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v.

BRITISH FOSSILS.

On the STRUCTURE of the BELEMXITUXE ; with a description of
a more complete Specimen of Belemnites than any hitherto
known, and an account of a New Genus of BELEMNiTnxfi,
Xiphoteuihis, by THOMAS H. HUXLEY, F.R.S., Professor of
Natural History, Royal School of Mines.

THE fossil shell called Belemnites consists fundamentally of (1) a
hollow cone, the phragmoconus, with a thin, shelly, wall which may be
termed the conotheca, and which is divided by transverse septa, more
or less convex towards the apex of the cone, and concave towards its
base, into chambers, or loculi. Each septum is traversed, close to the
conotheca, in a direction which corresponds with the median ventral
line of the body, by a canal, the siphunculus. More or less extensively
enveloping the apical part of the phragrnocone is (2) a more solid body,
the guard, or rostrum, composed of calcareous matter arranged in prisms,
or fibres, perpendicular to the planes of lamellae. These are disposed
concentrically around an axis, the so-called apical line, which extends
from the extremity of the phragmocone to that of the rostrum.

All observers are agreed as to the presence of the parts hitherto
mentioned in a Belemnite, but a great diversity of opinion prevails
respecting the nature, and indeed the existence, of a third constituent
of the fossil, the so-called " pen " or " osselet." As the part which
commonly goes by the name of " pen " in the Belemnite, however,
corresponds to only a part of the structure already known as the
" pen " in recent Cephalopoda, I shall endeavour to avoid ambiguity,
by using for it the appellation of pro-ostracum.

In a paper " On the discoveiy of a new species of Pterodactyle, and
;i nl*o of the faeces of the Ichthyosaurus, and of a black substance re-
" sembling Sepia or Indian ink, in the Lias of Lyme Regis," read before
the Geological Society, on the 6th February 1829, by Dr. Buckland,
the following passage occurs.

" 3. Fossil Sepia. — An indurated black animal substance, like that
in the ink-bag of the cuttle fish, occurs hi the Lias at Lynie Regis ;
and a drawing made with this fossil pigment, three years ago, was
pronounced by an eminent artist to have been tinted with sepia. It
i> nearly of the colour and consistence of jet, and very fragile,
with a bright splintery fracture ; its powder is brown, like that of
painters' sepia; it occurs in single masses, ' nearly of the shape and
size of a small gall-bladder, broadest at the base, and gradually con-
tracted towards the neck : these are always surrounded by a thin
nacreous case, brilliant as the most vivid Lumachella ; the nacre
seems to have formed the lining of a fibrous, thin, shelly substance,
\vhich, together with this nacreous lining, was prolonged into a hollow
cone like that of a belemnite, beyond the neck of the ink-bag ; close

A 2

355817

4 BRITISH FOSSILS.

to the base of the ink-bag there is a series of circular transverse
plates and narrow chambers, resembling the chambered alveolus
within the' cone of a belemnite ; but, beyond the apex of this alveolus,
no spathose body has been found.

" The author infers that the animal from which these fossil ink-bags
are derived, was some unknown cephalopod, nearly allied in its
internal structure to the inhabitant of the belemnite ; the circular
form of the septa showing that they cannot be referred to the
molluscous inhabitant of any nautilus or cornu-ammonis."

This is, so far as I am aware, the first notice of the existence of a
pro-ostracum in a Belemnite.

Voltz, writing in 1830, had no direct knowledge of the existence of
any such structui'e, though he was led by observation of the lines of
growth of the conotheca, to conclude that it was prolonged dorsally ;
but, in the same year, Count Minister* published descriptions and
figures of Belemnites with complete pro-ostraca.

" In the Belemnites there is an empty prolongation, similar to the
last open chamber, of the guard, which is as long as the thick and
chambered shell, but infinitely more delicate, as in the Orthoceratites.
and hardly of the thickness*of the skin of a bladder."

The species which exhibited this pen in the best state of preserva-
tion was Belemnites scmisulcatus, complete specimens of which, from
the Solenhofen slates, are represented in Count Minister's third and
fourth figures.

Dr. Bucklaud returns to the subject in his '' Bridgewater Treatise "
(1836), page 374, note.

" In 1829, I communicated to the Geological Society of London a
notice respecting the probable connection of Belemnites with certain
fossil ink-bags, surrounded by brilliant nacre, found in the Lias at Lyme
Regis. (See Phil. Mag., N.S., 1829, p. 388.) At the same time I
caused to be prepared the drawings of fossils engraved in Plate 44",
which induced me to consider these ink-bags as derived from Cephalo-
pods connected with Belemnites. I then withheld their publication.
in the hope of discovering certain demonstration, in some specimens
that should present these ink-bags in connection with the sheath or
body of a Belemnite, and this demonstration has at length been
furnished by a discovery made by Professor Agassiz (October 1834), in
the cabinet of Miss Philpotts, at Lyme Regis, of two important speci-
mens which appear to be decisive of the question. (See PI. 44',
figs. 7-9.

" Each of these specimens contains an ink-bag within the anterior
portion of the sheath of a perfect Belemnite ; and we are henceforth
enabled with certainty to refer all species of Belemnites to a family in
the class of Cephalopods for which I would, in concurrence with
M. Agassiz, propose the name of Belemnosepia."\ Such ink-bags are

* " Nouvelles Observations sur les Belemnites, in Boue's Memoires Geologiques,"
translated from " Bemerkungen zur niiheren Kenntniss der Belemniten." — 4to. Bai-
reuth, 1850, p. 296.

fBuckland it is obvious, proposes Belemnoscpia as a name for the animal, or ' family '
of animals to which the shell called " Belemnite " belongs, and not as a name for the
particular specimens which he examined. Those who have considered these speci-
mens not to belong to the family of the Belemnitida: at all, therefore, but to the
Teuthidcp or Loligida*, are not justified in using Belemnosepia as a generic title — for
the intention of the authors of that name is to employ Belemnosepia as the equivalent
of Belemnites in its ordinary acceptation; and as naturalists have refused to do so,
and continue, as I think, wisely, to nse "Belemnites" for the genus of animals
which fabricate the shell called " Belemnite," Belemrwsepia remains as a mere syno-
nym, and can be employed in no other sense.

BRITISH FOSSFLS.

occasionally found in contact with traces of isolated alveoli of Belem-
nites ; they are more ^frequently surrounded only by a thin plate of
brilliant nacre."

In his statement of the component parts of a Belemnite (L c., p. 372),
Dr. Bnckland yery clearly demies the characters of the part which he
had discovered ; it is, says he, ** a conical, thin, horny sheath or cup,
•• commencing from the base of the hollow cone of the fibro-calcareoui
" sheath, and enlarging rapidly as it extends outwards to a considerable
" distance. PI. 44', fig. 7 b, e, e', e". This horny cup forms the
** anterior chamber of the Belemnite, and contained the ink-bag (e) and
~ tome other viscera."

In Dr. Buckland's restoration of the " Beleinnosepia," PL 44' fig. 1,
the cup in question, or pro-ostracum, e, et is made to extend nearly to
the anterior end of the mantle of the animal.

In the preceding year (1835) Professor Agassiz communicated a
short note, a Ueber Belemniten," to Leonhard and Bronn's " Jahrbuch."
in which he states that " he has made out with certainty that the so-
called Onychotenthif prisca with the ink-bags, as they are figured by
Van Zieten (as Loligo, Tab. XXV'.), are nothing but the anterior pro-
longation of a Belemnite, and, indeed, of B. oralis, a? is shown by a
perfect specimen from the Lias of Lyme Kegis, in Miss E. Philpotts'
collection. ** The Belemnites have, therefore, the plate of Onychoteu-
" tki&, as a prolongation of the alveolus, and, internally, the ink-sac of
- Sepia."

-hall find reason to believe, however, that what Von Zieten
figures in the plate referred to are pens of Loligidte. And it must be
particularly observed that Agassiz and Buckland. though apparently in
agreement, are not really so. Dr. Bnckland neither assents to the
proposition that the pens figured by Von Zieten belong to Belemnites,
nor does he agree with Professor Agassiz' opinion that Miss Philpotts'
specimens exhibit traces of such a pen.

