Ail, oie

Pet we PP ow
oy em Pee
e™

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a
2
=
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:
5
+
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oe
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ao
e 3

TAB. IL.
CORALS FROM THE LONDON CLAY.

DIPLHELIA PAPILLOSA (p. 28).

Fig. 1. A branch of this compound corallum ; natural size.

1 a. Terminal portion of one of the corallites, much magnified, and having half of
its calice cut away so as to show the structure of the septa and the thickness
of the walls.

14. Calice, magnified.

OcULINA CONFERTA (p. 27).

Fig. 2. A small mass of this compound corallum ; natural size.
2a. Terminal portion of one of the corallites magnified, and cut down so as to
show the structure of the columella, the pali, the septa, and the thickness
of the walls.
2. Calice, magnified.

TuRBINOLIA BowERBANKII (p. 16).
Fig. 3. Adult specimen ; natural size.
3 a. The same magnified, so as to show the characters of the intercostal furrows.
36. Calice, magnified.

‘TURBINOLIA FIRMA (p. 20).
Fig. 4. Adult specimen ; natural size.
4a, The same, magnified.
46. Calice, magnified.

TURBINOLIA MINOR (p. 19).
Fig. 5. Adult specimen ; natural size.
5a. Magnified view of the same.
5 6. Calice, magnified.

Sea ne iar oe

TAB. II.
CORALS FROM THE LONDON CLAY.

Tursinoiia Dixonit (p. 15).

Fig. 1. Adult specimen ; natural size.

1 a. The same magnified, to show the characters furnished by the coste, and the
mural furrows.

14. Vertical section, showing the columella, the lateral surface of the septa, the
wall, and the intercostal striee.

1 c. Calice, magnified ; the numbers surrounding the septa indicate the cycla to
which each of these belong.

1d. Fragment of the wall in which the lamellar coste have been worn down, so
as to show that the intercostal dimples are produced by transverse dissepi-
ments, and are not pores, perforating the wall.

TuRBINOLIA FREDERICIANA (p. 17).

Fig. 2. Adult corallum; natural size.
2a. The same, magnified.
26. Calice, magnified.

In one of the systems the septa are numbered with reference to the cycla to which
they belong.

TURBINOLIA SULCATA (p. 13).

Fig. 3. An adult corallum; natural size.
3a. An individual showing a variety of forms.
3 6. The first of the preceding corals, magnified.
3c. Calice, magnified.

TURBINOLIA HUMILIs (p. 18).

Fig. 4. An adult specimen ; natural size.
4a. The same magnified.
4 6. Calice, magnified.

By a mistake of the artist, the third cyclum of septa is here represented as being
complete, whereas in reality these septa do not exist in two of the systems.

TURBINOLIA PRESTWICHII (p. 20).

Fig. 5. An adult specimen ; natural size.
5a. The same magnified.
5 6. Calice, magnified.

LEPTOCYATHUS ELEGANS (p. 21).

Fig. 6. Side view of the corallum ; natural size.
6 a. The same magnified.
64. Under surface of the same, magnified.

6c. Calice, magnified ;—1,1,1,1,1,1, Septa of the first order; 2, Septa of the
second cyclum; 3, Septa of the third cyclum; 4, 5, Septa of the fourth
and fifth orders constituting the fourth cyclum.

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TAB. IV.

CORALS FROM THE LONDON CLAY.

PARACYATHUS CRASSUS (p. 23).
Fig. 1. Two adult corals and a young one, cemented together by their basis;
natural size.
la. A young specimen, with its basis spreading very much, viewed from above.
14. A young specimen, magnified to show the structure of the costa.
le. Calice, magnified ;—1,1,1,1,1,1, Septa of the first cyclum.

PARACYATHUS CARYOPHYLLUS (p. 24).
Fig. 2. An adult specimen, complete ; natural size.
2a, 26, 2c. Worn specimens ; natural size.
2d. "wo young individuals cemented by their basis.
2 e. A remarkably short specimen.
2 f. A complete specimen, magnified so as to show the structure of the wall.

2g. Calice, magnified.

PARACYATHUS BREVIS (p. 20).
Fig. 3. A specimen, the base of which is somewhat worn down; natural size.
3a. The same, magnified.

30. Calice, magnified.
3 ¢. Interior cast of a corallum, which is taller than the preceding one, but appears

to belong to the same species.