In the paper entitled " Bemerknngen uber das genus Belemnosepia
** und uber den fossilen Dinten-sack in den vorderen Kegel der
" Belenmiten." in fact, Buckland speaks of Von Zieten's specimens a?
" species of Loligo " (p. 39. note), and in the Bridgewater Treatise,
p. 308, when treating of the pens and ink-bags of " Loligo" in the
English Lias, he says, —

" We learn from a recent German publication (Zieten's Versteine-
rungen Wurtembergs, Stuttgart, 1832, PI. 25 acd PL 37), that similar
remains of pens and ink-bags are of frequent occurrence in the Lias
shales of Aalen and Boll."

Taking for granted the correctness of Professor Agassiz' interpreta-
tion of the pen called " OnycJioteuthis prisca," as the pro-ostracmn of
Belemnites oralis, .M. Voltz contributed much towards the acceptance
of that interpretation by essaying to prove (Ueber Qnychotevthis pritca
von Munster ; Leonhard und Bronns Jahrbuch, 1836, p. 323), that the
arrangement of the lines of growth in the former corresponds with that
of certain striations upon the couotheca of Belemnites to which he had
drawn particular attention in his " Obeervationes sur les Belemnite? "
(1830). These are two systems of very remarkable striae visible on
the outer surface of the lamellated test composing the conotheca :
" the one kind are straight and set out from the apex, they are
" analogous to the longitudinal striae of all univalve and bivalve shell?;
" and the others are more or less transverse." The first set are
usually well seen only in the ventral or internal face. " Commonly,
" the external lamina of the conotheca shows them more distinctly
" than the internal lamina?, whilst the other striae are seen equally

6 BRITISH FOSSILS.

" well on all the laminaj. These last indicate the mode of its pro-
" gressive growth, and, consequently, the form which the opening had
" during the whole period of its growth.

" These striae of growth form a series of transverse semicircles,
parallel with the sutures of the septa, on the ventral face of the phragmo-
cone. There are always many on each alveolar chamber, and they are
the closer together the nearer they are to the apex. This arrangement is
seen only on the ventral face of the phragmocone ; and when the striae
reach the lateral regions they assume an almost hyperbolic curvature
to approach the straight lines which pass from the apex of the cone
and run, between the side and the back, as far as the aperture. I
shall call these lines the asymptotes, and the lateral regions where the
striae have a hyperbolic curvature, hyperbolic area (regions hyper-
boliques) ; the region between the asymptotes I shall term the dorsal
area (region dorsale). The transverse strias sometimes unite in groups
into a single line, when they take the hyperbolic curvature and ascend
towards the asymptote. The width of the dorsal area, that of the hyper-
bolic area?, the quantity and the curvature of the strias of these different
areae, vary according to the species ; but this variation is very slight.
The width of the dorsal area is usually about one-fourth of the circum-
ference, and that of each of the hyperbolic areas an eighth.

" The stria? of the dorsal area are less numerous than those of the
rest of the test, and are usually less pronounced than the latter, being
sometimes imperceptible ; they form ogive arcs, the apex of which is
turned towards the aperture of the shell. Often, one sees a slightly
raised straight line, which sets out from the apex of the cone and in-
tersects the summits of all the ogives ; at other times a groove traverses
the region ; and occasionally the ogives are not visible.

" It would appear from these facts that the ventral edge of the
aperture of the phragmocone is parallel to the sutures of the septa,
and that, on the sides, it curves round almost at a right angle to form
an elongated lobe, which terminates in an ogive arch on the dorsal
side."

The existence of these conothecal strife has been noted by all
observers, and Voltz's clear description of their distribution and
direction has been largely confirmed. I shall have to point out, how-
ever, that one species of Belemnite, at any rate, exhibits a different
pattern.

The view taken by Agassiz and Voltz of the nature of the Liassic
pens, formerly referred to Loligo and Onychoteuthis, met with strong
opposition in an essay by Prof. Quenstedt, entitled " Loligo Bollensis
1st kein Belemiten Organ," and published in Leonhard and Bronn'-s
Jahrbuch for 1839.

Prof. Quenstedt points out, with great justice, firstly, that the mark-
ings on the pens are quite different in character from those on the
phragmocones of any known Belemnites ; and, secondly, that the
posterior ends of the pens are complete, and certainly were not united
with any such structure as a Belemnite, while it is impossible to
imagine that the latter should have been attached to the anterior ends
of the pens.

In 1 840, however, M. Voltz, in his " Observations sur les Belopeltis
ou lames dorsales des Belemnites,"* brings forward new arguments in
favour of Agassiz' opinion.

Memoires de la Societe d'Histoire Naturelle de Strasbourg — June 1840.

BRITISH FOSSILS. 7

In the first place he ,gives a clear view of the structure of Beleni-
nites in general, repeating and extending his previous statements, and
more especially denning his opinion as to the meaning of the conothecal
stria?.

"As the stria? of growth represent the successive openings of shells,
one is always enabled to form an exact idea of the form of a shell at
any stage of its growth, when the whole length of these striae is fol-
lowed out. so that an exact idea of the form of the opening of the cono-
theca (test alveolaire) of a Belemnite may be formed by following one
of the lines of growth.'*

Volte gives a restoration of the Belemnite shell constructed upon
this principle in the third figure of the third plate of the work cited.

The Liassic and Oolitic pens are, for M. Voltz, the pro-ostraca of
Be lemnites ; but as these Belemnites are unknown, he proposes for
them the new generic name of Belope his. In anticipation of obvious
objections, M. Voltz writes : —

" It might be supposed that the Belopeltes belong to some other
genus of Cephalopods than Belemnites, or than any other known form
of Acetabulifera ; and the shell of which, though without a guard, had
much analogy with the alveolar test [conotheca] of the Belemuite.
But then one would ask why these Belopeltes are always incomplete
at their apices, a fact which is fully explained, and, so to speak, be-
comes a necessity, when these fossils are referred to Belemnites. It
would also be necessary to explain why no fossils we ever found which
can be referred to the apex of Belopeltis, and why, lastly, fossils are
never met with appertaining to the dorsal lobe of the alveolar test of
Belemnites : a very much elongated lobe, the existence of which in
entire and uninjured Belemnites cannot be doubted by any one who
has cai-efully examined the strife of growth of the alveolar test [cono-
theca] of the Belemnites." L.c., p. 21.

And further, in his " Observations Supplernentaires " (l.c. p. 31) : —

•• M. Buckland gives, on his Plates 44' and 44", figures representing
fragments of Belemnites found &t Lyme Regis and still containing the
ink-bag. The figure 7, Plate 44', represents Belemnites ovalis with its
ink-bag.

" It is to be regretted that M. Buckland has not published a figure of
the counterpart of this specimen ; for M. Agassiz, who studied this very
important fossil in 1834 or 1835, and who discussed it at length with
M. Buckland, says, in the German translation of the Bridgewater
Treatise, that the counterpart of the fossil exhibits the dorsal region
of the alveolus with striae similar to those which are seen in Plates 28,
29, and 30 of this work,* so that not the smallest doubt could be enter-
tained as to the justice of the union of Belopeltis with Belemnites, a
union the necessity of which was made obvious to my friend, M.
Agassiz, by his first inspection of the fossil.

" M. Buckland states in his work that the specimen presents a
nacreous test showing transverse and waved stria?. M. Agassiz says,
on the other hand, that he does not comprehend this explanation, and
that these striae are the traces of the sutures of the septa with the
alveolar cone. The mere inspection of the figure suggests the same
idea, for these stria? appear to be more marked than simple striae
of growth would be, and they are placed at relative distances which
correspond well to the intervals between the alveoli of Belemnites, and
which are too great to be striae of growth. If this be the case, it

Pen and ink-bags of Loligo " are represented on these plate*.

8 BRITISH FOSSILS.

evidently follows that the ink-sac is not in its natural place, since it
occupies that of the septa, which would be impossible.*

" M. Quenstedt has just published, in Leonhard and Bronn's Jahr-
buch, a memoir, the intention of which is to prove that the Belopeltes
do not belong to Belemnites. He gives, in this memoir, the figure of a
Belopeltis, which does not appear to me to be exact, because it repre-
sents the fracture which is always observed at the posterior part of
these fossils, not as a fracture, but as the commencement of the shell ;
this would then be the point from which the successive growths Avhich
formed the shell started. M. Quenstedt had the goodness, in August
1839, to show me, in the museum of the University of Tiibiugen, the
original of this figure, but I was unable to agree with him on this
point. The striae of growth of this fossil present no point that can be
considered as the origin of the shell ; the asymptotes and the hyperbolic
striae can be very well observed cutting transversely the lines which
this naturalist takes for the origin of the shell ; therefore the part of
the shell whence its growth emanated, was inevitably situated beyond
that line. The origin must have been at the point of union of the two
asymptotes, and the shell is necessarily incomplete at its extremity.
Now the whole theory of M. Quenstedt is based on this obviously
erroneous mode of interpreting the fossil, so that it cannot be ad-
mitted."

In 1842, M. D'Orbigny, Paleontologie Francaise, Tome ler, p. 41,
and Planches II., III., IV., described with much confidence, and
figured, what he terms the " osselet corne " of the Belemnites.

"It varies but little in form, as I have been enabled to judge by the
examination of more than 15 distinct species, the rostra of which
are very different, while I have always found it to have the same
configuration. It is composed, in front, of a broad spatuliform plate,
exhibiting, in the middle, a wide dorsal regionf (a, Fig. 1, PI. IV., et 3,
PI. III., the part comprised between the lines b, 4), the angle of which
always exceeds ten degrees, covered with ogive-like striae of growth,
Avhich unite on each side at the metlian line, which is sometimes
projecting or slightly grooved.

" On each side of the dorsal region are the lateral expansions,^ Avhich
pass from this region and form, on each side, delicate horny plates,
which are marked with lines of growth passing obliquely from above
downwards, and from the dorsal to the ventral face. These expansions
accompany the " osselet " through its whole length (see PI. III., fig. 2,
PI. IV., fig. 1, the parts which lie between the lines c, d), and diminish
in width from above downwards as far as the inferior part, where they
form a longer or shorter conical cup, which appears to constitute about
a third of the whole length. On the sides, at the point of junction of
the lateral expansions with the terminal cup, the lines of growth
suddenly become sinuated, form curves with a dowinvard convexity,
and become transverse in the whole ventral region, to give rise to the
terminal cup, a kind of reversed horny cone, in which the chambers are
developed successively as the animal grows."

It appears, however, (see p. 43 of the work cited) that all this elabo-
rate description of the " osselet corne " is not based upon the exami-

* It would be interesting to know -what the striae seen upon the counterpart by
Professor Agassiz really were, and why Dr. Buckland nowhere says anything about
them. I suspect that they were the conothecal striae.

f The Asymptotes of M. Voltz's Memoir, p. 3.

J It is these lines, convex when the cone is reversed, which form what M. Yoltz
calls " hyperbolic regions."

BRITISH FOSSILS. 9

nation of a specimen of any such structure, but that it is deduced from
the character of the markings upon the surface of the phragmocone
originally described by Volte. In point of fact I/Orbigny made no
real addition to the discoveries and conclusions of the latter excellent
observer.

Quenstedt (Die Cephalopoden. p. 389) agrees with Yoltz in the descrip-
tion of the lineations of the sheath of the phragmocone, bat declines to
deduce thence the existence of a pen. Making no distinction between
the views of Bnckland and those of Agassiz, and continuing to deny
the existence of evidence justifying the connexion of such pens as those
of Loliffo Bollenfit. with Bclemmtrj. he falls into the error of doubt-
ing Buckland"s identification of the nacreous pro-ostraca^ &c. of Lyme
Regis with Belemnites ; and, after a critical examination of Btlem-
nilft semismlcatu*, he is disposed to admit, at most, ~ that the shell
~ of the Belemnite alveolus (= phragmocone) did not end superiorly
** by a circular lip, but in a unilateral parabolic process, which can by
'* no means be safely compared to a true Loligo pen."

M. Duval Jouve (" Belemnites des Terrains Cretaces InfenemW
1841), observed no pro-osrracum attached to any of the specimens he
studied.

In describing the specimens upon which he founded the genus
Bclemmolfutkis, in the " Proceedings of the Geological Society "for
1842, Mr. Channing Pearce indicated the existence of a " sepiostairc "
in that genus, in addition to the phragmocone and guard.

Professor Owen, in his memoir " On the Belemnites " (Phil. Trans.
1844), having mistaken BelemnoteiUkis for Btleimnitet. describes the
rudimentary rostrum of the former as the conotheca of the hitter : with
regard to the existence of a pro-ostracmn in Belemnites generally, he
follows Dr. Bnckland (Lc., p. 66).

In 1848 Dr. Mantell (Observations on Beleniniie-. &c~ Philosophical
Transactions) gave a more complete account of the pro-ostracum of a
Belemnite ( Belemmiltx 'aifeittiatits) than had previously appeared. In
describing the specimen figured in his Plate XV., fig. 3, he writes, —

** This fossil comprises the following parts : — 1. The capsule or
pfriotfraeum. This external investment (c1. e1. e1), which consists of
a thin, shelly, or corneo-calcareons integument that closely embraces
the guard, and, gradually enlarging upwards, finally surrounds the
peristome of the phragmocone. constituting; the thin horny laminated
sheath or receptacle (<% r), has been described by all previous'
observers as an extension of what they termed the sheath, or capsule ;
within this receptacle the ink-bag and other viscera were probably
contained * * ~ * » * *

The phragmocoue enlarges upwards, and anteriorly to the siphonated
part constitutes a large chamber, from the margin of which are pro-
duced two or more long, upright, shelly or calcareous processes, as
shown in PL XV., Fig. 3 ft, ft1."

In a subsequent memoir* Dr. Mantell shows that there were but two
of these processes, that they were situated nearer the dorsal than the
ventral aspect of the phragmocone, and that they were continued
downwards into * nacreous bands or plates, finely striated ** upon the
outer surface of the chambered cone.

A- I have already indicated, Quensiedt (Die Cephalopoden, 1849)
discusses the question of the presence or absence of a pen in the
Belemnites at great length, without arriving at any decided result ;

* On the •traettnv of the Briemnhe sad Brteanratevtfcis.— Pbikwophkal Tbansae-

10 BRITISH FOSSILS.

and Bronn (Letho>a, dritte Aufla<re, Bd. II., 1851-2) increased the
confusion which had already spread over the subject, by following
Qnenstedt in his scepticism respecting the truly Belemnitoid character
of Belemnoteuthis, and by referring the Belemnosepia of Buckland to
the family of the Loligidce and the genus Belopeltis. At least he
does this at p. 407 of the work cited, while at p. 385 lie seems to
entertain a, different opinion :

" The animal of the Belemnite is perfectly unknown ; what Buckland,
Agassiz, and R. Owen say of it must be referred to Belcmnoteuthis,
which only on account of accidental juxtaposition has been taken for
a part of a Belemnite. Of an ink-bag also no trace is ever seen."

That the specimens described by Professor Owen as the animals of
the Belemnite are Belemnoteuthes, as Pearce, Cunnington. and Mantel!
demonstrated, admits of no doubt, and has since been acknowledged
by Mr. Owen ("Palaeontology," p. 113), and that Agassiz was in error
in identifying Von Zieten's Onychoteuthis prisca with a Belem-
nite, also seems to be clear ; but Buckland is in a totally different
position, and it will be seen that his interpretation of the Lyme Regis
specimens was in the main correct.

Of the pro-ostracum (hornige Dute) Bronn says that " it has never
" been found entire, and rarely in a substantive form, but its compo-
" sition must be judged by combining fragments and their lines of
" growth." In fact he essentially adopts Voltz's view of its structure
and extent.