Dasmt1a SowErsyl (p. 25).
Fig. 4. Side view of the corallum ; natural size.
4a. The same magnified, so as to show the structure of the cost.
44. Calice, magnified ; the central part is filled up with extraneous matter.

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TAB. \.
CORALS FROM 'THE LONDON CLAY.

SvYLOCENIA EMARCIATA (p. 30).

Fig. 1. A small mass of this compound coralluin ; natural size.
la. A portion of the calicular surface of the same, magnified.

This specimen is somewhat weather-worn.

StyLocania MonticuLarta (p. 82).

fig. 2. A small, somewhat gibbose mass of this compound corallum ; natural swe.
2a. A portion of the calicular surface, magnified so as to show the structure of the
calices and of the marginal processes.
26. Transverse section of the compound corallum, slightly magnified, to show the
cavity circumscribed by its under surface.

ASTROCENIA PULCHELLA (p. 33).
ig. 3. A small mass of this compound corallum, m which most of the corallites have

been pressed together so that their calicular edges have become polygonal
and completely united ; natural size.

3a. Another group, in which most of the corallites have preserved their original
circular form and free calicv.ar margin.

36. A portion of the specimen fig. 3, magnified.

3c. A portion of the specimen fig. 3 ¢, magnified.

STEREOPSAMMIA HUMILIS (p. 37).
Fig. 4. A group of corallites ; natural size.
4a. Terminal portion of some of these, magnified.
4.6. Calice, magnified.

TAB. VI.

CORALS FROM THE LONDON CLAY.

BALANOPHYLLIA DESMOPHYLLUM (p. 35).
hig. 1. Side view of a complete specimen ; natural size.
la. The same, magnified.

1d. Calice, magnified.

Le. Side view of the upper part of the same, with half of the calice cut away in
order to show the structure of the septa and the depth of the fossula.
DENDROPHYLLIA DENDROPHYLLOIDES (p. 36).
Fig. 2. A large group ; natural size.
2a. Calice, magnified.

One of the branches, magnified to show the structure of the walls.

2c. Two young individuals that have not yet produced young by gemmation, and
are cemented together by their basis.
STEPHANOPHYLLIA DISCOIDES (p. 34).

Vig. 3.

Side view of a specimen, magnified ; the natural size is indicated by the length
of the line placed below.

3a. Calicular surface, magnified.

36. Inferior surface, or mural disc, magnified.

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TAB. VIL.
CORALS FROM THE LONDON CLAY.

Lirnarxza WapstERri (p. 38).
Fig. 1. A mass of this compound corallum adhering to the surface of a pebble ;
natural size.

la. Calicular surface, magnified.
14. Vertical section of two corallites, magnified to show their mternal structure.
le. A transverse section made at a considerable distance below the calice, and

magnified so as to show the structure of the columella, the septa, and the
wails.

HloLAREA PARISIENSIS (p. 4.0).

fig. 2. A fragment of this cylindroid compound corallum magnified ; the length of the
line placed below indicates its real diameter.

2a. Transverse section of the compound corallum, showing the vertical section of
the corallites, magnified.

Mors costata (p. 42).

Fig. 3. A large specimen ; natural size.
3a. A fragment, magnified.

GRAPHULARIA WrtTrERELL (p. 41).

Fig. 4. Fragments of the quadrangular portion of the sclerobasis ; natural size.

4a. A fragment of the cylindrical portion of the same.

14, 4c. Fragments of both forms, magnified.

4d, 4e. Transverse sections of the same magnified, so as to show their radiate

structure.

WEBSTERIA CRISIOIDES (p. 43).

Fig. 5. Fragment of a branch ; natural size.
5a. Portion of the same, magnified.

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TAB. VIII.
CORALS FROM THE UPPER CHALK.

PARASMILIA CENTRALIS (p. 47).
Fig. 1. A young specimen; natural size.

la. The same, magnified to show the structure of the wall.

14. Vertical section, magnified, so as to show the structure of the columella, the
dissepiments, We.

le. Calice, magnified; 1, septa of the first cyclum ; 2 secondary septa; 3, septa of
the third cyclum ; 4, 5, septa of the fourth and fifth orders, constituting the
fourth cyclum.

1d, 1 e. Specimens remarkable by their great length.

ParasMiLia Mantriii (p. 49).
Fig. 2. Side view of the corallum ; natural size.
2a. ‘The same, magnified.
PARASMILIA SERPENTINA (p. 51).
Fig. 3. Side view of the corallum ; natural size.
3a. Calice, magnified.
36. Portion of the wall, magnified.

C@LOSMILIA LAXA (p. 52).
Fie. 4. Side view of the corallum; natural size.
4a. A specimen, the growth of which has been intermittent ; natural size.
4%, The specimen No. 4, magnified, to show the structure of the wall.
4c. Calice, magnified.

PARASMILIA CYLINDRICA (p. 50).

Fig. 5. Side view of the corallum; natural size.

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TAB. IX.
CORALS FROM THE UPPER CHALK.

CYATHINA L&VIGATA (p. 44).
Fig. 1, 1a, 146. Specimens of different forms; natural size.
1c. Calice of the specimen fig. 1, magnified. It is to be remarked that im this
specimen there are only nine pali; these organs not existing in the half
systems, where the septa of the fourth cyclum are not developed.
1d. Calice of the specimen fig. 1 6, magnified, and showing the twelve pali and
the complete fourth cyclum of septa.

Parasmitia Frrtont (p. 50).

Fi Side view of the corallum; natural size.

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2a. Specimen of a different form, magnified to show the structure of the wall.
24. Calice, magnified.

CORALS FROM THE LOWER CHALK.

SYNHELIA SHARPEANA (p. 93).
Fig. 3. A branch of this compound corallum ; natural size.
3. Portion of the same magnified.

STEPHANOPHYLLIA BowERBANKTI (p. 54).
Fig. 4. Calicular surface ; natural size.
4a. Side view of the same, magnified.
46. Calice, magnified.
4c. Mural disc, magnified.

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CORALS FROM THE UPPER GREENSAND.

PEPLOSMILIA AUSTENI (p. 57).
Fig. 1. Restored figure of the corallum ; natural size.
1 a. Calice; natural size.

14. A broken specimen, showing part of the epitheca, the columella, and the
structure of the septa.

‘TROCHOSMILIA TUBEROSA (p. 58).
Fig. 2. Side view of the corallum ; natural size.
2a. Calice, magnified.
PARASTREA STRICTA (p. 59).
Fig. 3. A mass of this compound corallum ; natural size.
3a. Portion of the calicular surface, magnified.
MIcRABACIA CORONULA (p. 60).

Fig. 4. Calicular surface ; natural size.
4a. Side view, magnified
46. Calice, magnified.
4c. Mural disc, magnified.

CORALS FROM THE LOWER GREENSAND.

HoLocystIs ELEGANS (p. 70).

Fig. 5. A globose mass of this compound corallum ; natural size.
5a. Portion of the calicular surface, magnified.
5 6. Vertical section, magnified, of the visceral chambers in which the septa have

been partly cut away, in order to show the tabular arrangement of the
dissepiments.

TAB. XI.
CORALS FROM THE GAULT.

CyatHina BowErBaNKit (p. 61).

Fig. 1. A weather-worn specimen ; natural size.
la. A specimen showing the wall, but not the calicular margin.
14. Horizontal section made near the calice, and magnified, to show the position
of the pali, &e.

BATHYCYATHUS SOWERBYI (p. 67).
Fig. 2. Side view of a specimen, magnified, so as to show the structure of the wall.
The line placed on the side shows the natural size of the corallum.
2a. Calice magnified ; the wpper half is represented in its natural state, but in the
under half the septa have been cut down; the centre is clogged up with
extraneous matter.

CycLocyatuus Firront (p. 63).
Fig. 38. Side view of the corallum, magnified.
The line placed below shows the natural size of the specimen.
3a. Calicular surface, magnified.
36. Under surface, or mural disc, magnified.

‘TRocuocyatuus Harvreyanus (p. 65).
Fig. 4. Side view of the corallum ; natural size.
4, Calicular surface, magnified.
44. Under surface, magnified.

TROCHOCYATHUS CONULUS (p. 63).

Fig. 5. Side view of the corallum ; natural size.
5a. Calice, magnified.

‘TROCHOSMILIA SULCATA (p. 68).
Fig. 6. Side view of a specimen, the upper part of which is broken on one side ;
natural size.
6a. Calice, magnified.
64. A restored specimen, magnified.

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