Professor Quenstedt's final opinions appear to be stated in his

" Handbuch der Petrefakten-kunde " (1852). After describing the cono-

thecal lines, he says (p. 385), " It is but rarely that all these markings

" are distinctly visible, and especially they do not agree with those on the

" pen (schulpe) of the parabolic Loliginites, as was for some time wrongly

" asserted, and as some still maintain. On the other hand, they indicate

" the end of the couotheca (alveolarschale), as k has long been known

" from Solenhofen, and has been lately figured by Mantell, also from

" the Oxford clay at Trowbridge in Wiltshire (Philosophical Trans-

" actions, 1848). From two corresponding specimens, which I have

" lately obtained from Soleuhofen, the fig. 13 of plate 31 is constructed of

half the natural size. The conotheca (A) is chambered up to its upper

part, but when the chambers cease, the lip also ends upon the ventral

side, as it seems by a horizontal boundary, which would answer to

the horizontal lines, b, upon the conotheca of B. giganteus. Dorsally,

on the contrary, a high parabolic shield extends, at the edges of which

" two sometimes deeply coloured bands, h,h, are clearly perceived, and

" end in points like sharp ears superiorly. These are the hyperbolar

" regions, which, where they bend down below from the margin, have

" quite the same curvature as in Belemnites giganteus. Between these

" horns lies the region of the dorsal curves, a,a, with a median line

" r, in which the lines of growth are plainly curved upwards, just as

" the free margin of the shield is."

Professor Quenstedt's observations thus clearly confirm those of
Mantell, and go to prove that some Belemuites have a two-ribbed pro-os-
tracum ; but he is, as we shall see, in error in supposing that all
Belemnites have a pro-ostracum like that of his specimens ; and still
more in his assumption (p. 333) that Belemnoteuthis has no phrag-
mocoue. and is not one of the BelemnitidcB, but an Onychoteutln*.

Professor Pictet (" Traite de Paleontologie, "1854), follows d'Orbigny
(or rather Voltz,) as to the pro-ostracum of Belemnites, and with Bronn
separates Belemnosepia from the Bclcmnitidfr, Avhile. on the other

BRITISH FOSSILS. 11

hand, he is inclined to admit Belemnoteuthis as a distinct genus of the
last-named family.

Mr. S. P. Woodward (Manual of the Mollusca, 1851) states that
the " Pen " of the Belemnites is " represented by the nacreous bands on
" the dorsal side of the phragmocone, and produced beyond its rim in
" the form of sword-shaped processes (PI. II., fig. 5)." From this
description and the reference to the figure of Belemnites Puzosianus
from the Oxford clay ( = B. attcnuatus, Mantell), it is clear that Mr.
Woodward conceives that the pro-ostraca of Belemnites, in general,
are constructed upon the type of that of B. Puzosianus.

Mr. Woodward identifies Belemnosepia with Geoteuthis, and therefore
refers Buckland's specimens to the Teuthidte and not to the Belemnitidte.
In his Supplement, published in 1 856, the same author writes (p. 449), —

" Belcmnites.-*-Professor Buckman of Cirencester possesses a phrag-
mocone from the Lias, containing the fossil ink-bag."

Professor Owen (Palaeontology, 2nd ed., 1861, pp. Ill, 112), in speak-
ing of the remains of " Calamaries (Teuthidae)," states that "the most
" common form of the gladius has the shaft wide and longer than the
" wings ; it has a nacreous lining, and is usually accompanied by a large
" and well preserved ink-bag (Geoteuthis, fig. 34, 4). These were called
" Belemnosepia by Agassiz and Buckland, who supposed them to belong
" to the same animal with the Belemnite."

But I am not aware that any one has yet observed a calamary's
" gladius,' with a nacreous lining ; and we have seen that Buckland
entertained no such opinion as that here ascribed to him.

Belemnoteuthis is identified with Miinsters Acanthoteuthis . Of the
Belemnites, Professor Owen, adopting Dr. Mantell's results, says, " The
" last chamber is rarely preserved, and appears to have thinned off" into
ct a mere horny sheath with, sometimes, two pearly bands like knife-
" blades on the dorsal side. It must have been sufficiently capacious
" to contain all the viscera. The ink-bag has been very rarely found,
" and is even smaller than in the last genus, as if in relation to the
" more greatly developed shell."

From the preceding survey of the literature of the subject it appeal's
that very diverse opinions obtain respecting the nature and character
of the pro-ostracum in Bclemnitidte.

1. According to Dr. Buckland this part is a corneous, or shelly,
and more or less completely nacreous, extension forwards of the lip of
the phragmocone.

2. According to Agassiz it is a " pen " identical with that of the
so-called Loligo Bollensis, &c.

3. According to Voltz, it is a pen analogous to that of Loligo
Bollensis.

4. According to Mantell and Quenstedt, it is a broad dorsal plate,
more or less corneous in the middle, and with two strong calcified
asymptotic bands.

Furthermore, as to the existence of specimens proving that the
Belemnite was provided with an ink-bag ; some, like Buckland and
Woodward, affirm the fact as a matter of direct observation ; others
support the conclusion that the ink-bag existed in Belemnites, by the
analogy of Belemnoteuthis ; while yet others, denying Belemnoteuthis,
or Buckland's Liable specimens, to belong to the Belcmnitidte, doubt the
existence of any direct, or conclusive indirect, evidence of the existence
of this organ in the Belcninittda.

1.2 WIIT1SH FOSSILS.

So with regard to the acetabular hooks and beaks : that such slruc-
tures appertained to Belemnoteuthis was proved by Pearce and Cun-
nington and by Professor Owen ; but, at present, I am aware of no
direct evidence of their existence in Belemnites.

These lacunas in our knowledge of one of the most important of extinct
forms of life are filled up by the specimens described in the present
Memoir, which demonstrate the accuracy of Dr. Buckland's view of the
nature of the Lyme Regis fossils figured in his Plates 44' and 44"; and
furthermore, prove that at least three types of structure of the pro-
ostracum must be distinguished among the Bclemnitidce,

Some time ago, my friend Mr. Day of Charmouth was good enough
to direct my attention to a number of remarkable specimens of Belem-
nitida; obtained from the Lias in his neighbourhood, and noAv either in
his own possession, or in that of the Rev. Mr. Montefiore and Mr.
Henry Norri's, gentlemen to whom I am greatly indebted for the
readiness with which they have entrusted valuable and important
fossils to me for examination and description.

The most complete Belenmite in existence, to my knowledge, is
that .specimen, the property of the Rev. Mr. Montefiore, which is
represented in Plate I., fig. 1. Mr. Day informs me that it was
obtained from the Ammonites obtusus zone of the Lower Lias near
Charmouth.

In this remarkable fossil not only are the guard, phragmocoue, and
pro-ostracum preserved, but the general contour of the body is shown,
the beak is in its place, and irregular lines of hooks indicate the
position and extent of the arms.

The length of the whole animal, from the summit of the beak to the
apex of the guard, is 12^ in., Avhile its greatest breadth does not exceed
1^ inches. The arms, as indicated by the lines of hooks, cannot be
traced for a distance of more than 1^ in. from the beak, and they
diverge from one another, so as to include a triangular space, the
broad base of Avhich is superior, while its apex is close to the beaks.

The internal shell is 11 '8 inches long, and consists of guard, phrag-
mocoue and pro-ostracum. The guard and the phragmocoue occupy
rather less than the half of the whole shell (5*5 in. about). The guard,
about 0*3 in. thick in the middle, and nearly circular in section, remains
of about the same diameter for l^iu., widening, above, into the alveolus
for the phragmocone, while, below, it tapers to the apex. The inferior
narrowing commences at about half an inch from the apex, which is
marked by five indistinct, short, longitudinal grooves. Superiorly,
the guard spreads out over the phragmocoue, becoming gradually
thinner, and ceasing to be traceable, as guard, at the point (d'j, about
2f in. from the apex.

The guard is broken at the points b smd c ; at b the surfaces corre-
spond perfectly, but at c there is a slight loss of substance, so that the
portion of the guard, a, c, may not be in quite its natural position
relatively to the rest. I am inclined to suspect that there has been some
slight shifting of position at this point, from the circumstance that the
curved contour of the right side of the guard is somewhat more convex
than that of the left side, while the whole guard is slightly flattened
in the plane of the surface of the matrix. The right contour should
therefore be ventral, and the left dorsal. But there can be hardly any
doubt that the upper face of the body of the specimen is the dorsal
aspect, so that it Avould appear that the guard, where broken at c, has
undergone a certain twist upon its axis.

The primary chamber of the phragmocone probably lies in the guard
about the point indicated by the letter x. For the two lateral margins

BRITISH FOSSILS. 13

of the phragmocone, if produced, would meet thereabouts ; and further-
more, while at the line of fracture, r. the diameter of the phragmoeone
i* nearly equal to that of the guard, the latter forming a layer cot
more than ^ in. thick round it, at 6, the guard b solid throughout.
Where the guard ceases to be traceable on the phragmoeone at </, it Ls
O-To in. broad, and the phragmoeone gradually widens, until at e, the
furthest point to which it can be traced, it is H in* broad. It must
be recollected, however, that these dimensions are exaggerated by the
crushing and flattening of the specimen.

From e \of, a distance of 2\ in, the pro-t*?tracum is represented by a.
delicate lamella, of shelly substance, for the most part exhibiting a
beautiful nacreous lustre, and as wide as the upper diameter of the phrag-
moeone. The central part of this lamella is blackish, with metallic
reflections, and its edges are constricted in the middle, so that it appear?
saddle-shaped. At the sides it passes into a lamina of yellow nacre (A),
which dips down towards the ventral side of the body* and Is traceable,
on the right side, as far as the point (A'> Beyond this, no remains of
any shelly matter are distinctly visible, but the surface of the matrix
exhibits an irregular impression., extending as far as (fj, as of a, thin,
broad, partially crushed, oval extension of the pro-ostraeum. I presume-
that the mantle of the animal also terminated at this point. Beyond it.
the impression of the head is indistinctly traceable ; and it is worthy
of note that the head seems to have been small as compared with the
size of the body. The oral circle, embraced by the bases of the short,
uncinated, arms (/ >. is particularly narrow, so that these bases are closely
approximated. What was the precise number of the arms, and whether
any long tentacles did or did not exist, cannot be ascertained.

The remains of the beak (*), about half an inch long by O4 in. wide,
are so crushed and broken that there is some difficulty in the way of
interpreting the appearances it presents. I believe, however, that the
two beaks are fractured transversely, the dorsum of the dorsal beak,
and the edges of the ventral beak, having been left in the absent matrix;
and I take k to be the fractured edge of the dorsal beak surrounded by
k\ that of the ventral beak.

The substance of the beak is black and carbonized, and exhibits no
evidence of any calcareous coat. The irregularly dispersed hooks do
not seem to have remained in place upon the bases of more than two
of the arms. There are indications that they were disposed in double
rows of opposite hooks along each arm.

The most perfect of these hooks (PL L, fig. la) measures about one sixth
of an inch in a straight line from its base to it* apex. The basal part
seems to be nearly square, and is hollow ; from the base the hook is
continued at first in nearly a straight line, and then bends sharply
round to its acute point. The cavity of the base is traceable through
the hook, and probably terminates by an aperture at, or close to, its

The ink-bag is not very clearly distinguishable (a dark spot at m only
place) in this specimen ; the great value of wT

specimen ; the great value of which con-
demonstration which it affords of the co-existence of horny
hooks and beak, a nacreous pro-ostracum, and the ordinary guard and
phragmoeone of a Belemnite ; and, incidentally, of the justice of Dr.
Bnckland's identification of the Lvme Regis* « B<lrim*otrpia> " with
Befcaudfe*.

So much difficulty attends the identification of the species of the
Belemnitft, that I hesitate to attach any specific name to this specimen.
In many respects it is closely allied to the Btlemmitet eloityatm* of
Sowerby ; but the Self mm/ft Brmottifriamtu of ITOrbigny is abun-

14 BRITISH FOSSILS.

dant in the bed in which it was found, and my colleague, Mr. Etheridge,
is of opinion that it belongs to that species, "though it has strong
affinities with B, Foumeliamts (D'Orb)."

In the collections of Liassic fossils to which I have referred, and
chiefly in that of Mr. Day, there is a series of fragmentary Belemnitic
remains, consisting for the most part of ink-bags, associated sometimes
with more or less of the pro«ostracum, sometimes with hook? and imper-
fect beaks in very nearly natural relative positions ; sometimes with
more or less of the phragmocone, but hardly ever with a guard. That
these belong either to the species already described, or to a closely
allied one, is highly probable ; in any case, the study of the features
presented by some of them may help to throw light on the structure of
the Belemnitidce generally.

1. In a great many Belemnites I have observed conothecal stria?
having the arrangement described by Voltz (Plate I., figs. 6 and 7) ;
but a large phragmocone from the Ammonites obtusus zone in the Rev.
Mr. Montefiore's collection, the apex of which is broken off, but which
still has a width of 2^ in. and a length of five inches, exhibits a dis-
position of the conothecal lines different from any which 1 have met
with, or seen described. (Plate I., fig. 4a.)

Only a small portion of the conotheca is preserved in this specimen,
coating the cast of a phragmocone in calcareous spar, which exhibits
the remains of the siphuncle very distinctly along the middle line of
that face of the specimen which is turned to the right in the figure.
This is, therefore, the ventral line, and the face turned to the eye is the
left lateral face of the phragmocone, the figures not having been
reversed. Now it will be observed that instead of one asymptote as
usual, there are two, separated by an interval equal to about £th of the
circumference of the phragmocone. Sharply arched hyperbolic lines,
the ends of which pass into the asymptotes, and which are convex
upwards, occupy the space between the two asymptotes ; of which the
one may be termed the dorso-lateral, the other the ventro-lateritl
asymptote.

Faint curved lines run obliquely upwards from the dorso-lateral
asymptote towards the middle line of the dorsal region, so that the
dorsal area of the conotheca doubtless had its usual set of upwardly
convex curved lines. The ventral area, on the other hand, enclosed
between the ventro-lateral asymptotes, exhibits no very distinct
markings, though faint indications of transverse lines are discernible.

The conotheca in this case, therefore, differs from the ordinary type
in having three sets, one medio-dorsal and two lateral, of upwardly
convex curved striae, and in possessing four asymptotes instead of
two.

According to D'Orbigny (" Paleontologie Fran£aise," Terrains Juras-
siques, Atlas, PI. 16, fig. 1), the conothecal lines of Belemnites
Puzosianus have the ordinary arrangement, and the lateral bands of
the pro-ostracuni of this species would seem to correspond with the
asymptotes.

If the arrangement of the conothecal lines, then, indicates the form of
the pro-ostracum and vice versa, the majority of Belemuites ought to
have a two-banded pro-ostracum like that of B. Puzosianus ; and, on
the other hand, the peculiar arrangement of the conothecal lines of the
present phragmocone ought to indicate that it was associated with a
different kind of pro-ostracum ; and, so far, ihere may be ground for

BRITISH FOSSILS. \'j

suspecting that it belonged to some of the species which have pro-
ostraca like that of Belemnites Bruguierianus.

But I am by no means satisfied of the justice of Voltz's assumption,
which D'Orbigny and others adopt, that the conothecal lines must
indicate the form of the pro-ostracum, since the hitter may readily
have been modified by the deposition of shelly matter upon its exterior,
after its first formation.

2. The guard of the typical specimen of Belemnites elonyatus, now
in the British Museum, is covered by a superficial, smooth, thin,
whitish, cuticular pellicle ; and a better developed cuticle of the same
kind has been brought under my notice by Mr. Day in specimens from
the Upper Lias. A small example of the guard of apparently the
same species (PL I., figs. 3, 3a, 3b), pointed out to me by Mr.* Day,
exhibits a much more developed cuticle. This is thrown into fine
longitudinal wrinkles in its upper part, but, inferiorly, the wrinkles
pass into minute ridges and tubercles. Both these and the wrinkles
are larger, and extend farther up, on the dorsal than on the ventral,
aspect of die guard.

I? the existence of this cuticular pellicle an indication of the com-
pletion of the growth of the Belemnite '!

3. A splendid specimen in the collection of Mr. Norris (Plate II.,
figs. 1 and la) shows very clearly the association of a phragmocone with
a nacreous pro-ostracum and a large ink-bag. On the one face (fig. 1)
this fossil exhibits the dorsal part of the pro-ostracum and its con-
tinuation into the guard, while, on the other face (fig. la) the huge
ink-bag is displayed. The saddle-shaped, highly iridescent, region of
the dorsal part of the pro-ostracum (a) terminates in well defined
margins, both laterally and in front, the portion of the pro-ostracum
with which it was continuous, at the sides, having broken away from
this central region. On the left side, however, the lamellar continua-
tion of the pro-ostracum towards the ventral surface (b) is well shown ;
and, like the dorsal portion, it is highly iridescent. When subjected
to an oblique light, the pro-ostracum exhibits a shallow medio-dorsal
longitudinal groove and indistinct lines of growth, which are convex
upwards. The surface which continues the direction of the iridescent
part of the pro-ostracum upwards (c) has a granular pitted surface ;
but I am doubtful whether this appearance is due to the structure of
the pen in this region, or to the manner in which fossilization has
taken place.

The ink-bag is flask-like, 8 inches long, and If inches wide at
widest

4. In Mr. Day's collection, there is a specimen (No. 9) from the
Ammonites obtusus zone, consisting of the upper part of the phrag-
mocone, with almost the whole of the pro-ostracum, and the remains of
many hooks in place.

A length of about 2| inches of the phragmocone is preserved ; its
upper end is 2-^ inches wide, its lower end somewhat more than 1 inch,
but both ends are greatly crushed. Nacreous shelly substance coats
the exterior of the upper part of the phragmocone, and extends upwards
over more than the lower half of the pro-ostracum, which has an oval
form, and is nearly 10 inches long by 3£ inches wide.

The upper four or five inches of the middle portion of the pro-ostra-
1 cum is formed of a thin plate of shelly matter, which is not iridescent,
and beneath which there is no iridescent nacre. In the lower part of
the pen the external non-iridescent substance has a subjacent, beautifully
iridescent layer. In this, a> in other cases, the nacre is bounded by "a
well-defined upper contoiT. which in this instance is con .

16 BRITISH FOSSILS.

The hooks of one arm have remained in position, and are arranged in
two rows, and opposite to one another. One hook is so imbedded in
the matrix as to expose its outer or convex side. In this, as in the
lateral position, the base is much .wider than the shaft of the hook.

The guard is not preserved in any of the preceding fragmentary
specimens, while the ink-bag is but indistinctly traceable in the entire
one first described. But any hypercritical doubt that might remain
as to the possession of an ink-bag by a true Belemnite, must be removed
by Mr. Day's specimen of Belemnites elongatus represented of one-half
the natural size in Plate I., fig. 2, which exhibits the guard and phrag-
mocone complete, with a large and full ink-bag in situ. The ink-bag
is pear-shaped, and tapers off to its duct. The length from the ex-
tremity of this to the base of the bag is 1 • 4 inch, the widest part of the
bag measuring 0*55 of an inch. The shell from the apex to the mouth
of the phragmocone is 5 • 35 inches long. The guard from its apex to
the point at which it begins to expand over the phragmocone measures
about 2\ inches, and is 0 • 25 of an inch wide at widest.

These measurements may enable one to form a rough estimate of the
size of guard which appertained to any detached ink-bag, and rice versa.

I have not been able to make out more than six or seven arms in
any specimen, nor has any exhibited traces of elongated tentacula,
though the shortness of the arms which have been preserved would
lead one to suspect their existence. The hooks in the middle of the
length of each arm seem to have been largest ; those at the ends of the
series, especially at the base, smallest.

The foregoing descriptions demonstrate that certain true Belemnites
were provided with hooks upon their arms ; horny beaks ; large ink-
bags ; and with a pro-ostracum (into which iridescent, nacreous, shelly
matter entered more or less largely) prolonged as a broad spatula te plate
along the whole length of the dorsal region of the mantle, and pro-
duced laterally and interiorly, for an unknown distance, along the lateral
and ventral regions of the body.

But it by no means follows that all Bclemnitida were provided with
a pro-ostracum of similar form and character. On the contrary, it
appears to me to be certain that there were at least two other kinds of
pro-ostracum in this family.

Thus the Oxford Clay Belemnite, described by Mantell (Phil.
Trans., 1848), under the name of attemiatus, a name which appears,
like B. Owenii, to be only a synonym of B. Puzosianus (D'Orbigny)
has a pro-ostracum which was very thin and apparently horny, or
imperfectly calcified, in the dorsal region, and was supported laterally
by two thin calcareous bands, or pillars, which, inferiorly, expand upon
the conotheca.

A third very distinct type of pro-ostracum is exhibited by that
i-emarkable Belemnitoid originally figured and described under the name
of Orthocera elongata, by Sir Henry De la Beche,* who says in a note
(1. c., p. 28), " I have ventured to class this specimen as an Orthocera, as*
" it possesses more of the character of that genus than of the Belemnite,

* On the Lias of the coast in the vicinity of Lyme Regis, Dorset. — Transactions
of the Geological Society, ser. 2nd, voL ii. (1829), PI. IV., fig. 4.

BRITISH FOSSILS. 17

" the external shell not exhibiting the radiating fracture of the latter,
" and I have given it a specific name from its great length in proportion
" to the diameter. Mr. White, to whom I am indebted for the specimen,
" informs me that it was originally considerably longer than at present."

As this specimen (now in the Museum of the Geological Society) is
by no means well represented in the plate accompanying De la Beche's
memoir, I have had a correct sketch of it made (Plate IH., fig. 3).

It consists of an imperfect sub-cylindrical guard 3*2 inches long;
fractured above and below, and having, in its lower part, a diameter
of rather less than one-fifth of an inch. It contains the remains of a long
tapering phragmacone, the chambers of which have been completely
filled with transparent calcareous spar. The rounded, bead-like apical
chamber of the phragmacone, not one-fortieth of an inch in diameter, is
situated at about 0*2 of an inch from the fractured extremity of the
guard. The chambers gradually increase in length and in breadth, until
at 2 • 25 inches from the apex they are 0 • 2 of an inch long by 0 • 25 wide.
Beyond this point, the phragmocone is broken away, but the impressions
of three chambers are left on the inner surface of the eonotheca, which
adheres to what remains of the attenuated, upward, prolongation of the
guard. Altogether, there seem to have been about 30 chambers in
the 2*9 inches length of phragmocone, but the 10 outermost chambers
take up 1 * 8 inches of this extent. The conotheca is a thin lamella of
a much paler colour than the guard, to the walls of the alveolus of
which it adheres.

The real nature of this " Orthoceras " was first revealed by the
beautiful specimen obtained by Mr. Day, which is represented in
Plate HE., fig. 1, reduced to four-fifths of the natural size. Here, the
apex of a phragmocone of similar character is inclosed within a
subcylindrical guard, obtusely truncated at its free end. For an inch
and a half from its distal end, this guard is entire, but beyond this
point (6) it is split, and the dorsal has come away from the ventral
half, leaving the phragmocone (c) exposed. The chambers of the
phragmocone are filled with transparent spar, and their casts, thus pro-
duced, are exposed to view inferiorly. Superiorly, they are coated
over by a thin pellicle of quite a similar character to the theca of
the phragmocone in the original specimen ; and, indeed, at the sides,
this layer dips down between the fractured edge of the guard and the
phragmocone, showing clearly that it is the conotheca.

At 3-6 in. from the end, the specimen is transversely fractured, and
the section of the guard cannot be traced further than the fracture; but
a layer of shelly matter (a) quite similar to that which forms the cono-
theca, and which was obviously continuous with it, coats what appeal's
to be the upper termination of the phragmocone, and passes into the
remarkable pro-ostracum, the extreme point of which is broken off :
when it was entire it measured about 11^ inches. Posteriorly it is
a flat band 0.35 in. wide, which slowly narrows until its width is
about 0 • 2 in. ; it then widens to 0 • 5 in., and, finally, gradually tapers to
its point. Where it widens it thickens, and its surface, from being flat,
becomes convex from side to side, so that its section acquires the
form of a not very depressed ellipse, and this form is retained close to
the apex at/.

The surface of this singular pro-ostracum is polished, but is covered
with transverse wrinkles, or ridges, which are especially numerous
where the flat portion passes into the rounded part.

The sections of the pro-ostracum exhibit its structure. Like the guard
of an ordinary Beleinnite, it is composed of concentric lamella?, each
of which consists of fibres disposed perpendicularly to the plane of the

18 BRITISH FOSSILS.

lamella, whence the cut surface presents concentric and radiating
structure lines.

What the proper structure of the guard may be is more doubtful.
In all •specimens I have examined the texture of the guard is dense
and thoroughly calcified, and any indications of structure are of a
crystalline and not an organic character. The guard of the specimen
longitudinally and vertically bisected, of which a diagram, twice the
natural size, is given in Plate III., fig. 3, presents a dark longitudinal
axial line, a dark terminal transverse line, and another less dark
transverse line rather above the middle of its length.

The terminal transverse dark line is visible on the exterior of the
specimen, and looks at first like a colour band, a sort of indication
of the natural termination of the guard. But, on closer exami-
nation, these transverse markings are seen to arise merely from the
presence of plates of calcareous spar ; in other words, the calcareous
infiltration is transparent in these parts of the fossil. The dark axial
line appears to me to result from similar conditions.

The internal shell just described has not yet been found associated
with ink-bag, hooks, or beaks. The peculiar form of the pro-ostracum,
the long narrow phragrnocone, and the cylindroidal guard, distinguish it
generically from all the other Belemnitidce. I, therefore, propose for
this new generic type the name of Xiphoteuthis, and retain for the
present, the only known species of the genus, De la Beche's term of
elongata.

Is the guard entire in these specimens of Xiphoteuthis elongata, or
has its apex been broken off ? Was it originally solid and composed of
fibrous lamella?, or was it, like Belemnites tubularis, hollow through a
greater or less part of its extent ? The specimens which have passed
through my hands do not enable me to give a definite reply to these
questions.

I suspect that a thoroughly well-preserved specimen of Belemno-
teuthis will some day demonstrate the existence of a fourth kind of
pro-ostracum among the Belemnitida?. Mr. Pearce, as we have already
seen, speaks of a " sepiostaire " in this genus ; .and Mr. Woodward
ascribes to it " a horny dorsal pen, with obscure lateral bands."
A specimen of Belemnoteuthis from the Oxford Clay, in the British
Museum (Plate II., fig. 2), shows very distinct traces of a pro-os-
tracum of this kind. The fossil is unfortunately much crushed, but
from one lip of the phragmocone there obviously proceeds the horny-
looking plate (a, a), the two lateral contours of which, obscurely defined
from the matrix, pass into one another at an acute angle at b. A
narrow band of horny-looking matter, marked by oblique stria?, is dis-
cernible at c, and is quite distinct from the remains of the mantle (/),
under which it seems to pass.

Is the triangular plate part of the ventral pro-ostracum, and the band
c the remains of the dorsal portion of that structure ? I am inclined
to think so, though the .state of the fossil is not such as to encourage
positive assertion.

It has been seen that at least two genera of Belcmnitida;, viz.,
Belemnitcs and Belemnoteuthis, have hooks, arranged in double rows,
upon their arms. Now similar hooks, sometimes isolated, some-

BRITISH FOSSILS. 19

times associated with more or less complete remains of the animal to
which they belonged, have been discovered in abundance in the Solen-
hofen slates, and have been referred by Wagner and Miinster to the
genus Acanthoteuthis. The interesting question therefore arises, was
there, in the Mesozoic epoch, a cephalopod (Acanthoteuthis) with
hooked arms, distinct from Belemnoteiithis and Belemnites ; or are the
Solenhofeu fossils in question to be referred to one or other of these
genera ?

Count Minister's two memoirs on Acanthoteuthis are to be found in
his -Beitrage zur Petrefacten Kunde" (Erstes und Siebentes Heft.
Zweite Auflage).

In the first memoir, the genus is founded upon specimens of four
species, consisting either of hooks alone, or of remains of the body and
arms, the latter retaining their hooks.

In specimens of Acanthoteuthis speciosa, the first species, the mantle
is said to be preserved, and in it " the broad sword-like pen, devoid
" of any distinct ridges, is visible " (1. c.. p. 105); and, in both of these,
" hooks are to be seen near the upper or cephalic end, perfectly similar
" to those figured in Plate IX." The hooks, situated in double rows
along the arms, are marked by two ridges, one of which runs near the
convex, and the other near the concave, side.

The second species is Acanthoteuthis Ferussacii (A. prisca of
D'Orbigny), the only specimen of which exhibits an elongated mantle
with a largish head, and short arms, provided with a double row of
hooks. Each hook has only one ridge, situated towards the concave
side.

The third species, A. Lichtensteinit, has short round hooks without
ridges, and the fourth, unnamed, has hooks with two fine grooves on
each side. But Count Miinster communicated an important observa-
tion, bearing upon the present question, in a letter to Professor Bronn,
published in Leonhard and Bronn's Jahrbuch for 1836 (p. 583) : —

" From Solenhofen I have the large Phragmacone (Alveolkegel) of
a Belemnite. with the unchambered hollow continuation of the shell,
beside which lies the injured body (Sack) of a very large Onychoteu-
this ; round about are seen a few minute hooks from the arms of the
Cephalopod. The two fossils lie so close together, and partly in super-
position, that one might, at first, be led to believe them to belong to
one and the same animal ; but more careful examination shows that they
proceed from two different animals, Belemnites semisulcatus and Onij-
choteuthis speciosa (the largest fossil kind with which I am acquainted).
Notwithstanding all the trouble which I have taken to find a Belemno-
sepia of Buckland in the slates of the Lias and of Solenhofen, I have
as yet met with no success ; in no German collection with which I am
acquainted is there any true Bclemnosepia, for which, at first, I took
the fossil just described."

Professor A. Wagner (" Die fossilen Ueberreste von nackten Dinten-
fiachen," I860), however, having had the opportunity of carefully ex-
amining all Minister's specimens, and of collating them with others,
leads us to believe that the earlier opinion was more correct than the
later. He says (p. 820), that he was at first of the same opinion as
Count Munster. but that he is now perfectly persuaded, " that on the
" slab in question there are not two examples of different genera, but
" only a single individual specimen. The phragmocone is, in fact,
" directly connected with the posterior margin of the mantle, and
<; exhibits the same structure as that of Acanthoteuthis (Belcmnoteu-
9909. c

20 BRITISH FOSSILS.

" this) antiqua, although only a coarse impression of it is left. The
" whole length of this individual, from the base of the arms to the
" apex of the phragmocone, is above 14 inches."

At a previous page (777) Wagner states that a few hooks lie beside
the head of this specimen, and that the form of its body is exactly like
that of A, Ferussacii. " But what giv.es this specimen its greatest
" value is the circumstance that, in the posterior part of the mantle-sac,
" at its posterior as well as at its two lateral edges, a few delicate
" fragments of a brown, horny, irregularly fissured pen (Schulpe) are
" visible."

In the second essay ("Ueber die SchalenlosenCephalopoden des oberen
Juragebirgs), Miinster says he is not certain what kind of mantle or
pen might have belonged to Acanthoteuthis speciosa, nor has he any
knowledge of the pen of A. Ferussacii, or of A. Lichtensteinii ; but
he proceeds to describe some new species, prefacing his account of them
with some general remarks, as follows : —

" The bodies of all the species known to me have a narrow elongated
form, which sometimes is elliptical, sometimes ovate, sometimes fusiform,
or even conical. Since, in a few specimens, impressions of booklets
are discoverable at the upper part of the body, which agree perfectly
with the three foregoing, and besides, coprolites not uncommonly occur
in the slates, which consist exclusively of the remains and undigested
parts of these naked cephalopods, namely, of the middle keel of the pen,
which is crushed into many short pieces, and of the booklets of the
arms, which, sometimes large and sometimes small, lie scattered round
the fragments of the pen in great numbers ; I have not hesitated to
ascribe all these bodies and pens to the genus Acanthoteuthis, until
this view is upset by complete specimens," p. 57.

Thus * Acanthoteuthis ' speciosa turns out to be one of the Belem-
nitidce, but the statements before us leave it doubtful whether it was
like Belemnoteuthis, devoid of an elongated guard, or whether it is
really a Bclemnites scmisulcatus with the guard broken off.

With respect to " Acanthoteuthis" Ferussacii, of which only one speci-
men exists, Wagner is uncertain as to its distinction from the former
species, and believes it to be identical with A. Lichtensteinii ; and at any
rate, as the head and trunk have left only an impression, and not a
trace of any internal parts is to be seen (Wagner, I.e., p. 775), there is
no evidence to show that it, also, may not be a Belemnites, or a Belem-
noteuthis.

Of the other Acanthotetithes enumerated in the second memoir,
Count Miinster does not profess to have found hooks associated with
A, angusta, A. lata, A. subovata, A. subconica, A. acuta, A. brevis, A.
intermedia, A. rhomboidalis, A. semistriata, and A. tricarinata, all of
which are referred to a different genus, Plesioteuthis, by Wagner ; while
Wagner, after examination of the same specimens, denies the existence
of hooks in A. Orbignyana and others, to which Munster ascribed
them.

Thus, the existence of Acanthoteuthis as a genus apart from Belem-
nites, or Belemnoteuthis, becomes exceedingly doubtful. But it does not
follow from this that no other Mesozoic Cephalopoda were provided with
hooked arms, and indeed there is evidence that at least two genera,
Plesioteuthis (Wagner) and Celceno (Munster) were. In the first place
Count Miiuster afiirms, and Professor Wagner agrees with him, that
coprolites are not unfrequently found in the Solenhofen slates, "which
" consist exclusively of the remains of undigested parts of naked cephalo-
" pods, namely, of the middle keel of the pen, which is crushed into

BBITISH FOSSILS. 21

u many short pieces, and of the booklets of the arms, which, sometimes
" large and sometimes small, lie scattered round the fragments of the
" pen, in great numbers." Wagner adds to this (l.c., p. 785) that the
fragments of the pen are part of the keel and of the lateral wings of
pens, appertaining, almost wholly, to animals in which the latter are
sword-shaped and thin, and for which Wagner proposes the generic
name Plesioteuthis. It would therefore appear that Plesioteuthis
had hooks, though Wagner's statement that he had never, either in
Minister's collection, or any other, found hooks associated with these
sword-shaped pens,* is, so far as negative evidence goes, somewhat
against that conclusion.

In the next place, Professor Wagner (l.c., p. 783) describes an im-
pression of Celceno conica displaying hooks similar to those of " Acan-
thoteuthis Ferussacii" and, in addition, the remains of acetabula.f

Upon the whole it becomes plain that the Acanthoteuthes of Munster,
so far as they are known only by hooks and impressions of soft parts,
may have been either Belemnites, or Belemnoteuthes, or Plesioteuthes,
or may have belonged to the genus Celceno ; and that, with the evi-
dence before us, it is impossible to say whether Acanthoteuthis speciosa
and Ferussacii belong to Belemnites, or to Belemnoteuthis.

Under these circumstances, it appears to me that there is no good
ground for abandoning the name Belemnoteuthis, applied by Pearce to
one of the best known and most clearly definable of fossil Cephalopoda,
for Acanthoteuthis. Though it is quite possible that either A. spe-
ciosa or A. Ferussacii may be really a Belemnoteuthis, we have no
certain knowledge of the fact ; and even if such be the case, it would be
better to separate these forms as Belemnoteuthis, and to retain Acan-
thoteuthis for the Plesioteuthis of Wagner.

The genera hitherto enumerated in the family of the Belemnitidce,
characterized among the Dibranchiate Cephalopoda by possessing a
straight, chambered, siphunculated, internal shell, or phragmocone, are
Belemnites, Belemnitella,\ Belemnoteuthis, Beloptera, and Conoteuthis.
To these Xiphoteuthis must now be added, and I think it very probable
that by-aud-by it will be found necessary to subdivide Belemnites, the
difference between the pro-ostraca of B. Bruguierianus and B. Puzo-
sianus being, probably, of generic importance.

The extent of our knowledge of the structure of these different
genera is very unequal. Of Belemnoteuthis, the body and arms, hooks,
ink-bag, and internal shell are all known, few fossilized animals having
left more complete remains ; of Belemnites, the specimens described in
this paper have made known, for the first time, the form and proportions
of the body and the arms, the hooks, the ink-bag, one type of pro-

* Out of coprolites, that is to say.

t Wagner speaks of these as "hitherto never observed in fossil Cephalopoda"
(p. 783). but he has overlooked a paper " On the fossil Cephalopoda constituting the
" genus Belemnoteuthis," by Mr. J. C. Pearce, F.G.S., published in the " London
" Geological Journal," Xo. II., February 1847, in which the acetabula of Belemnoteu-
this are described and figured. (PI. XVI.)

J See, however, -with respect to BelemniteUa and Actinocamcu-. the important
observations of Saemann, " Observations sur Belemnites quadratus, Defr." — Bull, de la
Sociuto Gcologique de France, 1862. M. Saemann brings forward evidence to show
that these apparently distinct generic types arise merely from the defective calcifica-
tion of the upper part of the rostrum of a Belemnite.

355817

22 BRITISH FOSSILS.

ostracum ; and, less perfectly, the beak ; of XiphoteutJiis, the almost
complete internal shell is known ; of Conoteuthis, the phragmocone and
part of the pro-ostracum ; of Bcloptera and Belemnitella, only the
phragmocone and guard; but with the hooks, ink-bag, or soft parts of
these last four genera we have no acquaintance.

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COUNTRY around NOTTINGHAM. By W. T. AVELINE, F.G.S. Illustrating 71 N.E. Price 8d.
The GEOLOGY of PARTS of NOTTINGHAMSHIRE, YORKSHIRE, and DERBYSHIRE. Illustrating Sheet 82

By W. TALBOT AVELINE, F.G.S. Price 8d.
The GEOLOGY of SOUTH BERKSHIRE and NORTH HAMPSHIRE. Illustrating Sheet 12. By H. W. Jlar

and W. WHITAKEE. Price 3s.
The GEOLOGY of the ISLE OF WIGHT, from the WEALDEN FORMATION to the HEMPSTEAD BEDS incli

with numerous Illustrations, and a List of the Fossils fouud in the Island. Illustrating Sheet 10. By H. W. BRIS

F.R.S. Price 6s.

The GEOLOGY of EDINBURGH. Illustrating Sheet 32 (Scotland). Price 4s. By H. H. HOWELL and A . GEIKIE.
The GEOLOGY of the COUNTRY around BOLTON, LANCASHIRE. ByE. HULL. B.A. Illustrating Sheet 89S.E. 1'ri
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Tin- GEOLOGY of Iho COUNTRY around OLDHAM. By E. HULL, B.A. Illustrating 88 S.W. Priro 2s.
> I .< >< : Y of PARTS of MIDDLESEX. &o. Illustrating Shoot 7. By W. WuiTAKER, B.A. Priro »„-.

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AA 001172047 i

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